Homarid Lobster Hatcheries: Their History and Role in Research, Management, and Aquaculture

Home , Homarus gammarus, Maine Avenue Fish Market

FRANK NICOSIA and KARI LAVALLI

Introduction from the scientifi c research they con- investigated within these hatchery set- ducted. Bardach et al. (1972) empha- tings; questions concerning the state of Most historical reviews on lobster, sized the activities conducted at the the fi shery and its preservation were also Homarus spp., hatcheries have dealt hatchery on Martha’s Vineyard, Mass. addressed (Herrick, 1894, 1895, 1911a; with the specifi c objectives of hatchery In each case, the reviews focused on the Scattergood, 1949b). The information operation: hatching eggs and release economic and biological successes or gained thus far (reviewed in Cobb, 1976; of Stage I larvae or the rearing and failures of hatchery efforts (see Mead, Cobb and Phillips, 1980; McVey, 1983; release of Stage IV postlarvae (Nigrelli, 1910; Scattergood, 1949b; Taylor, 1950; Cobb and Wang, 1985; D’Abramo and 1936; Carlson, 1954; Thomas, 1964; Carlson, 1955; Taylor and Dow, 1958; Conklin, 1985; Mackenzie and Moring, Wilder, 1972; Dexter, 1986). Kenslor Prudden, 1962; Dow, 1969). However, 1985; Waddy, 1988; Aiken and Waddy, (1970) reviewed lobster hatcheries and, the current usefulness of lobster hatch- 1989, 1995; Lee and Wickins, 1992; to a limited extent, the benefi ts accruing eries in advancing knowledge about the Chang and Conklin, 1993; Conklin and life-history of homarid lobsters has not Chang, 1993; Factor, 1995; Waddy and Frank W. Nicosia is with Tri-Aqua Gold Lobster been fully explored. Aiken, 1995) has accumulated from over Farms, P.O. Box 2, Smithtown, NY 11787. Kari L. Lavalli is with the Department of Biology, From as early as 1858, experimental 100 years of detailed experiments. In Southwest Texas State University, 601 Univer- lobster culture has provided a large retrospect, the facts obtained from the sity Drive, San Marcos, TX 78666. Mention of trade names or commercial fi rms does not imply volume of information on the species’ early work of hatcheries formed a sound endorsement by the National Marine Fisheries life history (Scattergood, 1949a; Lewis, foundation from which current lobster Service, NOAA. Views or opinions expressed or 1970; Nowak, 1972). Anatomy, phys- research and management evolved. implied by the authors do not necessarily refl ect the position of the National Marine Fisheries Ser- iology, development, general habits, This paper reviews and summarizes vice, NOAA. behavior, and preferred habitats were the literature on past and present homarid lobster culture, hatchery activities, and stock enhancement programs, and ABSTRACT—This paper provides an his- descriptions of life stages, behavior, phys- gives recommendations for their future torical review of homarid lobster fi sheries, iology, etc.—has generally been confi rmed the development and usage of lobster hatch- rather than refuted and has stimulated fur- use. Most of the emphasis will be on eries, and much of the research infl uenced ther research important for an understand- the American lobster, Homarus ameri- by hatchery-initiated studies on natural his- ing of the life history of homarid lobsters. canus (Fig. 1) with references made to tory, physiology, and morphological devel- The connections between homarid fi sheries comparable data for H. gammarus (pre- opment of the lobster, Homarus spp. Few and hatchery operations (i.e. culturing of the viously H. vulgaris). The three species commercial lobster hatcheries exist in the lobsters), whether small- or large-scale for world today, yet their potential usage in fi eld and laboratory research, are important of clawed lobsters, H. americanus, H. restocking efforts in various countries is con- to understand so that better tools for fi shery gammarus, and Nephrops norvegicus, stantly being reexamined, particularly when management can be developed. This review have similar morphological and devel- natural stocks are considered “overfi shed.” tries to provide such connections. However, opmental trends (Gruffydd et al., 1975; Furthermore, many individual researchers the rearing techniques in use in today’s hatch- working on homarid lobsters use small- eries—most of which are relics from the past— Howard, 1989). Each species has a scale hatchery operations to provide the are clearly not effi cient enough for large-scale prelarval stage followed by three larval animals necessary for their work as well commercial aquaculture of lobsters or even stages and then a postlarval stage which as animals reared and provided by various for current restocking efforts practiced by sev- resembles the adult. These developmen- governmental agencies interested in specifi c eral countries today. If hatcheries are to be tal similarities imply behavioral and projects on larvae, postlarvae, or juveniles. used to supplement homarid stocks, to restock Such researchers can benefi t from the infor- areas that were overfi shed, or to reintroduce ecological similarities (Berrill, 1974). mation in this review and can avoid many species into their historical ranges, there is Thus, culture techniques and informa- pitfalls previously documented. a clear need to further develop culture tech- tion obtained for one species often can be The development of hatcheries and the niques. This review should help in assessments directly applicable to the others (Van Olst experimental studies that were generated from of culturing techniques for Homarus spp. and their activities have had a direct impact provide a reference source for researchers et al., 1980; Cobb and Wang, 1985). on much of the research on lobsters. The or governmental agencies wishing to avoid Although this is not an exhaustive past work arising from hatchery operations— repeating previous mistakes. review of all hatchery-inspired experi-

61(2), 1999 1 cotton or manila cord heads, tarred and strung to form a “funnel,” which was attached to an entrance ring made of spruce 15.2 cm (6 inches) in diameter (Herrick, 1911a). As bait, fi shermen used salted or fresh herring, halibut, hake, and cod heads (Cobb, 1901; Herrick, 1911a; Dow, 1980), or, to a lesser degree, synthetic substances, which consisted of a cloth bag fi lled with sand and saturated with uncooked herring oil, or mackerel pellets saturated with redfi sh oil (Prudden, 1962; Dow, 1980). Baited pots were weighted and then placed on the sea bottom, either singly, doubly, or by trawl (8–40 pots) with a rope or cord attached to a wooden fl oat (buoy). Hundreds of traps were pulled (hauled to the boat) by hand several times a day, while others were Figure 1.—American lobster, Homarus americanus. left overnight (Herrick, 1911a; Dow, 1949). When lobsters seemed to be less abundant or more widely scattered in ments and their results, this paper will were gaffed with a hook attached to a the 1880’s, fi shermen returned to the provide the reader with an insight into pole nearshore (Rathbun, 1884a, 1887; less common and older practice of set- how valuable the hatcheries have been Cobb, 1901; Herrick, 1911a; Dow, ting pots singly and altering pot posi- and what their future role could be. 1949; Krouse, 1989; White, 1991). Her- tion daily in hopes of capturing more For the enthusiast wishing to pursue the rick (1895) attributed this abundance of lobsters by covering more fi shing areas subject further, a bibliography of well nearshore lobsters to a bountiful food (Rathbun, 1884a) (Fig. 2). over 400 literature citations and selected supply, but explained that the number An ever increasing demand for the references is also given. To adequately and persistence of lobstermen had pro- lobster as a source of food resulted in review the origins of lobster hatcheries, found effects upon the abundance of tremendous increases in fi shing inten- we must start with a brief overview of larger lobsters. sity and annual landings. The change the history of lobster fi sheries. Lobsters were also taken by hoop from hoop nets to pots allowed so many nets used from small boats near the lobsters to be captured that supply some- History of Lobster Fisheries shoreline (Cobb, 1901; Herrick, 1911a; times exceeded demand, and, in the The American lobster and its Euro- Dow, 1949; Krouse, 1989). Hoop nets United States, lobsters were so abundant pean counterpart, H. gammarus, are were labor intensive due to their con- that they were used as agricultural fertil- among nature’s most valuable resources struction—they consisted of a 70 cm izer, cod bait, and for semicommercial for commercial and, to a limited extent, diameter iron hoop over which two half purposes in the 18th and 19th centuries recreational fi shermen. Until this cen- wooden hoops crossed. The iron hoop (Dow, 1980). By the early 19th century, these lobsters have been able to was attached to a shallow net bag and tury, what seemed to be an inexhaust- survive the hazards of nature and man- it was baited at the intersection of the ible supply of lobsters had begun to kind, despite commercial fi shing efforts wooden hoops. Because lobsters could decline (Wood, 1869; Rathbun, 1884b). which began as early as the 17th century exit as easily as they could enter the In an effort to ensure a continuous (Herrick, 1911a; Dow, 1949; DeWolf, nets, fi shermen had to pull the nets supply of lobsters, protective measures 1974; Bennett, 1980; Dow, 1980). They every 10–30 minutes (Rathbun, 1887; were passed in state legislatures which were reported as easily captured food Miller, 1995). included licensing fi shermen, leaving or sources in both Canada and New Eng- Around 1840, fi shermen modifi ed returning to the sea “berried” females, land in the early 1600’s and were so those hand practices to achieve more closing certain fi shing areas during par- plentiful that they were also used as fi sh effi ciency and began utilizing a trap ticular seasons, limiting the size of lob- bait and fertilizer in the 1800’s (DeWolf, or “pot” to capture lobsters (Herrick, sters caught, and culturing. 1974; Martin and Lipfert, 1985). 1911a; Dow, 1949; Krouse, 1989). Rang- Today, the pot shape basically remains During the summer months, lobsters ing in sizes from 0.76 to 1.2 m (2.49 unchanged, being rectangular (most were so common in the shallow littoral to 3.94 feet) long, 61 cm (24 inches) common), half-round, or squared (Firth, zone that fi shermen often gathered them wide, 45.7 cm (18 inches) high, these 1950; Everett, 1972; Dow, 1980; Krouse, by hand, dip net, and spear, or they pots were made of wooden laths and 1989; Miller, 1995); however, instead

2 Marine Fisheries Review Figure 2.—Above: Dory ﬁ shermen hauling lobster pots off Cape Ann, Mass. Below: Lobster ﬁ shing boats of Bristol, Maine. (Goode, 1887).

61(2), 1999 3 of being made of wood, the frames may now be made of aluminum, plastic, or vinyl-coated wire frames (Krouse, 1989) (Fig. 3). Nylon or plastic replaced the cotton and manila cord heads, and polypropylene is used instead of manila for the warps (Prudden, 1962; Krouse, 1989). Wooden or cork buoys were replaced by styrofoam or plastic buoys (Scarratt, 1980). Perhaps the most signifi cant change was the addition of an inner chamber, or “parlor” (Fig. 3b, c), which increased the effi ciency of the pot, as lobsters could not fall out of the end funnel during hauling and were less likely to escape on their own (Knight, 1918; Krouse, 1989; Miller, 1995). To reduce ghost fi shing by lost pots and to allow many undersized lobsters to escape before hauling, escape vents were added into the design of the trap in the 1980’s and trap doors or “ghost” panels are now closed with a material that bio- degrades in 1 year (Miller, 1995). In addition, the row boats and sail- ing dories previously used by fi shermen (Fig. 3) have been replaced by faster and larger boats, averaging at least 11 m (36 feet) in length (Gates and D’Eugenio, 1975; Pringle and Burke, 1993), equipped with electronic devices (e.g. radio, cellular phones, depth sound- ers, LORAN, GPS, radar) and sleeping berths. Small gasoline engines, used as early as 1900, were replaced by diesel engines in the 1970’s (Miller, 1995). Belt-driven haulers were also replaced by electric or hydraulic pot haulers (Krouse, 1989; Miller, 1995). While many pots are still baited with 2–4 fresh or salted fi sh, some lobstermen are once again using synthetic baits made from animal hide which are now proving successful (Stevens, 1993). These synthetic baits are cut into 8-ounce (227 g) strips, soaked in a salt solution to stabilize and preserve them, and infused with fi sh oils. One pound of this bait will fi sh a pot for 1 month (Stevens, 1993). Because of these modern materials and methods, fi shermen can now fi sh further from shore, fi sh more pots at a faster rate, and spend less time repairing pots. Early Protective Measures Figure 3.—Types of lobster pots. A) six-sided trap with three entrances and no parlors; The earliest lobster protective mea- B) lath trap with two entrances and one parlor; C) wire trap with two entrances and two sure, enacted by Massachusetts in 1812, parlors. Redrawn from Miller (1995) by Sapphire Tur-Caspar.

4 Marine Fisheries Review Table 1.—Changes in minimum legal sizes, state-by-state, for the American lobster, Homarus americanus.1

Maine New Hampshire Massachusetts

Total Carapace Maximum Total Carapace Total Carapace length length length length length length length Years (inches) (inches) (inches) Years (inches) (inches) Years (inches) (inches)

1874–1883 (10 1/2 (3 5/8)2 1874–1907 (10 1/2 (3 5/8)2 1874–1907 (10 1/2 (3 5/8)2 1883–1885 ( 9 (3 3/32) 1907–1933 ( 9 (3 3/32) 1907–1933 ( 9 (3 3/32) 1885–1889 (10 1/2 (3 5/8) 1933–1941 (9 7/32) (3 3/16 1933–1941 (9 7/32) (3 3/16 1889–1895 ( 93 to 10 1/2 (3 3/32 to 35/8) 1941–1988 (9 1/16) (3 1/8 1941–1950 (9 1/16) (3 1/8 1895–1907 (10 1/2 (3 5/8) 1988–1989 (9 11/32) (3 7/32 1950–1988 (9 1/16) (3 3/16 1907–1919 (10 1/2 (3 5/8) 1989–1993 (9 7/16) (3 1/4 1952–1989 (9 11/32) (3 7/32 1919–1933 (10 1/8) (3 1/2 1989–1993 (9 7/16) (3 1/4 1933–1935 (8 7/8) (3 1/16 4 3/4 1935–1942 (8 7/8) (3 1/16 5 1942–1958 (9 1/16) (3 1/8 5 1958–1960 (9 7/32) (3 3/16 5 3/16 1960–1988 (9 7/32) (3 3/16 5 1988–1989 (9 11/32) (3 7/32 1989–1993 (9 7/16) (3 1/4 5

Rhode Island Connecticut New York

Total Carapace Total Carapace Total Carapace length length length length length length Years (inches) (inches) Years (inches) (inches) Years (inches) (inches)

1874–1880 (10 (3 5/8) 1874–1875 (10 (3 15/32) 1874–1893 (10 1/2 (3 5/8) 1880–1895 (10 1/2 (3 5/8) 1875–1878 ( 8 (2 3/4) 1893–1935 ( 9 (3 3/32) 1895–1909 ( 9 (3 3/32) 1878–1895 ( 6 (2 1/16) 1935–1941 (9 7/32) (3 3/16 1909–1936 ( 4 1/8 (TBSL)4 (3 1/8) 1895–1909 ( 9 (3 3/32) 1941–1952 (9 1/16) (3 1/8 1936–1966 (9 7/32) (3 3/16 1909–1925 ( 4 1/8 (TBSL)4 (3 1/8) 1952–1989 (9 7/32) (3 3/16 1966–1989 (9 11/32) (3 7/32 1925–1955 (9 7/32) (3 3/16 1989–1990 (9 11/32) (3 7/32 1989–1993 (9 7/16) (3 1/4 1959–1988 (8 7/8) (3 1/16 1990–1993 (9 7/16) (3 1/4 1988–1989 (9 11/32) (3 7/32 1989–1993 (9 7/16) (3 1/4

1 Sources: 1 Kelly (1990). 1 K. H. Kelly, Maine Department of Marine Resources, West Boothbay Harbor, Maine 04575. Personal commun. 16 Sept. 1992. 1 Steven X. Cadrin, Massachusetts State Division of Marine Fisheries, Sandwich, Mass. Personal commun. 16 Dec. 1991. 1 J. I. Nelson, State of New Hampshire Fish & Game Department, Durham, N.H., State of N.H. Fish & Wildlife Documents and personal commun. 5 Feb. 1992. 1 Thomas E. Angell, Rhode Island Division of Fish & Wildlife, Coastal Fisheries Laboratory, 1231 Succotash Road, RR#1, Wakefi eld, R.I. 02879. Personal commun. 14 May 1992. 1 P. T. Briggs, New York State Department of Environmental Conservation, Stony Brook, N.Y. Personal commun. 18 Dec. 1991. 1 Public Acts of Connecticut, 1874–1971; Public and Special Acts of the State of Connecticut, 1971–1993 2 Numbers in parentheses are approximate equivalent measures. 3 9" was established for lobsters to be canned. 4 TBSL = total back shell length from tip of rostrum to center rear of carapace. prohibited nonresidents from fi shing in bition on holding, buying, and selling were permitted for cannery lobsters local waters without permission (Rath- recently molted lobsters was replaced until 1891 in the United States (primar- bun, 1886). Maine followed suit in by a closed season (Miller, 1995). ily Maine) (Kelly, 1992; Miller, 1995). 1823, requiring nonresidents to obtain In 1873, the fi rst minimum size limits In contrast, cannery lobsters continue to a permit to fi sh for lobsters (Dow, 1949; (76–95 mm carapace length (CL)) were be smaller throughout Canada, where Kelly, 1990). Later, Maine enacted a established for landed lobsters in the canning industry has had consider- law in 1872 prohibiting the catching, Canada; however, these size limits ably more infl uence (Miller, 1995). buying, or selling of “berried” lobsters. varied from region to region and con- The average size of lobsters marketed This was repealed in 1874, and a closed tinue to do so today. Several U.S. states in 1880 was about 92.3 mm CL (10.5 season was established on all lobsters followed suit in 1874 with a size limit of inches TL) in Portland, Maine, and New from 1 August to 15 October of each 10.5 inches total length ((TL) exclusive Haven, Conn.; 96.8 to 101.2 mm CL year (Dow, 1949). From 1917, ovig- of claws and antennae) being enacted (11 to 11.5 inches TL) in Boston, Mass.; erous females could be sold only to in Maine, Massachusetts (Wheildon, and 92.3 to 132.5 mm CL (10.5 to 15 the state, which marked them with a 1874), New Hampshire, Rhode Island, inches TL) in New York City (Rathbun, hole punched into their uropod and then and New York (Table 1). Connecticut 1884a). In weight, these lobsters ranged released them back into local waters as enacted a smaller size limit of 10 inches from 648.6 g (1.43 lb) at 92.3 mm CL property of the state (Miller, 1995). In TL (Wheildon, 1874). These sizes of to 1,941 g (4.28 lb) at 132.5 mm CL 1948, the uropod punch was replaced by 10 and 10.5 inches TL are roughly (Table 2). the V-notch (Miller, 1995). In Canada, equivalent to 87.8 and 92.3 mm CL, Since annual U.S. landings increased the holding, buying, or selling of berried respectively, which is the preferred mea- signifi cantly from 7,223 metric tons (t) females or recently molted lobsters was surement today. (Table 2 gives con- in 1880 to 13,958 t in 1889 (Anderson prohibited in 1873. In 1874, the prohi- versions of TL to CL.) Smaller sizes and Peterson, 1953) (Fig. 4), the estab-

61(2), 1999 5 lishment of a minimum legal size was mm CL (10.5 inches TL) remained in 3,175 t or 23% between 1889 and 1892 credited with the arrest of the declining effect for at least 15 years, these larger (Smith, 1898) (Fig. 5), and the average supply of lobsters (Rathbun, 1884b). lobsters did not actually maintain the size of landed lobsters decreased. Con- However, even though a size limit of lobster stocks. Landings in the New sequently, around 1893–95 New York, 87.8 mm CL (10 inches TL) to 92.3 England states alone declined more than Connecticut, and Rhode Island reduced their size limits to 78.9 mm (9 inches

Table 2.—Conversion of total length measurements to carapace length measurements with corresponding egg TL), 65.5 mm CL (7.5 inches TL), and production approximations and weights. 87.8 mm CL (10 inches TL), respec- Conversion of TL to CL1 Egg production2 Weight3 tively (Table 1), because smaller lobsters were more abundant. Similarly, TL (inches) TL (mm) CL (inches) CL (mm) (Approx. eggs) Pounds Grams landings in the Canadian provinces grew 6 152.4 2 1/16 52.1 0.25 113.9 7 1/2 190.5 2 19/32 65.5 4,269 0.50 226.8 until 1886 and then began a decline, 8 203.4 2 3/4 69.9 5,448 0.61 276.7 with minor upturns in the late 1890’s, 8 7/8 225.4 3 1/16 77.8 7,802 0.85 385.6 9 228.6 3 3/32 78.9 8,179 0.89 403.7 1930’s, and 1950’s. In the 1980’s, land- 9 1/16 230.2 3 1/8 79.4 8,354 0.90 408.2 ings again began to increase (Miller, 9 7/32 235.0 3 3/16 81.0 8,932 0.96 435.5 9 11/32 237.0 3 7/32 81.8 9,231 0.99 449.1 1995) (Fig. 4). Because of these intense 9 7/16 239.2 3 1/4 82.6 9,537 1.02 462.7 fi shing pressures which removed the 9 1/2 241.2 3 9/32 83.3 9,811 1.04 471.7 9 19/32 243.4 3 5/16 84.1 10,131 1.07 485.4 stock of larger lobsters considerably 10 254.0 3 15/32 87.8 11,705 1.23 558.0 10 1/8 257.1 3 1/2 88.9 12,204 1.27 577.1 faster than reproduction and natural 10 1/2 266.7 3 5/8 92.3 13,841 1.43 648.6 growth could replenish it, questions 10 27/32 275.4 3 3/4 95.3 15,409 1.57 712.2 12 304.8 4 5/32 105.7 21,810 2.15 975.2 arose concerning the age at which sexual 14 13/32 365.4 5 127.0 40,372 3.76 1,705.5 maturity was reached and the egg pro- 15 381.0 5 7/32 132.5 46,541 4.28 1,941.4 duction of mature lobsters. 1 TL = 2.8424 CL + 4.3922, r2 = 0.9906, n = 431, from Steven X. Cadrin, Massachusetts State Division of Marine Fisheries, Herrick (1894), working at Woods Sandwich, Mass. Personal commun. 16 Dec. 1991. 2 Hole, Mass., dissected over 100 females Herrick’s (1894) calculation: Log10 (Fecundity) = -2.4505 + 3.3542 Log10 (CL), from Saila et al. (1969). 3.0374 and determined that most reached matu- (Length )∗ 0.000692 3 Weight = from Thomas B. Hoopes, Division of Marine Fisheries, Salem, Mass. Personal rity between 69.9 and 105.7 mm CL 452 commun. 18 Dec. 1991. (8–12 inches TL). He further determined that the majority would reach maturity by 87.8 mm CL (10 inches TL). He estimated that a lobster of 69.9 mm CL averaged about 5,448 eggs, one of 78.9 mm CL averaged 8,179 eggs, one of 87.8 mm CL averaged 11,705 eggs, and one of 105.7 mm CL averaged 21,810 eggs (Table 2). To date, Herrick’s maturity and fecundity fi ndings have been generally confi rmed by various researchers in Maine, Massachusetts, and New York (Krouse, 1973; Briggs and Mushacke, 1979; Estrella and McKieran, 1989; Estrella and Cadrin, 1990; Graulich1). Herrick (1894) also concluded that all states with size limits less than 92.3 mm CL needed to raise their legal limits, but it was not until 1907 that any state responded. Maine raised its size limit to 95.3 mm CL (10 27/32 inches TL), New Hampshire and Massachu- Figure 4.—U.S. and Canadian landings of lobsters (in metric tons) for the years setts lowered their size limits to 78.9 1880 to 1992. All data from 1976 to 1995 are from preliminary reports of state agen- mm CL (9 inches TL), and in Rhode cies. Data for 1995 for the U.S. and for 1996 from Canada are preliminary. (U.S. Island, Connecticut, and New York size landings from Historical Fishery Statistics Summary of American Lobster Land- ings, Northeast Fisheries Center, MA, transmitted from R.L. Shultz; Canadian land- 1 Graulich, H. A. 1991. American lobster investi- ings from Douglas Pezzack, Lobster Biology & Assessment, Dept of Fisheries and gations in New York waters. U.S. Dep. Commer., Oceans, P.O. Box 550, Halifax, Nova Scotia B3J 2S7, Canada. Personal commun. NOAA, NMFS Compl. Rep., Proj. 3- IJ-11 under 21 April 1997.). Interjuris. Fish Act, 21 p.

6 Marine Fisheries Review limits remained the same at 78.9 mm CL (Table 1). Herrick (1898) further concluded that laws should be established to prohibit the taking of egg-bearing females. How- ever, despite all states enacting such a law by the turn of the century (Carl- son, 1955), annual landings continued to decline. U.S. landings declined from 13,958 t in 1889 to 5,227 t in 1905 and to about 4,407 t in 1924 (Ander- son and Peterson, 1953) (Fig. 4), while in Canada a much larger decline was evident, decreasing from 47,620 t in 1886 to 31,746 t in 1906 and then dramatically to 12,200 t in 1924 (Pringle et al., 1993). The European H. gammarus fi sheries also experienced similar declines (Herrick, 1911a; Bennett, 1980). Table 1 provides the changes in size limits throughout successive years for the various U.S. states. Combined landings from all states (Fig. 4) were used to interpret the overall condition of the U.S. lobster fi shery. However, because Maine landings represented about 50% or more of the catch landed each year (Fig. 5), the use of combined data could result in mis- interpretations about declines and/or increases in landings. For example, Figure 4 illustrates a continuous decline in the overall U.S. landings from 1889 to about 1935; however, Figure 5 depicts each state separately and shows that only Maine landings declined, while landings in the other states remained stable or increased. During this period of reduced landings in Maine, the legislature passed what came to be known as a “poverty gauge” of 4 3/4-inch back- shell length (slightly longer than CL) in 1907. In response fi shermen were thought to routinely land illegal lobsters for home consumption or illicit shipping Figure 5.—State-by-state landings of lobsters (in metric tons) for the years 1880 to 1992. A) Landings from Maine, Massachusetts, and Rhode Island; B) Landings and to smash undersized lobsters for from New Hampshire, Connecticut, and New York. Some data points are missing bait (Acheson, 1997). The legislature until 1942 and data from 1995 are preliminary. then changed the size limit to 3.5-inch CL in 1919, but this did not appreciably raise the landings nor decrease the violations of the law. Finally, after CL (4 3/4-inch CL) in 1933 (Acheson, explain the so-called “bust” in the early a closure of the fi shery along the cen- 1997). Beginning in 1935 overall land- 20th century Maine fi shery. tral Maine coast in the 1920’s and a ings increased both in terms of total further reduction of the minimum size pounds landed and in terms of pounds Environmental Factors limit to 77.8 mm CL (approx. 3 1/16- landed per pot fi shed (Fig. 5; Table and Landing Fluctuations inch CL), Maine put in place a max- 3). Acheson and Steneck (1997) fur- Insuffi cient information on the causes imum carapace measure of 121 mm ther discuss a series of hypotheses to of natural fl uctuations in landings brought

61(2), 1999 7 Table 3.—Historical landing data for the State of Maine, including number of pounds landed, number of pots ﬁ shed, minimum sizes in effect, approximate weights of lobsters landed, and calculated number of lobsters landed per not begin to mature until they attain a pot. size of 80 mm CL or larger (Krouse,

Item 1897 1942 1992 1973; Aiken, 1980; Aiken and Waddy, 1980; Waddy and Aiken, 1991). Tem- Number of pounds landed 10,300,0001 8,400,0001 26,830,0002 Number of pots fi shed 234,0001 187,0001 2,000,0002 peratures also affect the proportion Number of pounds per pot fi shed 44 45 13.4 of prerecruits entering (molting) into Number of lobsters per pot 31 53 13 Minimum size (mm CL) 92.3 77.8 82.5 the fi shery (Campbell, 1983; Estrella Approximate weight (in lbs.) of lobster of minimum size 1.431 0.851 1.021 Average weight (in lbs.) of lobster landed 2.633 1 1.242 and Cadrin, 1991). Furthermore, activ- Corrected number of lobsters per pot based on average weight landed 16.7 45 10.8 ity and catchability are associated with

1 Source: Dow et al. (1975). increased water temperatures (McCleese 2 Source: Jay Krouse, Department of Marine Resources, Marine Resources Laboratory, McKown Point, West Boothbay and Wilder, 1958; Dow, 1966; Flowers Harbor, Maine 04575. Personal commun. 22 March 1994. and Saila, 1972), which directly affect 3 Estimate based on the average lobster marketed in 1880, from Rathbun (1884b). landings (Taylor et al., 1957; Dow, 1961, 1977, 1978, 1980; Fogarty, 1988; new research efforts, directed at the sig- inated between 1874–1892, landings Estrella and Cadrin, 1991; Kelly, 1992, nifi cant role that environmental factors declined (Smith, 1898; Fig. 5), sug- 1993). played in regulating and controlling the gesting that increased egg production Despite the upward trend in landings survival, size at maturation, catchability, was indeed irrelevant. However, Fog- beginning in the late 1930’s, researchers and, ultimately, the supply of lobsters. arty (1995) suggests that small changes predicted that climatological conditions These factors include, but are not limited (such as an increase of 1%) in larval for the remainder of the century would to, food, light, salinity, disease, mutila- survival could dramatically increase the not improve or maintain the extant land- tion, social environment, and water qual- number of lobsters eventually recruit- ings (Dow, 1980). Furthermore, extant ity (Herrick, 1911a; Templeman, 1933, ing to the fi shery. Nonetheless, recent levels of fi shing were assumed to be 1936; Aiken, 1980; Aiken and Waddy, attempts to explain the low landings substantially greater than those which 1986; Ennis, 1986a). in Maine in the 1930’s and the higher would allow the greatest productivity Although each of these factors is landings of the 1990’s, showed little from the resource (Northeast Marine important, many have postulated that relationship between landings and tem- Fisheries Board, 1978). This informa- temperature plays a key role in increas- perature, particularly during the recent, tion caused concern for the long-term ing survival of the larvae and post- so-called, “boom” years (Acheson and viability of the overall fi shery with larvae, accelerating growth rates, and Steneck, 1997). Thus, while temperature respect to stock and recruitment. Efforts increasing both activity and catchabil- may improve larval survival, its posi- were directed toward unifi ed manage- ity. Temperature affects larval lobsters tive effects on subsequent survival and ment and eventually resulted in the throughout their course of development recruitment to the fi shery are uncoupled establishment of the American Lobster (Hadley, 1906a; Templeman, 1936), at some point. This uncoupling has led to Fishery Management Plan (FMP) in short ening their developmental rates the idea that there may be bottlenecks 1983. The FMP was designed to 1) pro- when elevated, and increasing their present in the life cycle of the lobster mote conservation, 2) reduce the pos- chances of survival (Templeman, 1936; that limit lobster numbers (Wahle and sibility of recruitment failure, and 3) Hughes and Mattheissen, 1962, 1967; Steneck, 1991). allow full utilization of the resource Caddy, 1979; Harding et al., 1983; Ennis, Elevated summer water temperatures by the U.S. fi shing industry. The main 1986a; Mackenzie, 1988; Corey2). There- not only accelerate growth rates, they objective, however, was to support and fore, the number of larvae surviving to can also induce early maturation (Aiken, promote the development and imple- settle is more dependent upon favorable 1980; Waddy and Aiken, 1991). Typi- mentation, on a continuing basis, of environmental conditions than on the cally, male lobsters mature at the rela- a unifi ed, regional management pro- number or size of spawning adults (Carl- tively smaller sizes of 50–70 mm CL in gram for American lobsters (Anony- son, 1955). Herrick (1895) remarked most areas, regardless of water temper- mous, 1983a). that the destruction of a few hundred ature (Krouse, 1973; Briggs and Mus- Recommendations included a uni- thousand eggs, or even millions, would chacke, 1979). In contrast, there is a form size limit of 81 mm CL (3 3/16- not appreciably affect the supply of lob- wide variation in maturation among inch CL) to be established in all U.S. sters at any given point along the coast. females in various geographical loca- areas by 1981 (Northeast Marine Fish- If Herrick was correct, then any increase tions. For example, higher water tem- eries Board, 1978). This size limit was in egg production within the same order peratures in Long Island Sound, N.Y., fi nally implemented in 1985 (Anony- of magnitude would not appreciably Buzzards Bay, Mass., and the Gulf of mous, 1983a, b). The objective behind improve recruitment into the fi shery. St. Lawrence, Can., cause females to the increase in minimum size stemmed In fact, when larger lobsters predom- mature at the smaller size of 60 mm CL, from studies of instantaneous mortal- with most maturing at 80 mm CL. In ity rates (fi shing and natural), together 2 Corey, S. 1963. Research on lobster larvae (Homarus americanus). Biol. Sta., St. Andrews, the colder waters of the Gulf of Maine with fecundity studies, general growth N.B., Can. Unpubl. manuscr., 16 p. or southern Nova Scotia, females may rates (from various areas), and yield

8 Marine Fisheries Review per recruit (Herrick, 1895; Saila et al., in any size increase beyond the 82.5 The management teams met their dead- 1969; Skud and Perkins, 1969; Krouse, mm (3 1/4-inch) CL size. These groups line; however, the Council failed to meet 1973; Briggs, 1975; Uzmann et al., claimed that these size increases put its deadline due to the objections of the 1977). However, the most important them at an economic disadvantage with states of Maine, New Hampshire, Rhode result of those studies was to determine Canadian lobster suppliers who were Island, and Connecticut to certain mea- the size at which females reached matu- allowed to take animals 81 mm CL (3 sures in the recommended amendment. rity (Northeast Marine Fisheries Board, 3/16-inch CL) or even smaller in cer- Because the NMFS requested that all 1978). tain regions (Miller, 1995). states commit to participation in the Amendment 4 to the FMP was amendment’s administration and four Fishery Management: implemented in 1991 and temporarily states refused, NMFS proposed the The Federal Government rescinded the next scheduled size withdrawal of the FMP in March 1996 and the States increases, such that all major U.S. lob- stating that the plan would no longer The National Marine Fisheries Ser- ster-producing states stood at the 82.5 meet the standards set forth in the Mag- vice (NMFS) and the U.S. lobster pro- mm CL size limit (Anonymous, 1991d). nuson-Stevens Act requiring the pre- ducing states came together in 1978 In 1994 the NEFMC adopted Amend- vention of overfi shing (Lockhart and to formulate a State-Federal Fishery ment 5 to the FMP to address the over- Estrella, 1997; Anonymous, 1998). Management Plan (FMP) for the lob- fi shed condition of the lobster resource Nonetheless the NMFS did not with- ster that would provide for a single, while maintaining the minimum size draw the FMP until an effective state unifi ed approach to managing the fi sh- of fi shable lobsters at 82.5 mm CL. management plan was completed to ery. The goals of this plan were to This Amendment was required to avoid replace it. The Atlantic States Marine adjust minimum size limits appropri- the next scheduled increase mandated Fisheries Commission (ASMFC) pro- ately, reduce incidental lobster injury by Amendment 4, which would have posed such a plan for state waters in and mortality during fi shing, establish changed the minimum size to 84.1 mm July 1996 and the plan was subse- the use of escape vents, prohibit pos- CL (Anonymous, 1994). Amendment quently adopted in 1997. Lobster fi sh- session of ovigerous females, standard- 5 also provided a defi nition for over- ing in Federal waters is now regulated ize gear marking, and license dealers fi shing, appointed committees (called under the provisions of the 1993 Atlan- by state and fi shermen by state and/or Effort Management Teams) for four tic Coastal Fisheries Cooperative Man- state and Federal waters (Anonymous, regional management areas (Gulf of agement Act (ACFCMA). 1998). This plan was referred to the New Maine nearshore, southern New Eng- In October 1996, the Sustainable England Fishery Management Council land nearshore, mid-Atlantic nearshore, Fisheries Act (SFA) amended the Mag- (NEFMC) in late 1978 for inclusion in and offshore), limited access to the fi sh- nuson-Stevens Act (MSA) to require the Magnuson-Stevens Act, but was not ery for 5 years (1995–1999), and pro- that the NMFS identify all overfi shed implemented until 1983. At that time, posed closed seasons, closed fi shing resources under the jurisdiction of fi sh- the FMP established a legal size limit areas, mandatory data reporting by all ery management councils (the MSA of 81 mm CL (3 3/16-inch CL), pro- active permit holders, and future length plan for American lobster, however, hibited the taking of ovigerous females, increases if deemed necessary (Anon- was withdrawn in 1999). In the case and required escape vents in all fi xed ymous, 1994; Miller, 1995). Amend- of an overfi shed resource which occurs lobster gear. Since its implementation, ment 6, approved in 1997, addressed predominantly in state jurisdictional the FMP has been amended six times. gear confl icts. Furthermore, under a waters along the Atlantic seaboard, the Amendment 1, approved in 1986, stan- separate legislative authority, under the ACFCMA provides for the develop- dardized gear marking in the offshore Atlantic Coastal Fisheries Cooperative ment, implementation, and enforcement fi shery. Amendment 2, implemented in Management Act (ACFCMA), the Fed- of effective interstate conservation man- 1987, imposed a series of four incre- eral government is allowed to suspend agement. The lobster was identifi ed as mental increases to the minimum fi sh- lobster fi shing within a state’s waters if an overfi shed resource in 1994. The able size of 0.8 mm CL (1/32-inch CL), that state is found to be in noncompli- ASMFC approved Amendment 3 to the effective in January 1988, 1989, 1991, ance with an affi liated interstate FMP Interstate Fishery Management Plan for and 1992, with the goal of reaching (Miller, 1995). Lobster in late 1997. Amendment 3 84.1 mm CL (3 5/16-inch CL) in 1992 According to Amendment 5, the Effort adopted an area approach to the man- (Anonymous, 1987b, 1998). Amend- Management Teams were required to agement of the lobster fi shery (similar ment 3, implemented in 1990, required establish stock rebuilding programs for to that in Amendment 5 of the with- the use of biodegradable escape panels their respective regions and to present a drawn FMP) with 7 Lobster Conserva- in traps to reduce the possibility of set of recommendations to the NEFMC tion Management Teams. It also adopted ghost fi shing by lost traps. by January 1995. The Council was then coastwide management measures such Despite these approved measures, required to submit a management plan as making it unlawful to possess lob- several industry associations, includ- by July 1995 designed to incorporate ster parts, speared lobsters, ovigerous ing the Maine and Massachusetts Lob- the recommendations of the manage- females, and V-notched females; requir- stermen Associations, requested a delay ment teams into the FMP for lobsters. ing ghost panels for non-wooden traps;

61(2), 1999 9 and limiting the landings of fi shermen In 1996, gear entanglements with chusetts, respectively, now comes from using nontrap fi shing methods to 100 whales, and specifi cally the endangered animals within one molt of legal size. lobsters per day. Additional coastwide right whale, also came to the fore. The In Rhode Island and central and west- measures included implementation of number of right whale entanglements ern Long Island Sound, egg production trap tags, designation of individual fi sh- occurring annually is unknown. Kraus within one molt of legal size is now erman’s areas of fi shing (beyond which (1990) estimated that 57% of the known 95%. If landings rely more and more on they could not fi sh), new escape vent right whale population exhibits scars newly-recruited lobsters to support the sizes, and maximum trap sizes. Three from entanglement. Generally, fewer fi shery, then the fi shery is compressing of the 7 areas were required to reduce than 10 entanglements are reported per egg production potential into a narrow the number of traps fi shed per fi sher- year, representing an unknown frac- size range (the prerecruits). The fear men each year with the goal of reaching tion of the total. Because the right of managers is that the abundance of 800 traps by the year 2000. The inshore whale population is so low, the poten- the prerecruits may fall, and the fi shery Gulf of Maine area (Area 1) also imple- tial removal of even one whale per will be severely affected (Lockhart and mented a maximum size gauge of 5 year requires regulatory action pur- Estrella, 1997). inch CL in January 1999. suant to the Endangered Species Act Furthermore, since size increases NMFS, citing requirements under the and Marine Mammal Protection Act have been implemented incrementally, Magnuson-Stevens Act to end overfi sh- (MMPA). In 1997 NMFS implemented and it takes about 6 years for stock ing, determined that Amendment 3 of an initial series of restrictions under the size changes (those in the size of the ASMFC’s plan did not fully address MMPA or Magnuson Stevens Act for spawning stock and thus in egg pro- the measures necessary to end over- lobster pot and gillnet gear to protect the duction) to affect recruitment, a period fi shing and thus did not provide ade- highly endangered right whale, as well of 20–30 years may be necessary before quate protection for conservation of as humpback, fi n, and minke whales. “so called” benefi ts, or the lack thereof, the lobster. However, NMFS did note These regulations included time/area would be apparent (Ricker stock size- that the ASMFC plan was an excellent closures and gear modifi cation require- recruitment model, Anonymous, 1987b). beginning for developing a seamless ments. As the take reduction plan However, based on the upward trend in co-management scheme within state evolves, additional restrictions are antic- landings since the late 1930’s, it would and Federal waters. NMFS was con- ipated to meet mandated MMPA goals. be rather diffi cult to assess the impact of cerned that trap reductions were not recent size increases on the fi shery. The low enough to signifi cantly reduce fi sh- Size Limits increase in landings might also be due ing effort and that the ASMFC plan Increases in the minimum size were to highly favorable environmental con- did not specify any conservation mea- considered a major regulatory tool for ditions (e.g. temperature) or a reduc- sures beyond 3 years despite commit- the lobster fi shery for the following rea- tion in lobster predators (Anonymous, ting to an 8 year schedule. In March sons: 1) larger size limits protect females 1991d; White, 1991; Pezzack, 1992), 1998, NMFS drafted a series of alter- so that they are allowed to molt, mate, although, Addison and Fogarty (1992) native management approaches in their and spawn at least once prior to being give an alternate view. However, since Draft Environmental Impact Statement captured, 2) fecundity increases with temperature also regulates the produc- (Anonymous, 1998). Public hearings increasing body size, and 3) higher yields tivity of predators (Estrella and Cadrin, on these measures were held through- in weight are expected with increased 1991), generally increasing their num- out the affected states through May 19, minimum sizes. However, since 1991, bers when a breeding stock is present, 1998 to obtain fi shermen’s views on no further size increases have taken the question of why lobsters were so the alternative options for managing place and the concern is that the fi shery recently abundant remains a mystery the fi shery in Federal waters. As per is now relying too heavily on animals (White, 1991). the Sustainable Fisheries Act, NMFS one molt away from recruitment into has until June 1999 to adopt a new the fi shery to provide the future gen- The Role of Shelter Federal management plan. In Decem- erations of the species. A recent stock Although the American lobster fi sh- ber 1999, Federal authority for manag- assessment for lobster (NEFSC, 1996) ery has experienced several fl uctuations ing lobster fi shing was transferred from noted that during 1983–94 in the Gulf over the last century of management, the Magnuson-Stevens Act to the Atlan- of Maine relative abundance of lobsters two specifi c incidents stand out. During tic Coastal Fisheries Cooperative Man- increased. But in 1995 relative abun- the early fi shery from 1874 to about agement Act (ACFCMA). Nonetheless, dance of recruits decreased while that for 1933, the major lobster producing state, many of the measures and prohibitions prerecruits increased; a similar trend was Maine, had a larger size limit for were carried over (limited access, no seen in Georges Bank (NEFSC, 1996). lobsters landed than that in effect in taking of berried, v-notched, or lobster Furthermore, egg production coming other states (Table 1), yet their land- parts, minimum sizes, etc.). New mea- from smaller size classes has been ings continued to decline. Then from sures included fi shing gear restrictions, steadily increasing since the 1970’s such 1933 (excluding a slight decline in trap tags, management areas, and con- that 60% and 90% of the egg produc- the 1970’s), landings remained stable sultations with ASMFC. tion in the Gulf of Maine and Massa- or were on an upward trend (Fig. 5).

10 Marine Fisheries Review These landings increased during the that occupy shelters will live closely previous and current landings statis- period of the taking of smaller-sized together, as long as they have individ- tics. During the early lobster fi shery, lobsters, despite assumptions of low ual shelter areas (Karnofsky and Price, larger lobsters were so numerous that average reproductive potential of the 1989). Shelter use and the number of crowding occurred (despite their pro- smaller females (and thus fewer poten- shelters used increases a few weeks clivities to disperse) and they dispersed tial recruits into the fi shery over time), prior to molting (Karnofsky et al., as much as possible, including to the lit- low survival of larvae and benthic set- 1989b), and lobsters may engage in rit- toral zone in waters so shallow that they tlers, and increased fi shing intensity. ualistic agnostic encounters for such were often exposed during low tide. Some of this increase in landings shelters (Scrivener, 1971). The premolt Pots were usually set out to a depth of may be due to the availability of shelter, condition may cause both the increase 36.6 m and captured lobsters that were which plays a role in the abundance of in aggression and the increase in shel- at least 92.3 mm CL (10.5 inch TL) in lobsters, infl uencing the sizes, number ter use (Tamm and Cobb, 1978; Atema size and about 648.6 g (1.43 lb) (Rath- of occupants, density, and survival in et al., 1979). Thus, the size distribution bun, 1884b). Increased fi shing inten- lobster populations (Scarratt, 1968; and/or availability of shelter may have sity removed virtually all of the larger Cobb, 1971; Howard, 1980). Cooper important ecological and evolutionary lobsters near shore. The availability of and Uzmann (1980) have documented consequences for the lobster. these near-shore grounds was further the use of shelter by lobsters from European lobster, H. gammarus, pop- reduced with the development of indus- their postlarval settling stage onward ulations seem to be locally size limited trial factories which discharged their throughout life, and shelter has been by the shelter characteristics of the sub- wastes directly into the coastal waters described by some as being a limiting strate (Howard, 1980). Wahle and Ste- (Cook, 1972) and with the develop- factor (Cobb, 1971; Fogarty, 1976; Fog- neck (1991) confi rmed that such size ment of coastal areas for marinas and arty and Idoine, 1986; Richards and limitation also exists within juvenile homes. Cobb, 1986). Artifi cial reef experiments American lobster, H. americanus, pop- The tremendous decrease in lobster have supported the theory that shelter ulations. Steneck (1989) reported that numbers near shore forced fi shermen is scarce, since lobsters readily occupy lobster population densities and body into deeper coastal waters. Because lob- reefs in numbers equal to or greater size corresponded to shelter availability sters were either removed from near- than those on natural grounds (Scar- which was controlled by regional geol- shore environments or forced offshore ratt, 1968, 1973b; Briggs and Zawacki, ogy. Population densities of 40–90 mm due to environmental degradation, the 1974; Sheehy, 1976). CL sized lobsters increased signifi cantly amount of space and shelter per lobster Recently, it has been suggested that when additional shelters were placed decreased offshore and, due to disper- settling juvenile lobsters are the most in the fi eld (Steneck, 1991). However, sal of the larger lobsters as seen by Ste- habitat-restricted (Hudon, 1987), and as lobster densities increased, the pro- neck (1991), the size of lobsters landed a “demographic bottleneck” has been portion of larger lobsters declined, sug- began to decrease. Once this situation proposed for this phase of life (Wahle gesting that larger lobsters move from equalized, populations of smaller lob- and Steneck, 1991). Lobsters must con- regions of higher density to lower den- sters began to increase. This hypothe- tinually fi nd larger shelters as they sity (Steneck, 1991). sis is illustrated in Fig. 6a–c showing grow, which subjects them to predation In addition, Skud (1969b) reported the increase in the density of lobsters risks that are inversely related to body that the size of lobsters occurring in an and demonstrating the decrease in the size (Wahle and Steneck, 1992; Wahle, area refl ected the fi shing pressure applied amount of space. The decrease in amount 1992a). Accordingly, the tight associa- to that population. Thus, the removal of of space corresponds to the fi ndings of tion with shelter during the early phases large lobsters would also result in popula- Steneck (1991) that lobsters lose their of life relaxes with an increasing body tions of smaller individuals, both inshore ability to live at higher densities when size, and larger lobsters can be found in and offshore. If these smaller individu- they get larger and that they are most the open without shelters, suggesting a als are then fi shed, the number of lob- abundant where they tend to be of smaller decline in the predation rate for inshore sters landed may increase, although the size. Furthermore, a shift in size, due lobsters (Wahle and Steneck, 1992). pounds landed would be less. Table 3 to the increase in the minimum legal Nevertheless, lobsters still need and shows that in 1942 when the legal size of size limits, may cause larger lobsters continue to use shelter throughout their lobsters was considerably smaller than to move into deeper offshore habitats entire lives. Shelter provides a place that in effect today or 100 years ago, the (Skud and Perkins, 1969; Steneck3). of protection during their vulnerable number of lobsters landed per pot was While increases in the size limits soft-shelled (molting) condition and for indeed greater. This suggests that smaller may force the larger lobsters to disperse overwintering, and is a prerequisite for lobsters are, indeed, more abundant than more widely, such that fewer are landed mating (Thomas, 1968; Atema et al., larger lobsters and suggests that studies 1979; Karnofsky and Price, 1989; Kar- similar to those by Steneck (1991) should nofsky et al., 1989a, 1989b, Cowan and be pursued. 3 Steneck, R. 1989. Ecological considerations on increasing the minimum legal size of lobsters. Atema, 1990). In naturalistic settings, Given those results, the following Univ. Maine, Ira C. Darling Mar. Cent., Walpole, an average of 70.3% of the lobsters scenario is hypothesized to explain the ME 04573. Unpubl. manuscr., 14 p.

61(2), 1999 11 Figure 6.—A) Inshore lobster fi shery (ca. 1800’s). Larger lobsters (> 90 mm CL) dominated this fi shery and were caught close to shore with traps set out to a depth of about 36.6 m (slashed line). Each square represents 10,000 m2 and each open circle 10 lobsters of a size > 90 mm CL. B) Inshore lobster fi shery (ca. 1930 to 1990). Smaller lobsters (< 90 mm CL) after removal of larger lobsters (> 90 mm CL), due to intense fi shing pressure. Smaller lobster densities equalized as a result of decrease in the minimum legal size limit. Each square represents 10,000 m2 and each asterisk 200 lobsters of a size < 90 mm CL. Slashed line is the general area where lobstermen set their pots, at a depth of about 36.6 m (121 feet). C) Inshore lobster fi shery (ca. 1990 to 1993). As a result of increased size limits, larger lobsters diffuse from regions of high density to those of low density. Likewise, once lobster size changes, lobster density also changes, such that fewer large lobsters live in the same area. Asterisk represents lobsters of a size < 90 mm CL; open circles represent lobsters of a size > 90 mm CL. Slashed line is the general area where lobstermen set their pots, at a depth of about 36.6 m (121 feet). per pot, intense fi shing efforts may abundance of the prerecruits (those one able clip. These new regulations render remove them quickly. Currently, a min- molt away from recruitment) declines the traps nonfunctional in about 1 year imum size of 82.5 mm CL (3 1/4-inch for whatever reason (habitat destruc- (Blott, 1978; Anonymous, 1987b, 1989, CL) is in effect for all major lobster- tion, unfavorable environmental con- 1991d). Once they cannot retain a lob- producing states (Table 1). The mean ditions, etc.). If the numbers of those ster against its will, they become added size of lobsters landed has increased recruiting into the fi shery is reduced, shelter. Steneck (1987) reported that to 87.2 mm CL in New York (Briggs, then landings would eventually begin removal of pots from a highly popu- 1992), 88.6 mm CL in Maine (Krouse to decline. Thus far, the relative abun- lated lobster area resulted in a consid- et al., 1990), and 88.8 mm CL in Mas- dance of prerecruits has been steady erable drop in abundance of lobsters sachusetts (Cadrin and Estrella, 1993). or increasing in most areas (NEFSC, after a 30–45 day period. Dow (1980) These sizes are similar to the 87.8 and 1996). reported that scuba divers observed lob- 92.3 mm CL minimum sizes in effect Pot loss, which provides shelter sters partially entering traps, consuming in 1889 and 1892, respectively. As in where it may already be limiting, may the bait, and returning to their burrows. those years, lobster landings declined also affect the density of lobsters. Shel- In areas where intensive fi shing is con- from 1991 to 1992 by 2,493 t or 9% don and Dow (1975) estimated that ducted, 80% of the lobster’s diet may in the United States (Fig. 4). Removal an average of 20–25% of pots were come from baited pots (Steneck, 1987). of the larger lobsters from the fi shery lost in the Maine lobster fi shery annu- Furthermore, escape vents used today over a relatively short period of time, ally, and 80% of these pots were capa- allow undersized lobsters to enter and will tend to result in a fi shery reliant ble of “ghost fi shing” for an average leave the pots at will and thereby pro- upon the newly recruited lobsters that of 2 years before they become non- vide them with food and temporary might initially be found at higher den- functional. However, effective 28 May shelter (Landers and Blake, 1985). sities. This could result in a spike in 1992, traps were required to possess These observations show the infl u- the landings, as was seen in the early either a ghost panel made from bio- ence that pots may have had, and may 1990’s. However, such a reliance on new degradable material or to have their continue to have, in increasing the abun- recruits might present problems if the escape vents attached with a biodegrad- dance of lobsters by providing them

12 Marine Fisheries Review with more shelter and a food source. (1990), Pringle and Burke (1993), and water to renew itself every 12 hours, As further support of this theory, the Miller (1995). following the tidal ebb and fl ow. Lob- lobster fi shery has utilized the pot to sters were fed shellfi sh and fi sh during catch lobsters since the mid-19th cen- The Development of confi nement (Moquin-Tandon and Sou- tury (Cobb, 1901; Dow, 1949). If we Artifi cial Propagation beiran, 1865). assume only a 1% loss of pots per year Despite stringent laws prohibiting the The Concarneau laboratory had a for the overall lobster fi shery (instead of landing of “berried” lobsters in both total of 85 tanks, small and large, where the 20–25% quoted for Maine alone), Canada and the United States, a sig- observations were made on lobster then the number of pots that have nifi cant number of females with eggs hatching, metamorphosis, and behav- been lost over a period of 150 years were scrubbed (eggs removed) and sent ior. In 1865, Guillou and Coste reared would have added signifi cantly to the to canning factories or to market (Her- a considerable number of lobsters and shelter made available for lobsters. A rick, 1895, 1911a; Smith, 1898; Anony- recorded their length and weight for decrease in space inshore and move- mous, 1906; Wilder, 1954, 1965). This Stages IV through XIV, and they were ment of larger lobsters offshore, along frequent sacrifi ce of eggs was believed reported by Moquin-Tandon and Sou- with added shelter inshore, could cause to be a major contributing factor in the beiran (1865) to have remarked, “we a clumping effect where smaller lob- lobster decline in the late 1800’s. Con- may hope that it will be possible to sters would live closely together. Such sequently, artifi cial propagation (cul- regenerate the fi shery on parts of our clumping effects have been observed by ture) of lobsters was seen as a way to coast.” Guillou and Coste noted that Karnofsky and Price (1989) and Kar - reverse the loss of the millions of eggs the rearing of lobster larvae was tech- nofsky et al. (1989a, b) in both the lab- destroyed by scrubbing. The eggs saved nically easy, which led to the idea of oratory and fi eld experiments, respec- could be hatched and released to pre- “reseeding” and enhancing the waters tively. If the above hypothesis holds serve, or possibly increase, the supply (Latrouite4). true, larger size limits for lobsters may of lobsters (Moquin-Tandon and Sou- have great implications for future in- beiran, 1865; Wood, 1869; Sars, 1879; Lobster Parks shore landings, as they did in the past. Ryder, 1886a; Rathbun, 1892; Herrick, Because of the successes in France, The highest densities of large lobsters 1894, 1895; Bowers, 1900; Bumpus, the establishment of similar operations (>90 mm CL) reported is 50 per 10,000 1901b; Mead and Williams, 1903; in the United States for H. americanus m2; that same space can accommodate Herrick, 1911a; Scattergood, 1949b; was recommended (Wood, 1869). Two 5,000 smaller lobsters (<90 mm CL)—a Wilder, 1965). Various culture methods “parks” were established: one in Massa- hundredfold increase in the number of were employed, some of which approx- chusetts in 1872 and the other in Maine animals (Steneck3). Thus, smaller size imated the natural conditions of tidal around 1879 (Rathbun, 1886). Accord- limits result in the taking of more lob- ponds and man-made (enclosed) basins ing to Smith (1898), “parking” involved sters since they occur at higher den- or parks, and others which used care- retaining egg-bearing female lobsters sities and cause landings to increase fully controlled laboratory conditions. in natural, enclosed basins where they (Skud, 1969a; Skud and Perkins, 1969; were allowed to hatch their eggs. This Uzmann et al., 1977; Fogarty et al., Early Work form of culture was also used in Stavan- 1982). Clearly more research is needed In 1858, Guillou began culturing ger, Norway, in 1873–75 and in Canada to establish estimates of minimum and experiments with the lobster H. gam- around 1903 (Corrivault and Tremblay, maximum densities for variously sized marus at the laboratory of Concarneau 1948, cited in Scattergood, 1949b). In lobsters, with respect to the amount of in South Brittany, France (de Maude, Norway, man-made enclosures (parks) space they require and the habitat limi- 1858, as cited in Latrouite and Lorec, in the natural environment were built, tations they face. 1991; Latrouite4). Adult lobsters were and egg-bearing lobsters, H. gammarus, For comprehensive reviews on other maintained in natural and man-made were placed inside cork boxes about protective measures and regulations for fi sh ponds and/or in tanks in the labora- 1.5 m square × 0.6 m deep within the the management of both H. america- tory. The fi sh ponds consisted of natu- enclosures. Lobsters were successfully nus and H. gammarus lobster fi sheries rally enclosed basins formed by rocks, hatched and reared through their larval see Herrick (1911a), Bennett (1980), occupying an area of about 1,500 m2 stages until they reached Stage IV or Cobb and Phillips (1980), and Cobb where the bottom consisted of sand, V when they exhibited a crawling habit and Wang (1985). For emphasis on the mud, and rocks. These ponds were (reported by Rasch, 1875; Sars, 1879). American lobster fi shery see Anony- divided into six different sizes, made Appelløf (1909a) also conducted lob- mous (1983a, b; 1986, 1987b, 1989, by thick barriers capable of withstand- ster park experiments in 1892, with 1991d), MacKenzie and Moring (1985), ing the pressure of the sea. They had a limited success in Kvitsøy, Norway. and Northeast Marine Fisheries Board gate which opened freely to allow the Despite these early successful efforts, (1978). For specifi c reviews on regula- it was soon determined that this form tions during the course of their develop- of natural larval culture was not an effi - 4 Latrouite, D. 1992. Ifremer, Centre de Brest, B.P. ment in Canada and Maine see Wilder 70-29280 Plouzane, France. Personal commun., cient way to replenish the supply of lob- (1965), Pringle et al. (1983), Kelly 24 Jan. sters; therefore, these experiments were

61(2), 1999 13 discontinued (Smith, 1898; Bowers, at Wickford, R.I., began a series of sys- 1900; Appelløf 1909a; Corrivault and tematic experiments to fi nd a way to rear Tremblay, 1948, cited in Scattergood, postlarvae (Bumpus, 1901b; Barnes, 1949b). 1906a; Emmel, 1908; Mead, 1910). Many different devices were adapted Hatching Jars and tried (e.g. artifi cial pools, enclo- Meanwhile other researchers actually sures made of wire screen which were hatched lobster eggs in jars. In Scotland, submerged or fl oating in water, glass Saville-Kent (1884) hatched the eggs jars of various sizes, and huge canvas of H. gammarus and kept the larvae in bags and boxes), but all proved inef- small jars where they were maintained fective (Bumpus, 1901b; Sherwood, and fed with minced fi sh, and their water 1905). However, between 1898 and was changed every day. He reared lob- 1899, Bumpus succeeded in rearing sters to the size of 1 inch, or about several hundred larvae to Stage IV in Stage V–VI, when they would hide upon scrim bags (bags made of light, loosely release onto rocky grounds (reported in Figure 7.—McDonald Hatching Jar. woven cotton or linen) at Woods Hole Williamson, 1905, and Herrick, 1911a). From McDonald (1883). (Mead, 1905, 1910). That same year (1883), the Norwegian Despite this success, the previous G. M. Dannevig, began experimenting uous supply of lobsters in the future failures to rear larvae resulted in the with hatching of “detached eggs.” He (Herrick, 1911a, b; Barnes, 1939). For suggestion that environmental condi- successfully hatched the eggs, but the a full description of the automatic tions at Woods Hole were not ideal for lobsters experienced high mortality after hatching jar (the Downing, Chester, the development of the young lobsters. the fi rst larval stage. This mortality was or McDonald hatching jars) and the The same apparatus used at Woods attributed to disease and/or cannibalism McDonald tidal box or Nielsen incuba- Hole was tested at various other loca- (Dannevig, 1885a, cited in Scattergood, tor, the reader is referred to McDonald tions (e.g. Annisquam River, Glouces- 1949b). Dannevig continued his work, (1883), Brice (1898), Galtsoff (1937), ter, Mass.; Orr’s Island, Maine; and and in 1885 he reared about 200 lobsters Havinga (1921), and Roché (1898, cited Wickford, R.I.) (Bumpus, 1901b; Sher- through the fi rst three larval stages, with in Scattergood, 1949b). wood, 1905). Based on the results of a survival rate of around 47% (Dann- these experiments (Sherwood, 1905), evig, 1885b). Several of these lobsters Scrim Bags the other stations were abandoned for were held and reared for 9 weeks while Despite the success of the McDonald the season, and all efforts were trans- being fed on the soft parts of crabs. They jar, the question of larvae survival after ferred to Wickford, R.I., which had molted fi ve times, measured 21 mm release into nearshore waters arose, the highest water temperatures and the (total length) on Stage V, and were of a and Herrick (1894) speculated that not best availability of natural plankton greenish-gray color (Dannevig, 1885c). more than 2 in 10,000 larvae survived (Bumpus, 1901b; Sherwood, 1905). their pelagic life. Later, other research- McDonald Jars ers estimated that not more than one- Water Agitation In 1883, a seaside laboratory was tenth of 1% survived (Mead, 1905; Research was then directed towards set up at Woods Hole, Mass., by the Sherwood, 1905). Therefore, research- the design of an apparatus for keeping U.S. Commission of Fish and Fisher- ers concluded that hatching eggs and the water agitated in the scrim bags so ies (Smith, 1908), where small quanti- liberating the Stage I larvae was not an larvae would not settle to the bottom ties of H. americanus lobster eggs were effective way to replenish or improve where cannibalism was most prevalent. successfully hatched. However, large- the natural supply of lobsters (Dann- Originally, paddles were used to stir scale artifi cial propagation did not begin evig, 1885a, cited in Scattergood, the water to keep it in constant motion. until 1886, after completion of the new 1949b; Herrick, 1894; Mather, 1894). However, this method was too time laboratory in 1885 with fl owing seawa- Instead it was recommended that the consuming because it had to be done ter (Rathbun, 1892; Bowers, 1900). In larvae be reared to their bottom-crawl- manually. In 1901, new equipment was 1886, experiments progressed so well ing (benthic) stage, in an effort to reduce devised, consisting of a two-bladed pro- that millions of eggs were detached from larval mortality that occurred in nature. peller designed and installed by Sher- female lobsters and hatched in McDon- Mather (1894, 1900) suggested that the wood (Mead, 1902, 1910; Fig. 8). This ald hatching jars. The McDonald larvae be reared individually in a tank propeller kept the water in constant cir- hatching jar (Fig. 7) gave the best sur- or in small compartments. However, he culation inside the rearing bags (0.9 m vival—as high as 93% from egg to the concluded that it would not be possible, in diameter and 1 m deep), and with fi rst stage larvae (Smith, 1898; Bowers, at that time, to feed a million or more this new system 8,974 Stage IV lobsters 1900). Billions of fi rst stage larvae individually housed lobsters. were successfully reared from eggs that were hatched and released directly into In 1898, Bumpus, Mead, and their were stripped from females (Sherwood, nearshore waters to ensure a contin- associates at Woods Hole, Mass., and 1905). The bags were suspended from

14 Marine Fisheries Review frames attached to a fl oating house-boat (laboratory) directly on the water (Bea- sley, 1904; Middleton, 1909). Later, the bags were enlarged to 3.6 m2 by 1.5 m deep, so that rather than scrubbing eggs, female lobsters could hatch their eggs within crates placed directly into the bags (Barnes, 1906a). The fi rst system is fully described in Mead (1902) and Sherwood (1905), and that of the second system is in Mead and Williams (1903). It should be noted that Mead’s “original method” bears some resemblance to Nielsen’s fl oating incubator (Firger, 1974), which may have been modifi ed for Mead’s system. The success of the enlarged bags was evident by the high output of lobsters which followed. By 1920, Wickford’s hatchery reached the million mark in rearing Stage IV and some Stage V lobsters (Havinga, 1921, cited in Scatter- good, 1949b). The highest production was obtained in 1936, when Wickford reared 1.7 million Stage IV lobsters (Carlson, 1955). In conjunction with rearing Stage IV lobsters, success was also achieved in rearing Stage V lobsters in substantial numbers. The meth- Figure 8.—Agitation unit used at the Wickford, RI hatchery for hatching and rear- ods were similar to those of rearing ing larvae. A) scrim bag or car where lobster larvae are contained; B) engine to drive × Stage IV, and survival rates were esti- propeller system; C) propeller or fan, 14 inches long 5 inches wide, made from cypress and screwed into a piece of maple, the other end of which is connected to a mated to be up to 80%, with an average tee. (Note: second bag removed to show propeller system.) From Sherwood (1905). of 60% to Stage V (Barnes, 1907). Not- withstanding the above success, many future hatcheries were still built upon Cobb, 1932). Barnes (1939) (and in et al., 1974; Fig. 11). The fl ow pattern the idea of hatching eggs and releasing Anonymous, 1930) reported that this of this system resulted in constant tur- Stage I lobsters. system was preferable to that used at the bulence which maintained a homog- Wickford, R.I. hatchery. His conclusion enous distribution of the larvae and The Banning Box was based on the fact that this system their food and minimized interactions In 1929 at the Noank hatchery in could be used indoors, had greater pos- between larvae that could culminate in Connecticut, Capt. Banning constructed sibilities of refi nement, was less expen- cannibalism (Hughes et al. (1974) gives an indoor rearing system, based upon sive to run, and resulted in higher construction and hydraulic characteris- the Norwegian plan (Carlson, 1955), survival rates of Stage IV lobsters. John tics). At densities of 2,000 larvae per consisting of a hatching trough (Fig. Hughes, the fi rst director of the Mas- kreisel, survival in this system was esti- 9) in which female lobsters were held sachusetts State Lobster Hatchery and mated at 75–85%, with larvae being fed until they hatched out their larvae, and Research Station, further refi ned the Artemia salina brine shrimp (Hughes et a square wooden box about 400 mm Banning Box by replacing the square al., 1974). × 400 mm × 350 mm (Fig. 10), into wooden units with cylindrical, fi ber- In Helgoland, Germany, Greve which the larvae were transferred. The glass pots (350 mm deep, 400 mm in (1968) also designed a “planktonkrei- latter became known as the “Banning diameter). The square circulators were sel” for culturing and rearing planktonic Box” (Dexter, 1986). Circulation in the also replaced by round ones (Fig. 11; marine organisms. Unlike the “Hughes box was made by introducing water Anonymous, 1965). Pot,” an inside-sand fi lter arrangement into a circulator containing small holes (Fluchter, 1964) was incorporated at the bottom from which fi ne jets of The Hughes Pot or Kreisel directly inside the kreisel to provide water emerged in an upward motion. These refi nements resulted in the devel- self-contained fi ltration of the culture Larvae and food were kept in a constant opment of the “Hughes Pot,” “plankton- medium (Greve, 1968). Although this upward motion (Anonymous, 1930; kreisel,” or just simply “kreisel” (Hughes system was successful for the culture

61(2), 1999 15 Figure 9.—Hatching trough used at the indoor Connecticut State Hatchery in connection with the Banning Box. The trough is 14 inches (35.56 cm) wide × 8 inches (20.32 cm) deep and consists of 7 compartments (A) which hold female lobsters. Each compartment is separated by a 3/8 inch mesh screen (C) which allows the larvae to pass through. The last compartment’s (D) fl ashboard has its top cut away so that the larvae fl ow through the opening to the next compartment (B) 20 inches long by 14 inches wide. Larvae are prevented from entering the outfl ow by the #18 mesh screen (E) at the end of the compartment. All compartments are covered, except for compartment B, which is left open to allow the larvae to display their phototactic responses. From compartment B, the larvae are transferred to the Banning Box. From Anonymous (1930).

and rearing of ctenophores, chaeto- gnaths, and meroplanktonic crustaceans, the “Hughes Pot” became the most popular kreisel for culturing larval lobsters. Since then, other modifi cations have been added to the basic kreisel. Ser- fl ing, Ford and Van Olst incorporated 16 kreisels into four rearing systems and developed added features of temperature control, fi ltration (Serfl ing et al., 1974a; Fig. 12), and automatic feeding devices for the larvae (Serfl ing et al., 1974b; Fig. 13). Stocked at densities of 4,000 larvae per kreisel and with an average survival rate of 75%, these four rearing systems could yield 48,000 Stage IV lobsters every 10–12 days at temperatures of 22°C (73°F)—in other words, nearly half a million larvae could be produced in about 120 days (Klop- fenstein and Klopfenstein, 1974; Ser- fl ing et al., 1974a). Color Morphs Due to the seeming lack of evidence that hatchery operations (particularly in the case of “seeding” coastal waters with postlarvae) were having a positive effect on lobster landings, researchers began to investigate the use of color morphs of the lobster. Through controlled mat- ings, red, blue, white, orange, and Figure 10.—Banning Box. Redrawn from Anonymous (1930) and Cobb (1932). multiple-colored lobsters could be cre-

16 Marine Fisheries Review ated (Anonymous, 1966, 1967; Hughes, 1968a; Shleser, 1971). These oddly colored lobsters are extremely rare in nature, appearing only once in every 15 million lobsters (Syslo, 1986); thus, it was hoped that color morphs could serve as a natural tag through which hatchery efforts could be assessed. Today, this is actually being accomplished (Plante, 1989; Irvine et al.5; see also the section Aquaculture Potential: Resource Enhancement), although survival rates of these color morphs has not been assessed, compared to survival of wild type coloration lobsters. Hatchery Establishment The success of these early experimental culturing stations resulted in the construction of more than 20 hatcheries in the United States, Canada, Norway, France, and the United King- dom between the years 1885 and 1954 (Scattergood, 1949b; Carlson, 1954, Figure 11.—Hughes Pot or Kreisel. From Hand et al. (1977). Used with permission. 1955; Kenslor, 1970; Bardach et al., 1972). In Canada alone, there were 14 such hatcheries in the areas of Bay 27,700 fi rst stage larvae were success- three eyelet holes through their uropods. View, Nova Scotia; the Southern Gulf fully hatched (Anonymous, 1892). Egg- If these lobsters were later recaptured, of St. Lawrence, and on the outer coast bearing females were caught by local they could not be resold (Anonymous, of Nova Scotia (Wilder, 1965). lobstermen, and their eggs were hatched 1919, 1920). Originally, the purpose of these Fed- in McDonald hatching jars (Anony- eral and state-supported hatcheries was mous, 1899). In 1900 the Cold Spring Connecticut Propagation to hatch lobster eggs artifi cially. During Harbor hatchery released 2.4 million By an act of the legislature in 1905, their operations, experiments were con- larval lobsters (375,000 more than the Connecticut established a hatchery at ducted to determine both the best meth- previous year) into Long Island Sound Noank where the original purpose was ods of hatching the eggs and the best (Wood, 1901). Despite this success to hatch detached eggs in McDonald kind of apparatus. and that of succeeding years in which hatching jars. In 1906 alone, 20.1 mil- 2–3 million fi rst stage lobsters were lion Stage I larvae were hatched and New York Hatcheries released, this hatchery was discontin- liberated (Anonymous, 1906). By 1936, Due to the extremely low lobster ued in 1902 due to the lack of a boat for the Noank hatchery was using the Ban- populations in New York waters during seeding efforts and a dispute between ning Box and successfully reared and the late 1880’s, lobster propagation was New York and Connecticut over fi sh- released over 500,000 Stage IV lob- begun in 1886, when Fred Mather, ing territories (Wood, 1903; Walters, sters into the coastal waters of Connect- superintendent of the Cold Spring 1904; Anonymous, 1909). Around 1909, icut (Carlson, 1955). The hurricane of Harbor Fish Hatchery, obtained 50,000 an auxiliary hatchery opened at Fort 1938 destroyed the Noank hatchery, but eggs and 5,000 larval lobsters from Pond Bay, Montauk, N.Y. Rather than it reopened in 1940, and continued its the U.S. Fish Commission at Woods using methods currently in practice, this operations, rearing in excess of 3 mil- Hole, Mass. Although all of the eggs operation hatched fi rst stage-larvae in lion Stage IV lobsters during 1940–54. died in transit, 4,000 of the larvae sur- fl oating boxes used for hatching shad Then the Board of Fisheries and Game vived and were liberated into Cold (Walters, 1911). In 1910, 7,005,180 fi rst concluded that hatchery operations were Spring Harbor (Mather, 1887). By 1891, stage larvae were hatched, while in 1911 uneconomical and recommended its 45,100,000 larvae were hatched (Anon- closure (Anonymous, 1955). Carlson ymous, 1912). This hatchery continued (1955) gives a complete evaluation of 5 Irvine, C. M., B. Beal, R. C. Bayer, S. Chap- man, and D. A. Stubb. 1991. The effi cacy of blue operations until 1918 when the state the recommendations. When the Noank colormorphic lobsters in determining the feasi- passed a regulation that required offi - hatchery was transferred to the Uni- bility of hatch release programs. Manuscr. Rep., 16 p. Avail. from The Lobster Inst., Univ. Maine, cials to purchase egg-bearing females versity of Connecticut, egg-bearing Orono, ME. from lobstermen and mark them with female lobsters were purchased from

61(2), 1999 17 affected species. NMFS approved funds to place six small artifi cial cobble reefs off Narragansett Bay into which hatchery reared, microwire tagged postlarval lobsters will be seeded onto three reefs, while the other three reefs will be used to make comparisons (Nor’Easter, 1996). These reefs are about 10 × 20 m each, separated by 30 m and were placed along the shoreline at Dutch harbor to a depth of about 3 m. Each reef consists of two sub-areas 10 × 10 m each, one with cobble (10–20 cm in diameter, 25 cm above the seabed), and the other with boulders (20–40 cm in diameter, 50 cm above the seabed) (Castro, 1997). With funds from Uni- versity of Rhode Island Sea Grant the reefs will be monitored over a period of 5 years to determine growth, immigra- tion and emigration rates, competition between hatchery-reared and wild lobsters, and additional recruitment to the reefs by wild postlarvae. Some believe that this study may be the fi rst step to the reestablishment of a Rhode Island hatchery (Nor’Easter, 1996). During the summer of 1997, tagging of lobsters and monitoring of the reefs were conducted with a total of 1,036 lobsters being caught at three sites: 1) artifi cial reefs, 2) off the reefs, and 3) Dutch Island. These lobsters ranged in size between 1.5 inch CL to 4 inch CL, with an average of 0.86 lobsters per m2 (Castro, 1997). In 1998, Mount Desert Oceanarium Lobster Hatchery in Maine Figure 12.—Recirculating culture system incorporating several kreisels with the shipped 2,000 postlarvae to the Univer- added features of temperature control and fi ltration. From Serfl ing et al. (1974a). sity of Rhode Island Fisheries Center Used with permission. to join the 1,500 postlarvae that were reared at the university hatchery and the 300 provided by the New England the lobster fi shermen and returned to duction to 1.7 million Stage IV lob- Aquarium. Each lobster was to be the state waters as a conservation mea- sters (Carlson, 1955). Unfortunately, a micro-tagged for identifi cation and sure (Anonymous, 1955). 1938 hurricane destroyed the hatchery placed on the artifi cial reefs for further at Wickford, R.I., but it was rebuilt. studying (Tuttle, 1998). Rhode Island Propagation Another hurricane destroyed the Wick- The Wickford, R.I. hatchery, which ford hatchery again in 1944, but from Maine Propagation was highly active after the turn of the 1940 through 1943, it hatched, reared, As early as 1883, the legislature in century, was the site of many of the and released into local waters over 5 Maine passed an act allowing the artifi - new efforts to fi nd the most effective million Stage IV lobsters. Although lob- cial propagation of lobsters. By 1887, rearing technique. By 1920, Wickford’s ster rearing ended in 1944 due to the R. T. Carver was granted the right to hatchery reached the million mark in second hurricane, the artifi cial hatching propagate lobsters in Carver’s Pond, rearing Stage IV and some Stage V lob- of eggs continued until 1951 (Carlson, Vinal Haven, Maine. Unfortunately, this sters in scrim bags with water agitation 1955). attempt was a failure, owing to the mud (Havinga, 1921, cited in Scattergood, In 1989 a large oil spill resulted in a in the pond which killed all the larvae 1949b). By 1936, they increased pro- settlement to the state for restoration of (Cobb, 1901). However, that did not

18 Marine Fisheries Review deter further attempts, and by 1904, a Federal hatchery was established at McKown’s Point, Boothbay Harbor. Activities consisted of hatching detached eggs in McDonald hatching jars and releasing Stage I larvae, until about 1938 when a few Stage IV lobsters were reared experimentally and released (Taylor, 1950). Meanwhile, during the mid-1930’s a Maine delegation visited the Connecti- cut hatchery and, on the basis of their observations, established a state hatchery in Boothbay Harbor, just southeast of the Federal one (Dow, 1949; Carl- son, 1955; Stickney, 1986). The Maine state hatchery ran in conjunction with Figure 13.—Automatic feeding unit for the recirculating larval culture system. From the Federal hatchery in that eggs were Serfl ing et al. (1974b). Used with permission. hatched in the Federal facility and transferred to the state hatchery for rearing (Taylor, 1950; Taylor and Dow, were not ideal for raising postlarvae, and Canadian Propagation 1958). By following the design used in suggestions were made to try other loca- Connecticut with some modifi cations tions. Releasing of Stage I larvae con- In 1890, the fi rst lobster hatchery (Anonymous, 1936), Maine reared and tinued, however, until 1917, when the opened in Newfoundland. Originally released in excess of 2.3 million Stage Gloucester hatchery was closed (Sher- eggs were incubated in glass jars with IV lobsters into their state waters over wood, 1905; Carlson, 1955). Around aeration, but by 1893 this method of a 10-year period (Taylor, 1950; Taylor 1937, the Gloucester hatchery reopened hatching eggs was replaced by the and Dow, 1958). to hatch lobster eggs artifi cially and rear Nielsen incubator (Roché, 1898). By them to postlarvae, but in 1953, this 1894, a total of 21 hatching stations Massachusetts Propagation hatchery once again ceased its activities. were established with anticipation of In 1885, lobster culture was con- In 1939 Massachusetts appropriated the number reaching as many as 68. ducted on a very small scale by Rich- money to build a state lobster hatchery Each station hatched about a million ard Rathbun and Captain H. C. Chester (Barnes, 1939) in Oak’s Bluffs on Mar- eggs each year (Anonymous, 1895). In working in Woods Hole and Glouces- tha’s Vineyard; however, the actual con- 1891, a hatchery, consisting of a build- ter, Mass. By 1886, several million eggs struction did not begin until after World ing 45 × 35 ft, opened at Bay View, were hatched using the McDonald tidal War II (Anonymous, 1963). Unlike the Nova Scotia (Corrivault and Tremblay, box, the Chester jar, and the McDonald hatcheries before it, this hatchery’s pur- 1948). Eggs were collected from lob- hatching jars (Bowers, 1900). Larval pose was both to rear and release Stage sters supplied by the canning factories development was fi rst observed at this IV lobsters and to study the basic biol- and were incubated in McDonald hatch- time (Ryder, 1886b). During 1887–90, ogy of the lobster (Anonymous, 1963; ing jars with aeration and agitation of over 17 million eggs were collected and Kenslor, 1970; Syslo, 1986). Beginning the water (Roché, 1898). From 1903 hatched with a 54% success rate. At the operations in 1951, the Massachusetts to 1912, 15 additional hatcheries were same time, the various methods were State Lobster Hatchery and Research established: 8 in Nova Scotia, 3 in New assessed, and the researchers deter- Station used rearing equipment similar Brunswick, 2 on Prince Edward Island, mined that the McDonald jars outper- to that developed by Capt. Banning at and 2 in Quebec (Corrivault and Trem- formed the McDonald tidal box and Connecticut’s Noank hatchery (Carl- blay, 1948). The combined releases of the Chester jar. From 1890 to 1897, son, 1954; Anonymous, 1964; Hughes larvae from these hatcheries was well billions of eggs were hatched in the and Matthiessen, 1967). From 1951 over 2 billion (Herrick, 1911a), but McDonald jars, with an 81–93% suc- through 1963, 2 million Stage IV lob- in 1917 it was concluded that 1) the cess rate (Bowers, 1900). sters were reared and released, averag- female lobster was a better incubator During 1898–99, emphasis shifted to ing 150,000 annually, with a survival than artifi cial methods and 2) there rearing postlarvae instead of larvae. Sev- rate of about 30% (Anonymous, 1963). was no evidence that hatchery efforts eral methods to do so were explored, Although annual releases increased to were enhancing the natural populations; including submerging the larvae in about 500,000, the hatchery’s opera- wooden cars and canvas bags (Sherwood, tions were terminated in 1997 so that 6 Estrella, B. 1998. Massachusetts Department of 1905). By 1903, researchers felt that its focus could shift to research projects Marine Fisheries, 50A Portside Drive, Pocasset, environmental conditions in Gloucester only (Estrella6). MA 02559. Personal commun., Feb.

61(2), 1999 19 Table 4.—Approximate number of days required to pass through larval and postlarval stages at various temperatures.

H. americanus1 H. americanus2 H. americanus3 H. gammarus4 Temp. (°C) I II III IV I-IV I-V I II III IV I-IV I-V I II III IV I-IV I-V I II III IV I-IV I-V

8 20 26 42 10 14 15 25 49 54 103 13 18 25 54 56 110 12 10 11 15 32 36 68 8 11 16 30 35 65 14 7 8 11 25 26 51 32 4 6 7 17 15 6 7 9 23 22 45 27 4 5 6 15 16 5 6 9 22 20 42 26 4 5 6 15 18 3 5 7 18 15 33 4 4 6 20 14 34 15 3 4 5 12 20 2 4 6 14 12 (26) 12 3 4 4 11 22 2 4 5 11 11 (22) 3 3 4 14 10 24 9 2 3 4 9

1 Source: Templeman (1936). 2 Source: Mackenzie (1988). 3 Source: Hughes and Matthiessen (1962). 4 Source: Havinga (1929), as cited in Scattergood (1949b). thus all hatcheries were closed (Knight, (1950) reviewed lobster-rearing efforts because of their naiveté. In the absence 1918). Through the years several hatch- in Maine, evaluating hatchery effi ciency of clear, easily recognizable tags for eries were established in Canada for based on the percentage of survival hatchery-reared lobsters, the impact transplantation programs (Ghelardi and from larvae (Stage I) to post-larvae they have on natural populations will Shoop, 1968, 1972), to supply Stage (Stage IV). Carlson (1955) reviewed continue to be diffi cult to assess. How- IV postlarvae for studies on their ecol- the effi ciency of the Connecticut hatch- ever, the best estimates of the economic ogy in coastal regions (Roberts, 1984), ery, considering biological, social, and impacts released lobsters have made and with the purpose of exploring the economic factors. Both reports empha- upon the natural population at very low biology of the lobster and aquaculture sized that hatcheries were not the proper survival rates is presented in Table 4 potential (Waddy, 1988). vehicle for enhancing natural lobster (Syslo7). The focus then shifted to broodstock stocks. However, during the years that management, where females and males the Wickford hatchery was in opera- Laboratory and Field were kept in captivity on a temperature- tion in Rhode Island, landings increased Research Stemming From photoperiod cycle that mimicked that of from <180 t to >726 t, which some Hatchery Experimentation their natural habitat (i.e. one where the researchers attributed to the releases of water temperature rises above 12°C in postlarvae (Carlson, 1954). Additional During early hatchery operations, the summer and drops to 5°C or lower anecdotal evidence exists for other many aspects of lobster life history in the winter). Normal female molting stocking programs (New York, Rhode were examined. Ultimately these stud- and spawning cycles were maintained Island, and Massachusetts) where sight- ies were justifi ed as being important for and, by manipulating the photoperiod/ ings of abundant small lobsters have improvement of culture methods and for temperature cycle, year-round produc- been assumed to be a product of hatch- providing life history data important for tion of larvae was achieved (Waddy ery reared stock (Anonymous, 1899; fi sheries management. However, these and Aiken, 1991; Aiken and Waddy, Mead, 1905; Bardach et al., 1972; Syslo, studies also provided the basis for many 1995). Details of this broodstock facil- 1986). current experiments. While it is impos- ity located at the biological station at St. In contrast, although the Massachu- sible in such a review to describe every Andrews, N.B., can be found in Aiken setts stock enhancement program has experiment on homarid lobsters, brief and Waddy (1995). Despite the suc- released millions of Stage IV lobsters outlines of the most important work are cess of this program, the Department of into coastal waters, documented land- presented. Fisheries and Oceans closed this facil- ings have not increased signifi cantly. Embryology ity in the late 1980’s. Such results suggest that few postlarval Despite the successes in hatching lobsters released ever reach commercial Several early studies focused on H. eggs and rearing larvae beyond the post- size. Barnes (1939) concluded that it americanus and H. gammarus embryol- larval stage, the United States hatcher- is often diffi cult to determine just what ogy. Herrick (1891a, b; 1895) described ies, as well as those in Quebec, France, benefi ts derive from lobster enhance- the developmental rates of H. americanus embryos at temperatures of and Norway, all were closed by 1955, ment which supplements a naturally o o with the exception of the one in Massa- fl uctuating supply of lobsters, particu- 20 –22 C and their prelarval stages. chusetts. The reasons for their closure larly when protective measures are also From these data, he was able to calcu- varied from being “biologically unsuit- in operation. Spanier (1994) suggested late the approximate date of extrusion. able” to “economically unjustifi able” that lobsters reared in the absence of (Knight, 1918; Taylor and Dow, 1958; predators and other environmental cues 7 Syslo, M. 1989–92. Massachusetts State Lob- ster Hatchery and Research Station, P.O. Box 9, Prudden, 1962; Wilder, 1965, 1971, and subsequently released into the wild Vineyard Haven, MA 02568. Personal commun., 1972; Bardach et al., 1972). Taylor might be at a higher risk of predation var. dates.

20 Marine Fisheries Review Bumpus (1891) described structures associated with the beginning and end Aiken, 1980). More recently, Macken- and illustrated the various developmen- of the embryonic metanaupliar stage. zie (1988) examined the temperature tal stages. Although these stages of The characterization of the metanau- dependence of stage duration, survival, development were not documented with pliar molt cycle and the percent-staging and body size for larval and postlarval varying temperatures, Bumpus’ data system should prove useful for future stages. Time to reach Stage IV varied describe a “primary egg-membrane” investigations of neural, physiological, from 10 to 56 days at 21°–22°C and which proved useful in later studies on and ecological aspects of Homarus 10°C, respectively. Stage IV postlar- osmoregulation by embryos (Charman- embryonic life, as well as for evolution- vae required 14 to 54 days to reach tier and Aiken, 1987). Similar descrip- ary comparisons with other decapod Stage V at 22°C and 10°C, respectively. tions were provided for H. gammarus species (Helluy and Beltz, 1991). Sim- Total cumulative survival, defi ned as by Fullarton (1896). ilarly, Charmantier and Mounet-Guil- the number of larvae reaching Stage V Templeman (1940a) presented infor- laume (1992) determined the rate of divided by initial sample size of Stage mation on the time required for H. development for embryos of H. gamma- I larvae, was 4, 56, 64, 68, and 47% at americanus to reach the 16-cell stage rus for various temperatures by measur- 10°, 12°, 15°, 18°, and 22°C, respec- up to the formation of eye pigment at ing the size of their eyes and calculating tively. Dry weight of larvae increased various temperatures. Perkins (1972) the eye index according to Perkins nearly tenfold during development from then determined the rates of develop- (1972). They discovered that the slopes Stage I to Stage V, and Stage V lob- ment, the time required to complete of equations for H. gammarus are nearly sters cultured at 15°C and 18°C were the embryonic period, and subsequent identical to those for H. americanus, larger than those cultured at other tem- hatching time at various temperatures indicating similar effects of tempera- peratures (Mackenzie, 1988). In spite for H. americanus. Hepper and Gough ture on the developmental rate of both of the different methodologies used by (1978) examined the development of species. the above researchers, stage durations embryos of H. gammarus during dif- In nature, lobster ova develop inter- were within one standard deviation ferent times of the year. However, they nally for about 1 year (Aiken and (Mackenzie, 1988). Homarus gamma- did not manipulate temperatures during Waddy, 1980; Waddy and Aiken, 1991); rus experiences similar growth variations rearing conditions. Nonetheless, their after extrusion, development can vary at corresponding temperatures (Sund, information has been useful for calcu- from 9 to 12 months depending on 1914; Havinga, 1929, cited in Scat- lating time of hatching (Burton, 1992). temperature (Bumpus, 1891; Herrick, tergood, 1949b; Richard and Wickins, Richards and Wickins (1979) used Per- 1911a; Templeman, 1940a; Aiken and 1979; Beard et al., 1985; Burton, 1992). kin’s (1972) formula to provide simi- Waddy, 1980, 1986; Waddy and Aiken, Table 5 summarizes the development (in lar information for H. gammarus, but 1991). This also holds true for H. days) for larvae to reach Stage IV and V used only one temperature regime of gammarus (Fullarton, 1896; Branford, at various temperature regimes. 13o–15oC. The data of Perkins (1972) 1978; Hepper and Gough, 1978; Burton, and Richards and Wickins (1979) have 1992). The duration of the hatching Description of been used in several lobster hatcheries period is also determined by tempera- Morphological Changes to schedule year-round larval produc- ture and can vary from a few days to sev- Morphological changes during larval tion (e.g. Schuur et al., 1976; Castell, eral weeks (Herrick, 1911a; Hughes and development and metamorphosis have 1977; Waddy and Aiken, 1984a, b; Matthiessen, 1962; Goggins and Fort- also been extensively studied. The fi rst Beard et al., 1985). ier, 1964; Ennis, 1975a). Once hatch- descriptions were made by Smith (1872) More recently, Helluy and Beltz ing begins, the larva emerges from the using larvae sampled from Vineyard (1991) examined the embryonic period egg as a “pre-larva” and usually molts Sound, Mass. and adjacent waters. and subsequent hatching time at var- into the fi rst larval stage before being Smith observed three larval stages and a ious temperatures for H. americanus released by the female (Herrick, 1911a; fourth stage which resembled the adult. from the formation of the naupliar stage Davis, 1964; Ennis, 1975a; Charman- Fourteen years later, Ryder (1886b) until the emergence of the fi rst larval tier and Aiken, 1987; Charmantier et confi rmed Smith’s observations while stage. They provided a percent-staging al., 1991). The time required for Stage rearing larval lobsters in confi nement system based upon Perkins’ (1972) eye I larvae to molt to Stage IV can vary at the Woods Hole, Mass., hatchery. index, with a quantitative characteriza- considerably, from 11 to 42 days. Water These studies were followed by the tion of 10 embryonic stages. Anatom- temperatures of 22oC result in 11 days comprehensive and beautifully illus- ical and morphological observations of development, while 8oC results in 42 trated work of Herrick (1895, 1911a). of earlier researchers (e.g. Bumpus, days. Hughes and Matthiessen (1962) His line drawings of larvae and postlar- 1891; Herrick, 1895) were then related reported similar periods of 9 days at vae have been reproduced and modifi ed to this staging system. In support of 22oC and 32 days at 14oC to reach Stage many times (by Chaikelis, 1953; Taylor, both Bumpus’ and Herrick’s claims IV. The duration of the postlarval stage 1975; Cobb, 1976; Phillips and Sastry, of embryonic molts, Helluy and Beltz also varies with temperature from 11 1980; Aiken and Waddy, 1986, 1989; (1991) presented evidence for two molts to 49 days at temperatures of 22oC and Anonymous, 1988; Charmantier et al., prior to the fi rst larval stage which were 10oC, respectively (Templeman, 1936; 1991; White, 1991). Hadley (1906a)

61(2), 1999 21 Table 5.—Economic value of hatchery-reared lobsters seeded into local waters, based on various survival rates after release. Survival rates are based on the release of 500,000 Stage IV postlarval lobsters.

1986 Fisheries Total Fisheries Total Survival Pounds Ex-vessel Value to economic economic economic economic rates harvested worth lobsterman multiplier1 value multiplier2 value

5% 25,000 × 3.14/lb = $78,500 × 2.8 = $219,800 4.5 = $353,250 10% 50,000 × 3.14/lb = $157,000 × 2.8 = $439,600 4.5 = $706,500 15% 75,000 × 3.14/lb = $235,500 × 2.8 = $659,400 4.5 = $1,059,750 20% 100,000 × 3.14/lb = $314,000 × 2.8 = $879,200 4.5 = $1,413,000 25% 125,000 × 3.14/lb = $392,500 × 2.8 = $1,099,000 4.5 = $1,766,250

1 Source: Blake (1991) for 2.8 economic multiplier. 2 Source: Mike Syslo, Massachusetts State Lobster Hatchery and Research Station, P.O. Box 9, Vineyard Haven, Mass. 02568. Personal commun. 30 Mar. 1990. also produced beautiful line drawings, Couch (1843, reported in William- used to distinguish larval stages during but these have proven to be less pop- son, 1905) was the fi rst to describe culturing experiments (Richards and ular than those of Herrick and have the prelarval stage for the European Wickins, 1979; Beard et al., 1985; Bur- only been used occasionally (Fig. 14 lobster H. gammarus. Sars (1875) ton, 1992). and by Beasley, 1904; Mead, 1910; subsequently described and illustrated Nigrelli, 1936; Barnes, 1939; Cook, the three larval stages. Williamson Larval Migrations 1972; Factor, 1995). Herrick also pro- (1905) also described and illustrated Attempts to understand the vertical vided the criteria for distinguishing the the larval and postlarval stages, and and horizontal migrations of the larvae individual larval stages which are still Chadwick (1905) described the pre-lar- resulted in a fl urry of research that began used by researchers today (Harding val, larval, and postlarval stages. More over 100 years ago. Smith (1872, 1873) et al., 1982; Matthiessen and Scherer, recently, Nichols and Lawton (1978) was apparently the fi rst to sample for 1983; Gunn, 1987; Blake, 1988, 1991, diagrammed Stage I to Stage IV lob- larvae in Vineyard Sound and the adja- 1993; NUSCO, 1989, 1990). sters; these diagrams (Fig. 15) are now cent waters. Using a hand or towing net, he observed that the planktonic stages seemed to inhabit only the surface waters. Smith’s observations were confi rmed by subsequent researchers using simple plankton sampling methods (Mead and Williams, 1903; Her- rick, 1911a; Templeman, 1937; Squires, 1970; Lund and Stewart, 1970; Scar- ratt, 1973a; Harding et al. 1979, 1982; Hudon et al. 1986). With more sophisticated sampling techniques, however, Matthiessen and Scherer (1983) found the highest concentrations of larvae at a depth of 3 m. More recently, Harding et al. (1987) found that larval lobsters were capable of vertical migrations to depths of 30 m in oceanic waters. The phototactic responses of the larvae can infl u- ence such vertical migrations. Herrick, experimenting in 1894, was apparently the fi rst to discover that the behavior of larval lobsters is strongly infl uenced by light (Herrick, 1911a). However, it was not until the extensive experiments of Hadley (1905, 1908) with H. americanus and of Böhn (1905) with H. gammarus, that the effects of light intensities on larval behavior were understood. Hadley (1905, 1908) found that phototactic responses of larvae changed both Figure 14.—Larval and postlarval American lobsters (Homarus americanus) show- within and between each stage. Böhn ing Stage I, II, and III larvae and a Stage IV postlarva. (From Hadley, 1906a). (1905) observed similar trends in H.

22 Marine Fisheries Review gammarus (reported by Herrick, 1911a). Within the ﬁ rst few hours following hatching, Stage I larvae were attracted to a light source of high intensity, but this response was reversed by the second day. However, Stage I larvae were still attracted to reduced light levels (Hadley, 1908). Harding et al. (1987) concluded that Hadley’s results anticipated not only the vertical migration due to light intensity, but also explained why most Stage I larvae remained at the surface throughout the day as found by Lund and Stew- art (1970), Scarratt (1973a), Harding et al. (1987), and Strube (1989). Stage II and III larvae are also positively phototactic several hours before molting; postmolt they become nega- tively phototactic (Hadley, 1908). This result is consistent with the reduced numbers of Stage II and III larvae found in the surface samples of Lund and Stewart (1970), Scarratt (1973a), Har- ding et al. (1987), and Strube (1989) and suggests that Stage II and III larvae migrate downward to avoid high light intensities. Figure 15.—Larval and postlarval European lobsters (Homarus gammarus) show- While phototaxis plays an important ing Stage I, II, and III larvae and a Stage IV postlarva. (From Nichols and Lawton, part in regulating vertical migrations 1978). Used with permission. and may allow the larvae to maintain or control their position, traditional thought held that larvae were merely Other factors associated with the sur- explained the large numbers of Stage passive drifters transported by surface vival, growth, and development of lobster IV postlarvae caught in surface sam- currents (Herrick, 1894; Mather, 1894; larvae also have received much atten- ples (Stasko, 1977; Greenstein et al., Templeman, 1940b; Scarratt, 1964; tion. These include, but are not limited 1983; Hudon et al., 1986; Blake, 1988). Stasko, 1978; Matthiessen and Scherer, to, light, food, salinity, disease, mutila- Between the early and late parts of Stage 1983; Hudon et al., 1986). However, tion, social environment, and water qual- IV, the phototactic response changes detection of turbulence induced by wind ity (e.g. Templeman, 1936; Cobb, 1968, dramatically and permanently, becom- or surf close to the shore may trigger 1970; Ford et al., 1976; Sastry and Pech- ing negative (Hadley, 1908; Scarratt, vertical movements (Squires, 1965, enik, 1977; Aiken, 1980; Phillips and 1973a; Botero and Atema, 1982). This 1970; Squires et al., 1971; Caddy, 1979), Sastry, 1980; Charmantier et al., 1984; change in phototaxis explains the lob- enabling the larvae to utilize subsur- Aiken and Waddy, 1986; Jackson and ster’s observed preference for dark face currents moving in different direc- Castell, 1989; Burton, 1992; Corey2). shelters beginning with the postlarval tions as a mechanism to avoid long stage and continuing into the juvenile distance displacement by surface cur- Metamorphosis and Settlement stages (Cobb, 1971; Botero and Atema, rents (Ennis, 1986a). Metamorphosis occurs at the fourth 1982; Johns and Mann, 1987; Bou- Planktonic stages also react to pres- molt (from Stage III to IV) and results dreau et al., 1990). It may also par- sure, suggesting that modulation of in a postlarvae which is different, not tially explain the adoption of a benthic depth is possible (Ennis, 1975b). Thus, only in form, but also in behavior, from lifestyle midway through or near the researchers hypothesized that move- the previous stages (Hadley, 1906b). end of the stage (Herrick, 1895, 1911a; ments of larvae were progressively The postlarva is morphologically simi- Botero and Atema, 1982; Charmantier inshore due both to predominant inshore lar to the adult, but lacks the asymmet- et al., 1984, 1991; Cobb et al., 1989a; currents during the same months that rical claws and has a longer abdomen. Corey2), when postlarvae settle into the larvae are present, and the ability of the Hadley (1908) noted that Stage IV post- benthic environment (Scarratt, 1973a; larvae to regulate their depth (Stasko, larvae were positively phototactic at Cobb et al., 1989a). 1978; Harding et al., 1982, 1987; Hard- the beginning of the stage and actively Despite the early observation that ing and Trites, 1988, 1989). sought greater intensities of light, which postlarval lobsters were capable of

61(2), 1999 23 swimming with greater speed and pre- to decline to those typically found in the fourth stage (Berrill, 1974). In con- cision than any of the preceding stages poorer habitat types (i.e. eelgrass, mud, trast, the postlarval stage of N. norvegi- (Smith, 1873; Williamson, 1905; Her- etc.)(Wilson and Steneck8) cus is assumed to fi rst enter the burrow rick, 1911a), it was not until the studies The variability in timing of settle- of an adult before forming its own shel- of Ennis (1986b), Cobb et al. (1989b), ment observed by Cobb et al. (1989a) ter (Howard, 1989). Herrick (1911a) and Rooney and Cobb (1991) that the confi rms earlier observations of Her- remarked on the burrowing behavior, signifi cance of strong and well-directed rick (1911a) that such timing is not “. . . when a bottom life is adopted, the swimming for inshore recruitment was fi xed. Furthermore, both Cobb (1968) instincts of burrowing, hiding, wari- understood. Katz et al. (1994) point out and Botero (1980) observed that Stage ness, pugnacity and preying become that currents and wind-induced trans- IV lobsters can delay molting to Stage strongly accentuated, that at this stage it port alone are insuffi cient in and of V if not presented with a suitable sub- betrays fear and caution, digs burrows themselves for offshore recruitment of strate. These laboratory observations and hides.” Herrick (1911a) concluded, larvae to coastal populations, although were confi rmed by fi eld observations of “. . . burrowing is a kind of behavior in when combined with strong directional postlarval lobsters swimming over and which the lobster frequently indulges swimming of the postlarvae, they may repeatedly diving to examine substrates from the fourth stage [late Stage IV] allow long-distance recruitment from (Cobb et al., 1983). Havinga (1929, onward throughout life. In a word, their offshore to inshore sites. cited in Scattergood, 1949b) also noted behavior is no longer variable, but is in Other studies reveal that nearshore that the swimming abilities of H. gam- measure ‘stereotyped’.” environments are not necessarily where marus postlarvae could be very useful Cobb (1971) concluded that the postlarval lobsters settle. Briggs (1975, during their search for a suitable benthic behavior involved in burrow excava- 1985, 1987, 1989, 1990, and 1991) substrate. Experiments by Boudreau et tion is not “stereotyped,” but may be found juvenile lobsters as small as 7, al. (1990) supports the hypothesis that modifi ed to suit the type of burrow 16, 17, 22, 24, and 26 cm CL in con- settling postlarvae can make an active being constructed and the type of sub- siderable numbers from traps in the benthic choice of microhabitat and will strate. However, Botero and Atema deeper waters off Long Island Sound, delay settlement if not provided with (1982) confi rmed Herrick’s conclusion New York. Blake (1991) also found suitable conditions. Delays in settle- and described the stereotypical burrow- juvenile lobsters the size of >17 mm ment are supported by higher propor- ing behavior for H. americanus. Howard CL in samples from the Connecticut tions of late molt stage postlarvae found and Bennett (1979) have described side of Long Island Sound. Therefore, in plankton samples (Cobb et al., 1989a; behavior remarkably similar for H. gam- postlarval and juvenile lobster sampling Incze and Wahle, 1991). Cobb et al. marus, as have Dybern and Höisaeter should not be limited to nearshore envi- (1989a) and Bertran et al. (1985) pro- (1965) and Rice and Chapman (1971) ronments, but should also be conducted vide some theories as to how postlarvae for N. norvegicus. in deeper waters, so that we can under- may prepare for settlement into benthic Mead and Williams (1903) noted a stand what role deeper water juvenile environments for H. americanus and marked preference for certain nooks, populations may play in the recruitment H. gammarus, respectively. The post- burrows, and other places of conceal- to the fi shery. larvae obtain information about poten- ment. Cobb (1971), Pottle and Elner It should be noted that thermoclines tial settlement sites by diving down to (1982), Lawton (1987), Boudreau et (≥ 5°C difference) may provide a bar- the substrate, touching down directly al. (1990), Wahle (1992b), Dybern rier to most settling postlarvae. It is on its surface, and lifting-off, if it is (1973), Howard (1980), and Howard not until they are much further along deemed unsuitable. By reentering the and Bennett (1979) have confi rmed in Stage IV (15 days postmolt), that water column, the lobster can use the the preference that H. americanus and they will pass through such thermo- currents to sample a wider range of H. gammarus exhibit for certain sizes clines (Corey2). Recently, Boudreau et bottom types (Cobb et al., 1989a). of shelters. Others have described the al. (1992) and Hofe (1994) have more burrowing behavior in various sub- rigorously tested the effects of thermo- Burrowing Behavior strates, such as mud, silt/clay, rocks, and clines on settling postlarvae and have Burrowing behavior has also been eelgrass (Mackay, 1926, 1929; Ennis, determined that both larvae and postlar- closely examined. Mead (1901) deter- 1968; Cobb, 1971; Berrill and Stewart, vae remain above the thermocline if it is mined that burrowing behavior fi rst 1973; Botero and Atema, 1982; Pottle of at least a 5°–6°C difference, but post- appears in Stage IV and becomes more and Elner, 1982; Barshaw and Bry- larvae will pass through a thermocline pronounced in the succeeding stages of ant-Rich, 1988). Experimental intro- as the temperature difference between H. americanus. Cobb (1971) confi rmed ductions of H. americanus along the it and the upper waters decreases (Bou- those observations. Homarus gamma- Pacifi c coast of Japan (Kittaka et al., dreau et al., 1991, 1992). This may rus also begins burrowing activity in 1983) showed that 1-year-old lobsters be one of the reasons that the greatest used burrowing methods (under cement densities of new benthic recruits are blocks) similar to those described by 8 Wilson, C., and R. Steneck. 1998. Personal found in shallow waters. With increas- commun. 14 April and 14 June. Univ. Maine, Cobb (1971), Dybern (1973), and ing depth, densities of new recruits tend Darling Mar. Lab., Walpole, ME 04573. Cooper and Uzmann (1980).

24 Marine Fisheries Review Substrate Selection individuals (< 40 mm CL) being located in quadrats with 100% cobble cover. in the upper portion of large sod clumps These results are similar to laboratory Interest in the appropriate substrates and in smaller clumps of shallow slope experiments of Van Olst et al. (1976a), for settling postlarvae has also spurred areas. Larger lobsters (> 40 mm CL) where they observed densities of 6–30 much research. The stereotypical and occupied large rocks or were found under lobsters/m2 ranging in size from 14 effi cient burrowing behavior exhibited boulders in deeper waters. More recently, to 18 mm CL, depending on the sub- not only by H. americanus, but also Able et al. (1988) found postlarval lob- strate. Conversely, densities up to 62/m2 H. gammarus and N. norvegicus, sug- sters in densities of 2.1 individuals/m2, were observed on rocky bottoms in gested that lobsters were particularly ranging in size from 6 to 72 mm CL semienclosed basin experiments carried suited to soft substrates (Berrill and (mean of 26.7 mm CL), in peat beds out with H. gammarus (Bertran, 1984). Stewart, 1973; Berrill, 1974; Botero, near salt marshes of Cape Cod. While Wahle and Steneck (1991) report 1980). As early as 1895, Herrick con- While densities vary, substrates other that cobble may appear to be a preferred sidered that eelgrass may be a poten- than peat or eelgrass provide shelter for habitat for postlarval lobsters, others tial habitat for lobsters. MacKay (1920, recently settled lobsters. A. M. Olsen (MacKay, 1920, 1929; Morrissey, 1966; 1929) confi rmed this when he found (Senior Research Offi cer, DFO, CSIRO, Cooper and Uzmann, 1977; Able et al., small lobsters in mixtures of sand, mud, Australia) captured 32 lobsters (of about 1988) have shown that postlarval lob- and eelgrass. Barshaw and Bryant-Rich 17–69 mm CL) by hand in 35 minutes sters use various substrates; thus, eel- (1988) conducted a long-term study on off Richibucto, New Brunswick, during grass, peatbeds, or mud can and will be the behavior and survival of early juve- a visit (reported in Wilder, 1959). He used as alternate habitat where cobble nile lobsters in three naturalistic sub- observed that lobsters were not only may be lacking. strates: eelgrass, mud, and rocks. They numerous on rocky bottoms, but also Surprisingly, lobsters have also been found that postlarval lobsters took less on smooth, fi rm sandy/silt. Some were found in the intertidal zone. Krouse time to burrow into eelgrass and had partially or completely hidden under (1983), during a tag-recapture program higher rates of survival compared to large fl at stones. Many, however, were conducted from 1977 through 1982, postlarval lobsters in mud and rocks. seen moving freely about (reported by caught lobsters ranging in sizes from Furthermore, lobsters in eelgrass were Wilder, 1959). National Marine Fish- <10 to ~80 mm CL, by hand in the larger, despite the higher densities of eries Service (NMFS) divers examined intertidal zone of Maine’s Sheepscot animals. However, predation experi- mud substrates in central Maine harbors River. Substrate there ranged from fi ne ments using eelgrass (Barshaw and during July through September 1975 to coarse sand intermingled with broken Lavalli, 1988) suggest that eelgrass is and found densities of Stage IV to XI shell with scattered rocks of various a suboptimal environment for predator lobsters of 1–20/m2. Nearly all indi- sizes (from barely moveable to many avoidance but does sustain intermedi- viduals were retrieved from tiny bur- of grapefruit size). This substrate is ate levels of survival when compared to rows excavated next to solid objects found adjacent to large bedrock ledges that of sand environments. Heck et al. (e.g. lumber, discarded shoes, bottles) covered with seaweed (Krouse9). Simi- (1989) conducted trawl sampling from that are the type of refuse commonly larly, Cowan10 and Cowan (1999) found eelgrass areas on Cape Cod, Mass., and discarded into waters of an intensively small-to-moderate densities (0–8.6 indi- found only low densities of lobsters. utilized harbor. Very few lobsters were viduals/m2) of intertidal lobsters rang- Their study concluded that eelgrass found in mud substrates which lacked ing in size from 3 to 42 mm CL, at meadows were not signifi cant nursery overlying rocks, gravel, or other solid Lowell’s Cove, Orr’s Island, Maine, areas; however, their sampling method objects (Cooper and Uzmann, 1977). and in selected rocky areas throughout differed greatly from the more success- Ennis (1968) found that Stage VI the New England states (N.H., Mass., ful air-lift sampling technique of Able juveniles were capable of building R.I., Conn.). These lobsters are typi- et al. (1988) and Wahle and Steneck depressions in sand and mud after a cally found under scattered rocks on the (1991). Peat reefs are also intermediate period of time spent wandering over beach surface. in their protective quality (Barshaw et these substrates, but ultimately pre- Despite fi nding lobsters in vegetated, al., 1994); nonetheless, relatively high ferred rocks and would take shelter rock, and intertidal habitats, Wahle and densities of juvenile lobsters have been immediately upon contact with them or Steneck (1991) suggest that soft sub- found in peat. Morrissey (1966) discov- would shift to sheltering under rocks if strates of mud or those with vegetation, ered postlarval and juvenile lobsters in they were subsequently introduced onto such as eelgrass and peat, are rarely salt marsh areas and around sod clumps the mud and sand substrates. Hudon used by settling postlarvae. Support for in the Nauset Harbortown Cove area of and Lamarche (1989) found no postlar- Orleans, Mass. Approximately 135 m of val lobsters of 5–31 mm (CL) on sand 9 Krouse, J. S. 1992. Department of Marine Re- sources, Mar. Resour. Lab., McKown Point, West shoreline were sampled in each instance. or on sand and eelgrass, but reported Boothbay Harbor, ME 04575. Personal commun. On the three sampling dates, lobsters densities of 1.4/m2 in bare rocks and 6 July. captured ranged in size from 7 to 83 mm 1.3/m2 in rocks with algae. Wahle and 10 Cowan, D. F. 1992–96. Department of Biology, Bates College, Lewiston, ME 04240, and The CL. Morrissey (1966) also observed a Steneck (1991) observed 5–40 mm CL Lobster Conservancy, P.O. Box 193, Orr’s Island, vertical gradation by size, with smaller lobsters at maximum densities of 16/m2 ME 04066. Personal commun., var. dates.

61(2), 1999 25 their hypothesis comes from their exten- or rocks upon rocks (Lavalli and Bar- She also confi rmed the conclusions of sive sampling efforts in cobble envi- shaw, 1986; Johns and Mann, 1987; Bar- Hargrave et al. (1985) that as larvae ronments along the coasts of Maine, shaw and Lavalli, 1988; Wahle, 1991b; progressed through each stage, their New Hampshire, and northern Massa- Wahle and Steneck, 1991), as well as prey size increased. Similarly, Juinio chusetts (Gulf of Maine). Densities of small stones embedded in sand and boul- and Cobb (1992) found that copepods small lobsters (<40 mm CL) found by ders with macroalgae (Hudon, 1987; and crab larvae were the most common them are some of the highest reported; Hudon and Lamarche, 1989). These items in planktonic, postlarval lobster however, sampling effi ciency of the substrates either require minimal bur- stomachs, with fi sh eggs and insects also air-lift suction process they use has rowing activities or allow lobsters to frequently found. Based on gut fullness not been established for the varying probe spaces between rocks and immedi- and condition of gut contents, Juinio and environments (mud, cobble, eelgrass). ately occupy appropriately sized spaces Cobb (1992) concluded that postlarvae Wahle and Steneck (1991) sampled to (Wilder, 1957; Wahle, 1992b). However, fed throughout the day intermittently a depth of 15 cm, while Cooper and Roach (1983) experimenting with caged and at all stages of the molt cycle. Fur- Uzmann (1977, 1980) observed post- environments in the fi eld found that sur- thermore, by comparing growth rates larval and juvenile lobster burrows as vival and growth was higher in mud sub- of wild and laboratory-reared lobsters deep as 70 cm. strates, followed by vegetation and then via RNA:DNA ratios, Juinio and Cobb Able et al. (1988) observed that larger rock. Unfortunately, the lack of consis- (1994) confi rmed Wilder’s (1953) ear- lobsters (>40 mm CL) build burrows to tent trends (such as reduced predation lier observation that laboratory-reared a 99 cm depth. Howard (1989) reported and high survival and growth) from these larvae grow more slowly than their wild that N. norvegicus constructed burrows studies has made an accurate portrayal counterparts. 20–30 cm below the mud surface. Wahle of benthic recruitment for postlarval lob- However, there is a caveat to these and Steneck (1991) relied on the lab- sters diffi cult. results in that growth rates of labora- oratory studies of Berrill and Stewart tory-reared larvae will be highly depen- (1973) and on their own observations Feeding Activity dent on the type of brine shrimp used for determining their sampling depth. Research has also targeted the feed- during culture, and different brands of However, laboratory experiments on ing activities of larval and postlarval brine shrimp can vary tremendously in burrowing behavior in 8–12 cm sub- lobsters. Williams (1907) examined the lipid content and overall quality (Fujita strates found burrows to be 9 cm deep stomach contents of 100 larvae and post- et al., 1980; Eagles et al., 1984). Fur- (Berrill and Stewart, 1973). Lobsters larvae taken from the hatching bags at thermore, the water supply can also pro- may have been restricted from digging Wickford, R.I., and noted that they con- vide additional food items (see below), further by the bottom of the tank. With sisted primarily of copepods and dia- if it is not fi ltered. Thus, the culture substrates 8 cm deep, lobsters burrowed toms. Herrick (1911a) found that the techniques of Juinio and Cobb (1994) to a depth of 2.6 cm (Botero and Atema, stomachs of larval lobsters examined must be considered when comparing 1982). In contrast, Howard and Bennett from laboratory aquaria and Vineyard wild and laboratory-reared larvae. (1979), experimenting with H. gamma- Sound, Mass., contained diatoms, crus- Survival studies indicate that settled rus, used substrates 40 cm deep and taceans, bacteria, algae, and amorphous postlarval and small juvenile lobsters found that postlarval lobsters regularly matter. Templeman (1933) reported that are also capable of surviving and grow- burrowed to a depth of 15 cm, with a larvae reared on a full ration of copepods ing on planktonic diets. Emmel (1908) single instance of an 8 mm CL lobster exhibited higher survival rates than those was the fi rst to show this ability when burrowing to a depth of 25 cm. fed half, quarter, or one-eighth rations conducting feeding experiments with Considering the stereotypical bur- of copepods. Harding et al. (1983) dis- artifi cial diets (e.g. beef, clam, lobster rowing behavior and ecological simi- covered that Stage I and II larvae, taken muscle, shredded fi sh, and beef liver) larities of these three species, the actual from St. George’s Bay, Nova Scotia, for hatchery operations. Postlarval lob- depth to which the postlarvae can consumed cladoceran podons and cope- sters on artifi cial diets were compared burrow is quite variable. The differ- pods, while Stage III and IV lobsters to those fed nothing, getting only that ences in depth and sampling techniques consumed copepods, gastropod larvae, which entered with the water supply. used by Hudon (1987) and Wahle and and crab larvae. Hargrave et al. (1985) Instead of starving, Stage IV postlar- Steneck (1991) could explain the lim- found that the larval lobster’s natural vae were able to molt into Stage V, ited numbers of lobsters found in mud diet is composed of organisms in the albeit at a slower rate than those on a and eelgrass, respectively. Therefore, 210–610 µm range, and that prey size feeding regime. Barshaw (1989) also until sampling techniques are tested for increases as the larvae grow, with late showed that postlarval lobsters could their effi ciency in each substrate type, stage larvae preferentially consuming survive well on planktonic diets, but extrapolation of laboratory results for more crab zoea and megalops larvae. exhibited molt delays after Stage V guidance in sampling protocols should Gunn (1987) noted that larvae from when compared to those fed artifi cial be used with caution. Long Island Sound consumed mostly diets. However, her data were compli- Environments which provide the best crab larvae and copepods, with occa- cated by unequal amounts of food fed protection from predators include cobble, sional cladocerans, insects, and diatoms. for the two groups of lobsters.

26 Marine Fisheries Review Lavalli (1991) repeated and extended did provide several theories on the di- sters (e.g. telson, dorsal side of the Barshaw’s experiments, fi nding that rection and extent of lobster move- third segment of the large claw, between postlarvae were capable of surviving ments. Thereafter, many tagging studies the carapace and abdomen in the body throughout their fi rst season on diets of focused on tag retention. Templeman cavity, and in the elbow of the large claw plankton in the size range of 95–1,000 (1935, 1940c) clipped metal tags to the (Anonymous, 1965)). Results indicated µm; however, these lobsters were inca- telson of sublegal and legal-sized lob- that these tags could be retained through pable of surviving on diets consisting sters. Plastic or metal discs attached by only one molt (Anonymous, 1966), but mostly of diatoms. Lavalli (1992) deter- wire to the second segment of the che- the “sphyrion” tag, a modifi ed spa- mined with videoanalysis that juvenile liped were used by Wilder (1947, 1953) ghetti- dart tag, remained through molt- lobsters were capable of suspension in Canada, Thomas (1955) in Scotland, ing. This tag consisted of an anchor feeding. More recent studies have and Simpson (1961) in Wales. Wilder made of stainless steel wire with a shown that adult European lobsters can (1954) used a metal strip bent to form a double strand of polyethylene mono- also benefi t from suspension feeding hook at one end. This hook was inserted fi lament attached to a numbered disc on plankton (Loo et al., 1993). There- into the posterior margin of the car- (Scarratt and Elson, 1965). fore, it is not surprising that Japanese apace and kept in position along the The “spaghetti” or “sphyrion” tag researchers have successfully reared mid-line by an elastic band attached to is essentially an external tag, that is homarid larvae by utilizing a mixture of the other end and passed over the ros- anchored internally either between the phyto- and zooplankton cultures previ- trum. These tags were quite successful, carapace and abdomen or elbow of the ously developed for culture of penaeid but were not retained through succes- claw. By implanting the tag between the shrimp (Kittaka, 1990). Development sive molts. Another method developed carapace and abdomen, a higher reten- of the larvae to Stage IV at 20°C in by Appelløf (1909b) and employed by tion rate can be expected because this is outdoor tanks takes about 2 weeks. At Dannevig (1936), Wilder (1953, 1963), where the lobster exits during molting. a stocking density of 10 individuals/ and Wilder and Murray (1956) was to Improvements in these tags continued liter, the survival rate was 50% (Kit- punch or drill holes in the tail fan. to be made and, after the successful lab- taka, 1990). This method was not only successful oratory experiments made by Hughes but lasted through several molts until (reported by Anonymous, 1965, 1966), Tagging and Movement the fl esh grew back and fi lled the hole, Scarratt (1970) and Cooper (1970) inde- Examination of the movements of but it was not completely reliable and pendently began using a vinyl tubing adult lobsters and the development of depended on the grow rate of the lob- sphyrion tag instead of the numbered tagging methods also began in hatch- ster; Simpson (1963) provides a review. disc. With such a modifi cation, Cooper ery settings. Bumpus (1901a) tagged While these external tags provided some (1970) reported an 88% retention rate 497 female lobsters (with eggs recently information to lobster managers, they for lobsters which had molted once, and removed) with copper tags attached to were incapable of providing long-term the successful fi eld tests of these tags their rostrums. These females were lib- movement information, as well as infor- (Cooper, 1970; Scarratt, 1970) spurred erated at various points along the coast mation on growth rates. further, large-scale tagging studies in near Woods Hole, Mass. Within 4–89 In 1963, John Hughes, Director of the United States and Canada; Krouse days, 76 of the females were recaptured, the Massachusetts State Lobster Hatch- (1980a, 1980b), Stasko (1980), Miller some from as far as 25.6 km away. ery and Research Station, experimented et al. (1989) give reviews. Migratory females moved in a south- with several prototype tags. The “spa- While it is impractical to review all westerly direction, while nonmigratory ghetti-dart” was made of various col- tagging studies, a few will be summa- lobsters remained in the local waters for ored vinyl plastic tubing (about 2 mm rized to show how the resulting data is several weeks. Homing behavior was in diameter) attached to a small fl ex- applied to lobster management. Cooper observed as several lobsters returned ible double-barbed nylon dart. Only the and Uzmann (1971) and Uzmann et to the place where they were captured dart and a small portion of the tubing al. (1977) have documented the sea- prior to tagging and release. Mead and was inserted into the lobster, with about sonal migrations of larger, deep-water Williams (1903) tagged 112 adult lob- 62.5 mm remaining exposed (Anony- lobsters from the continental slope and sters in 1902, using the same technique mous, 1965). The multi-barbed tag with canyons to the shallow waters in the as Bumpus. They liberated these lob- a dart-like spear was about 25 mm by summer on Georges Bank. Uzmann et sters at Wickford, R.I., and within 1–11 6.4 mm with a 1.6 mm diameter, 75 mm al. (1977) hypothesized that lobsters days, 16 were recaptured. An additional long plastic “spaghetti” tube attached. maintain an 8o–14oC temperature regime 385 adults were tagged and liberated in Then a larger tag was tried (30 mm long during their onshore migration in order 1903, and within 1–59 days, 62 were by 8 mm wide) with a multibarbed dart to maintain growth, molting, and egg recaptured. Southerly movements of having an eight-pronged barb (Anony- extrusion. Recently, this hypothesis has up to 17.6 km were recorded (Barnes, mous, 1965). been confi rmed by Chandler (1991) who 1906b). Several tagging studies used these conducted a series of in situ studies on Although these tagging experiments tags in various locations along the outer Cape Cod, Mass., and recorded were exploratory and inconclusive, they bodies of sublegal and legal-sized lob- lobster size, sex, temperature range, and

61(2), 1999 27 behavior. From these data, she was able apparently the fi rst to experiment with marus (Sheehy et al., 1996). The size and to estimate population structure. Other this type of tag, but their tag was made of number of lipofuscin granules and the studies undertaken by Cooper (1970), 1.6 mm clear plastic tubing with a small carapace length are signifi cantly related Lund et al. (1973), and Ennis (1984) wire insert. This was injected inter- to the age of the animal, such that it reported inshore movements to be min- nally into the lobster with a hypodermic may be possible to determine the age imal. Similar results have been reported needle. Ennis (1972) successfully used of any lobster sometime in the future. for H. gammarus and N. norvegicus internal tags for growth-per-molt studies Tagging experiments with smaller lob- (Bennett et al., 1978; Chapman, 1980). of larger lobsters. More recently, a tag sters are important to determine the link- Besides the information on move- 0.25 mm in diameter and 1 mm long has age between the numbers of postlarval ments and behavior gained from such become popular for use with lobsters as benthic recruits with those eventually tagging studies, data in the form of small as 9 mm CL (Wickins and Beard, recruiting into the fi shery. growth, natural mortality, and fi shing 1984; Wickins et al., 1986; Cowan, Thus, lobster hatcheries, whether mortality has also been collected 1999). Consequently, many enhance- state-run or for experimental purposes (Thomas, 1973; Smith, 1977; Krouse, ment programs for the lobster H. gam- only, have contributed to not only the 1981; Briggs and Mushacke, 1984; marus now use the “micro-tag” to “seeding” of larvae or postlarvae into Miller et al., 1989) and used for lobster evaluate the survival rates of hatchery- coastal waters of New England states management (Northeast Marine Fisher- produced lobsters and compare these and European coasts, but also to much ies Board, 1978). The sphyrion tag has rates to wild stocks (Bannister et al., experimentation and description of the enabled researchers to gather signifi cant 1989; Cook et al., 1989; van der Meeren species’ natural history. Many of the life history data on the lobster in its nat- et al., 1990; Burton, 1992; Beard and early studies have had direct impacts on ural environment; however, these data Wickins, 1992; and Resource Enhance- recent and current projects, providing are limited to large juvenile and adult ment section). not only the background for newer and lobsters (>60 mm CL) (Krouse and Nut- An even smaller tag (0.25 mm diam- supposedly more sophisticated work, ting, 1990a), which leaves gaps in infor- eter by 0.5 mm long) has allowed the but also the culturing techniques (scaled mation for smaller juveniles. Larger tagging of Stage V–VI postlarval lob- down) which supply the larvae and lobsters traditionally have been used sters (Burton, 1991, 1992). In the United postlarvae necessary for this work. In for several reasons: 1) early research- States, Krouse and Nutting (1990a) suc- nearly every case, the work of the past ers were more interested in older life cessfully tagged H. americanus lobsters has been confi rmed, rather than refuted. stages, 2) until recently it was diffi cult 12–24 mm CL with micro-tags 0.25 by l The articles published in various state to collect lobsters smaller than 40 mm mm long. Their fi eld and laboratory tests fi sh commission bulletins have been CL, and 3) the sphyrion tag was thought suggest that this tag is useful for the mon- cited and referred to for decades and are to be unsuitable for smaller lobsters itoring of movements, growth, and pos- still being used. Herrick’s (1895, 1911) (Krouse and Nutting, 1990b). sibly mortality rates. Others have been monographs, although sadly out of date, Because the sphyrion tag was con- experimenting with yet another type of remain invaluable references for lobster sidered inappropriate for smaller lob- tag called the polyethylene streamer tag. biologists. An update to Herrick has sters, researchers began experimenting This tag was originally developed for been recently published, and contains with lobsters < 60 mm CL to develop shrimp, but has also been successfully several chapters devoted to the develop- a satisfactory tagging technique. Bern- applied to lobsters (Landsburg, 1991). ment and regulation of the fi shery and stein and Campbell (1983) developed a The anchoring method of this streamer aquaculture (Factor, 1995). Through- miniaturized persistent back tag for lob- tag makes it possible to tag larval and out all of these literature sources, the sters 20–25 mm CL. Krouse and Nut- postlarval lobsters (Landsburg11). Fin- successes of the former and current ting (1990b) modifi ed the western rock ally, a rather unusual method for tag- hatcheries have kept alive the hope that lobster tag and reported retention rates ging lobsters actually records the number lobsters might be someday commer- of 100% on 30 lobsters of 25–39 mm of molts. This method involves epider- cially farmed. CL. They recommended that this tag be mal implants inserted into the abdominal used in movement and growth studies haemcoel of the lobster. Each time the Aquaculture Potential requiring observations over prolonged lobster molts, a layer of cuticle is formed. Farming of the homarid lobster has periods of time. By counting these layers, the molt history been classifi ed into several forms of Another extremely successful method can be ascertained (Shelton and Chap- “aquaculture:” Resource enhancement, for tagging small lobsters was the man, 1987, 1995). Lipofuscin granule product enhancement, full grow-out Bergman-Jefferts “microtag.” Originally content in the brain also holds a poten- (Waddy, 1988; Edwards, 1989), and developed and used by Jefferets et al. tial for aging cohorts of both Homarus transplantation programs (Van Olst et (1963) for marking salmonids in en- americanus (Wahle et al., 1996) and its al., 1980). More recently, another form hancement experiments in the United European counterpart, Homarus gam- has been suggested—that of a soft- States, the “microtag” was improved shelled product (Wear, 1990). 11 Landsburg, W. 1991. Science Branch, P.O. Box for use in lobsters. Saila and Flowers 5030, Moncton, New Brunswick, Canada E1C Aquaculture involves manipulation (reported in Anonymous, 1965) were 9B6. Personal commun., 3 Nov. of the natural population by culturing

28 Marine Fisheries Review (Van Olst et al., 1980), which is depen- odology for lobster culture (Portersfi eld, rare alleles, the offspring will bear a dent on the ability to control and eco- 1982; Mead, 1989; Montauk Marine Sci- natural tag—the numbers of progeny nomically optimize all aspects of the ence Institute, 1993). In addition to this produced with these rare alleles will be biology and environment of the species “Blue Lobster Project,” which is still in greater than that expected in a natural cultured (Shleser, 1973). We present a operation, D’Agostino was also respon- population. This type of tag could also review of these forms of aquaculture sible for releasing millions of Stage IV be used to monitor the survival rates and the developments that have fol- lobsters into Fort Pond Bay, Montauk, and dispersion of hatchery-released lob- lowed from their operations. N.Y., during 1974–79. Though not yet sters (Pressick, 1974; Hedgecock et al., reported in the literature, Goldfi nger12 1975; Tracy et al., 1975; Hand et al., Resource Enhancement states that the effi cacy of this program 1977; Hedgecock, 1977). Resource enhancement began over is well-documented, but in unpublished 100 years ago because of unenforced data, and, as reported by the local fi sh- Hybrids laws, increased demand, increased fi shermen, an increase in lobster numbers is Mating H. americanus and H. gam- ing effort, and declining lobster landings. evident. Goldfi nger12 asserts that “there marus produces hybrids (Carlberg et As such, the homarid lobster received is no question that seeding areas that al., 1978; Kittaka, 1984b) which pos- much attention from researchers on both have natural lobster habitat with Stage sess the subrostral spine of H. gamma- sides of the Atlantic Ocean, and resource IV or V lobsters, will have a positive rus. They are therefore distinguishable enhancement has been practiced contin- effect on future stock.” At the Darling from H. americanus (Wickins, 1983). uously ever since. Marine Center in Walpole, Maine, Sam However, this method may not be eco- The objective of these enhancement Chapman proposed a similar program logically sound, as the release of nonin- programs was to alleviate the high mor- to hatch, raise, feed, and release “blue” digenous species can have major effects tality associated with the pelagic stages lobsters off the Maine coastline. His pro- on the natural communities into which in nature by releasing larval, postlarval, gram was initially funded (Plante, 1989) they are released (Syslo, 1986). or juvenile lobsters. Several countries are and had the long-term goal of evaluat- currently engaged in this form of aqua- ing hatchery release programs (Plante, Micro-tags culture, whether it be called “resource 1989; Irvine et al.5; see later section on The micro-tag designed after Jef- enhancement,” “stock enhancement,” or culture work in Maine) and following ferets et al. (1963) has been used to “seeding.” Until recently, it was not clear year-to-year survivability in the wild, create a binary coded-wire tag in sev- if stock enhancement via the release of but it has since been terminated. eral sizes. Researchers have success- hatchery-reared lobsters provided any fully implanted tags of either l mm × real benefi t to the preservation of the spe- Permanent Genetic Tags 0.25 mm or 0.5 mm × 0.25 mm into the cies. Lack of tagging methods to distin- Genetic studies (Pressick, 1974; Hedge- base of the fi fth walking leg of the lob- guish hatchery-reared stock from those cock et al., 1975; Tracy et al., 1975; Hedge - ster and have documented a 90% reten- settling naturally prevented documenta- cock, 1977) revealed that there is one tion rate (Wickins and Beard, 1984; tion of hatchery-reared lobster survival. variable locus, phosphoglucomutose Wickins et al., 1986; Krouse and Nut- However, several methods have been (PMG), in lobsters that could be used as ting; 1990a). This method is success- recently identifi ed to distinguish hatch- a marker for distinguishing introduced fully used by several stock enhancement ery-reared stock from those of the wild. lobsters from wild stock. An example programs (see sections on Great Bri- These include the use of color morphs, of how this can be accomplished has tian; Sea Fish Industry Authority, Scot- genetic tags, hybrids, and micro-tags. been reported by Pressick (1974) and land; North Western and North Wales is explained as follows: lobsters from Sea Fisheries Committee; Norway; and Color Morphs Woods Hole and Martha’s Vineyard France). John Hughes (1968a), of the Mas- possess a slow moving allele (PMG100) Only two of the above methods are sachusetts State Lobster Hatchery, pro- which is fi xed, as there are no hetero- currently in use: micro-tags and color posed using color morphs as natural tags zygotes detected thus far in this popu- morphs. These methods will be pre- to determine the survival and impact lation. In Maine, the lobster population sented with up-to-date information in the hatchery-reared lobsters had on the fi sh- has both a slow (PMG100) and a fast sections covering the individual stock ery. Later Anthony D’Agostino, of the (PMG103) moving allele. The frequency enhancement programs using them. New York Ocean Science Laboratory, of PMG100 is around 0.16, which indi- incorporated Hughes’ idea in his brood- cates that a homozygote for the rare Massachusetts stock maintenance and development allele could only be expected in about Until 1997, the Massachusetts State program using “blue” color morphs 3% of the individuals in Maine. By Lobster Hatchery and Research Station, (Rattner, 1986; Irvine et al.5). Initially, selecting and mating lobsters with the Vineyard Haven, Martha’s Vineyard, the project was designed to develop a maintained the longest continuously hatchery program for restocking New running stock enhancement program in 12 Goldfi nger, I. N. 1990. Marine Science Insti- York’s inshore wild lobster populations tute at Montauk, P.O. Box 820, Montauk, NY the world. Since 1949 this hatchery and to advance the technology and meth- 11954. Personal commun., 24 July and 20 Aug. contributed substantially to the devel-

61(2), 1999 29 opment of hatching techniques (Hughes spent entirely rearing and stocking, no bleach (Bullis and Syslo, 1996; Cogger and Matthiessen, 1962, 1967; Hughes, new publications were written on diet and Bayer, 1996). Both of these tests 1968a, b; Hughes et al., 1972; Hughes, or growth studies (Syslo7). However, indicate that lobsters exposed to chlo- 1972, 1973; Hughes et al., 1974; Schuur experiments using various types of sea- rine bleach can be detected up to 12 et al., 1976; Van Olst et al., 1980; weeds have been published by Syslo and days post-dipping. Syslo and Hughes, 1981) which are still Hughes (1981) and have demonstrated After a visit to the hatchery in Mas- applied to hatchery programs world- that eelgrass, Zostera marina; Japa- sachusetts, Harvard Medical School wide. While the hatchery released nese weed, Codium fragile; Irish moss, researchers studying neurological dis- approximately 500,000 Stage IV lob- Chondrus crispus; rockweed, Fucus orders decided to start rearing their own sters annually into the coastal waters of spiralis; and kelp, Laminaria sp., can larvae (Syslo, 1986). The Research Sta- Massachusetts, its reseeding effort was be used as a diet substitute for fi sh and tion aided their efforts by supplying terminated in January 1997 in order may provide a less expensive nutritional ovigerous females at the New England to allow the research station to focus supplementary diet which contributes Aquarium in Boston. The Research Sta- more directly on its research goals. The to growth (Syslo and Hughes, 1981). tion also assisted the Japanese Govern- station will conduct research on the Experiments using color morphs have ment by providing egg-bearing female incidence of multiple egg batch extru- also been successful, particularly those lobsters for their research on the feasibil- sion that occurs without mating, gather of breeding; however, the rearing of ity of introducing the American lobster, growth rates, and determine how much larvae to a size of 40–50 mm CL has H. americanus, into their coastal waters egg production can vary in the wild. proven less successful since mortali- in the hopes of stimulating a new fi sh- Management then can use the informa- ties are higher than those with wild- ery (Syslo, 1986; Kittaka, 1990; Syslo7). tion gathered to assess the health of the type coloration. Based on the lower sur- The Environmental Research Labora- lobster stock (Estrella6) vivorship, color morphs may not pro- tory of the United States Environmental Prior to its closure, the hatchery vide reliable indicators of the survival Protection Agency (EPA) in Narragan- operations began with egg-bearing lob- rates of hatchery-stocked lobsters in the sett, R.I., established a lobster rearing sters obtained from offshore fi shermen wild. Furthermore, since the brighter program in 1990 to hatch and rear larval by special permit. These females were color morphs themselves may invite lobsters employing similar techniques to placed into large fi berglass tanks, more predation, this part of the research those used at the Massachusetts hatch- approximately 274 cm long × 91.5 cm was discontinued several years ago ery. In 1992 they had over 5,000 lobsters wide × 30 cm high. During mid-May, (Syslo7). in culture which were used solely for when the water temperature rose to The illegal removal of eggs from ber- environmental testing purposes (John- about 15oC, larvae hatched and were ried (egg-bearing, or ovigerous) lob- son13); however, they too have since ter- caught in a plastic screened box as the sters was a long-standing problem minated their culture program. water drained from the tank through dating back to the early 1800’s (Her- an overfl ow pipe. The larvae were then rick, 1895). Several methods used for Maine transferred to rearing tanks (kreisels) at the removal of eggs were described Bayer (1982) submitted a feasibility stocking densities of about 3,000 Stage I by Templeman (1940d) and Hughes study for a hatchery-release program to larvae per kreisel. They were fed frozen (1965) and included the use of a stiff the Department of Marine Resources, adult brine shrimp, Artemia salina, and brush, compressed air, and a high pres- and in 1986 fi ve lobster hatcheries were were reared for about 1 month until they sure hose. Hughes (1965) working at established at Cutler, Walpole, Stoning- molted into Stage IV. Once the Stage the Massachusetts State Lobster Hatch- ton, Five Islands, and Bar Harbor, Maine. IV animals were removed for stocking, ery developed a method for detecting The fi rst two hatcheries were funded the kreisels were again stocked with scrubbed lobsters based on physiologi- primarily by the lobster industry, but Stage I larvae. Postlarvae were trans- cal and biological characteristics. This the towns themselves, local fi shermen, ported to an appropriate coastal area method was then enhanced by the use of and the University of Maine all contrib- and were released by snorklers at the biological stains (Karsson and Sisson, uted funds, equipment, and time to these surface over a bottom that provided the 1973). These techniques were useful for programs. Stonington and Five Islands maximum amount of shelter (Hughes detecting the removal of eggs with the hatcheries were funded and operated by and Matthiessen, 1962, 1967; Syslo7). naked eye (Morejon, 1975). However, the local fi shermen on a volunteer basis Besides stock enhancement, exper- recent evidence suggested that fi sher- (Plante, 1986). The Bar Harbor hatchery iments on growth rates of juvenile men were now dipping ovigerous lob- is funded and operated by the Bar Harbor lobsters, diets, color morphs, and poly- sters in a chlorine bleach solution which and Southwest Bar Harbor Oceanariums culture techniques were conducted. dissolved all of the residual cement so and tourist admissions (Schreiber, 1998). Animals and technical advice were that no obvious external signs could Although Five Islands ceased operation also provided to researchers (Syslo7), be seen by the naked eye. As a result, which contributed signifi cantly to many researchers from Massachusetts and 13 Johnson, M. W. 1991. Environmental Protec- tion Agency, Environmental Research Labora- experiments. Because the growth rate Maine developed tests to determine if tory, 22 Tarzwell Drive, Narragansett, RI 02882. work was so voluminous and time was lobsters have been subjected to chlorine Personal commun., 17 May.

30 Marine Fisheries Review Table 6.—Cost to produce Stage IV postlarval lobsters. in 1988, three of the remaining hatch- ous water circulation to minimize larval (adapted from Fitzhenry et al., 1986.) eries released Stage IV postlarvae until interactions which could otherwise lead Cost1 per Cost1 per 1991 in the following numbers: 544,681 to cannibalism. The entire volume of Individual individual (Cutler); about 20,000–30,000 (Stoning- seawater was replaced every 48 hours by Percent stage IV Percent stage IV ton); and between 93,000 and 95,000 water pumped from the harbor, ensur- survivorship lobster survivorship lobster Stage IV and V lobsters, including 8,000 ing a high level of fresh seawater at all 100 2.1 40 5.2 90 2.3 30 6.9 “blue” color morph lobsters (Walpole) times (Fitzhenry et al., 1989). 80 2.6 20 10.4 14 70 3.0 10 20.8 (Chapman ). Once the larvae hatched, they were 60 3.5 5 41.7 Operations at Cutler and Walpole immediately collected with a small 50 4.2 consisted of similar procedures for pro- aquarium net, weighed (for counting 1 Cost in cents. duction of Stage IV postlarval lobsters purposes), and added to a food-enriched (Plante, 1986), so, with this in mind, conical tank (Fitzhenry et al., 1988). Table 7.—Number of Stage IV postlarvae produced from only Cutler will be reviewed. Based To minimize larval interactions, initial egg-bearing females; adapted from Fitzhenry et al. (1986) on Beal’s (1988) work at the Univer- stocking density was low, at about 40 with estimated number of stage IV postlarvae at 80% sity of Maine at Machias, the variables individuals/liter. Additionally, a high survival rate. infl uencing larval survival were iden- rate of aeration and a high concentration Estimated number of: tifi ed. These included food type, aera- of brine shrimp, Artemia salina (about Carapace Eggs Egg-bearing Total Stage IV tion rate, and initial stocking density, 600–1,000 per larvae), was provided length (mm) per female females eggs lobsters and the methodology used at Cutler to prevent cannibalism (Beal, 1988; 80 6,704 10 67,040 53,632 85 8,119 8 64,952 51,962 took them into account (Fitzhenry et Fitzhenry et al., 1989). Unlike most 90 9,723 7 68,061 54,449 al., 1989). Broodstock (berried females) hatchery operations, Cutler fed both 95 11,534 6 69,204 55,363 100 13,561 6 81,366 65,093 were obtained from the local waters algae and brine shrimp to larval lob- 105 15,819 4 63,276 50,621 off Cutler, Maine, by specially licensed sters. This method was fi rst employed 110 18,322 4 73,288 58,630 115 21,082 3 63,246 50,597 fi shermen, but were also obtained from by Sam Chapman at the Darling Marine 120 24,114 3 72,342 57,874 as far away as Long Island Sound, N.Y., Center, Walpole Hatchery (Fitzhenry 125 27,430 3 82,290 65,832 in an attempt to extend larval produc- et al., 1986). The basic formula con- tion seasons. Because local broodstock sisted of about 290 l of warm seawater, was only available during the months of 100 l of cultured algae, Isochrysis gal- when the lobster is transferred from the June through October, it was hoped that bana, and about 50 ml of brine shrimp trap to the holding tank or when the these warmer-water, egg-bearing lob- eggs. Supplying brine shrimp with a lobsterman transfers the lobster to the sters would hatch out sooner, thereby nutritionally balanced food source thus hatchery. extending the season. However, obser- resulted in a superior food item for the For an accurate estimation of stock- vations from experiments revealed that dietary requirements of larvae through- ing densities, Cutler established a reli- after adjustment to the cold waters of out all their rearing stages (Fitzhenry et able technique to count both Stage I Maine, the New York lobsters took al., 1989). The algae was also produced and Stage IV larvae via a mass-to-count longer to hatch (Fitzhenry et al., 1988). at Cutler under a process described by ratio. The equipment needed to do this By using local lobsters and applying Fitzhenry et al. (1988). consists of a small 7.62 cm diameter the techniques employed at the Biologi- During the course of rearing the PVC pipe that is 12.7 cm deep and has cal Station in St. Andrew’s, New Bruns- larvae, temperature was maintained at a 175 µm mesh screen epoxied to the wick (Waddy and Aiken (1984a) provide an optimal level of 20oC so that Stage bottom. Larvae were placed into this details), Cutler was able to successfully IV was reached in about 11 days (Table tube and weighed. Their number was expand its production season for about 3). Approximate cost to produce Stage estimated by the equations below: 1 month (Fitzhenry et al., 1989). IV lobsters is listed in Table 6. The weight (gm) − 0.069 Broodstock were maintained in two number of females needed to obtain No. of Stage I larvae = (1) separate tanks: an 800 l tank for indi- the required amount of Stage IV larvae 0.00633 viduals 2–4 weeks away from hatch- is presented in Table 7; however, cau- or, if using postlarvae, ing and a three-tiered tank for females tion should be used when using this weight (gm) − 0.09 currently releasing larvae (Fitzhenry et table because of the indices being used. No. of Stage IV larvae = (2) 0.03832 al., 1988). For Stage IV lobster produc- Although the method described by Per- tion, Cutler used 10 (400 l) conically kins (1971) was used to calculate the This system can be used at any hatch- shaped tanks supplied with aerated sea- number of eggs per female lobster and ery, and Fitzhenry et al. (1989) give full water. The aeration not only oxygenated can be used as a rough guide to estimate details. the seawater but also provided vigor- how many lobsters are needed for a pro- Release of Stage IV lobsters was duction season, it is not very accurate made in tidal fronts near shallow sub- 14 Chapman, S. 1991. University of Maine at since the number of eggs per female is tidal eelgrass or kelp beds. The lobsters Orono, Dep. Oceanogr., Ira C. Darling Mar. Cent., Walpole, ME 04573. Personal commun., always less than predicted. These losses were released through a large funnel 25 Oct. are usually due to eggs being dropped that had a 5 to 6 foot weighted exten-

61(2), 1999 31 sion attachment mounted to a tapered grown to trapable sizes and were cap- ment could sustain 30–60,000 lobsters snout (Fitzhenry et al., 1988). Release tured (Bulkeley, 1993). Wahle (1991a) annually (conservatively valued at over occurred at dusk (to reduce losses to and Wahle and Incze (1997) also placed $100,000 annually) (Steneck et al.15). visual predators) on a rising tide. Other hatchery-reared “blue” lobsters onto Currently, although Maine’s lobster experiments were being conducted to artifi cial cobble plots on a featureless hatcheries have provided information test the hypothesis that survival suc- sand-bottom cove in Maine and moni- (e.g. rearing and hatching techniques), cess increased in cobble areas com- tored the growth and densities of the only one lobster hatchery remains in pared with an unvegetated mud area juvenile lobsters over time. operation at the Mt. Desert Oceanarium, (Fitzhenry et al., 1989). Experiments on the behavioral traits Bar Harbor. Hatchery techniques are Besides the goal of enhancing natu- (swimming, burrowing, walking, and modeled after Cutler and have allowed ral stocks, the Maine hatcheries were lack of activity) of “blue” lobsters were the release of 40,000 postlarval lobsters interested in the development of “blue” also conducted using six different sub- to Bar Harbor, Winter Harbor, Northeast color morph lobsters as biological tags. strates: sand, eelgrass over sand, cobble Harbor, Seal Cove, Southwest Harbor, A long-term project was initiated in over sand, pebbles over sand, mud, and Islesford, Maine. Besides reseeding 1987 by researchers from the Darling and eelgrass over mud (Irvine et al.5). efforts, this hatchery provides postlar- Marine Center, Walpole Hatchery, and All lobsters used for this study were val lobsters for research. At the Big- the University of Maine at Orono and postlarvae and had carapace lengths elow Laboratory for Ocean Sciences, Machias (Irvine et al.5). However, it between 4.0 and 6.8 mm. No behav- Dr. Rick Wahle has requested 10,000 was unclear if “blue” lobsters would ioral differences were detected between larvae to develop a tracking system survive or behave in a manner similar the “blue” and normal (control) lob- (Schreiber, 1998). to wild-type individuals (Irvine et al.5). sters (Beal et al., 1998; Irvine et al.5). Several researchers speculated that However, smaller lobsters (4.0–4.9 mm Great Britain “blue” lobsters might be more easily CL) swam more often over sand and Homarus gammarus natural stock detected by predatory fi sh and thus swam least often over cobble than those enhancement is being tested by three suffer higher mortalities (Portersfi eld, of 6.0–6.8 mm CL. Conversely, larger groups in the United Kingdom. Experi- 1982; Fitzhenry et al., 1986; Plante, postlarval lobsters burrowed more read- ments were initiated by scientists from 1989; Syslo7). ily than small ones, mainly on the eel- the Ministry of Agriculture, Fisheries, Several pilot-scale experiments were grass over mud substrates. Walking and and Food (MAFF) directorate of fi sh- conducted in 1988 and 1989 to deter- lack of activity in this study corre- eries research at Burnham, Conway, mine the survival and growth of hatch- sponded to the previous observations of and Lowestoft (Richards and Wickins, ery-reared “blue” postlarvae released Botero and Atema (1982) and Hudon 1979). Following MAFF’s methodol- in the natural environment. In 1988, (1987) in that lobsters 4.0–4.9mm CL ogy, trials began by two other groups: 5,600–6,000 Stage IV and V “blue” spent more time walking. These results Sea Fish Industry Authority (SFIA) lobsters were released in a 20 × l00 tend to support the hypothesis that in Scotland, and North Western and m cobble patch at Damariscove Island, younger and smaller postlarvae use their North Wales Sea Fisheries Committee Maine. A day after release, densities energy searching for an appropriate sub- (NWNW-SFC) in Wales (Anonymous, were about 10 times lower than ex- strate rather than for growth (Botero 1991a; Burton, 1992; Cook et al., 1989). pected, had all of the lobsters recruited and Atema, 1982; Hudon, 1987; Cobb The latter two programs are summa- into the cobble patch. Curiously, 1 year et al., 1989a; Anonymous, 1991c; Irvine rized separately. later, densities were about the same as et al.5). At MAFF, ovigerous females are the day after the initial release. The ini- Other studies have been directed obtained from the fi shery for larvae pro- tial losses have been assumed to be due toward placing cobble (10–125 cm duction, and a 900 l recirculating system to predation and dispersion in the water diameter) in areas where recruitment maintains the egg-bearing lobsters column (Steneck et al.15). In 1989, 70 exists but habitat is limited. Population (Richards and Wickins, 1979). Larvae “blue” lobsters ranging in sizes from densities within these cobble patches are reared in 100 l cone-shaped poly- 12 to 14 mm CL were released. Within averaged 6 lobsters/m2 compared to 0.1 ethylene hoppers (originally designed 1 week their numbers decreased about lobsters/m2 in the adjacent sand area to hold agricultural feed). Ten such 40% to > 2/m2, but the population den- (Steneck et al.15). Thus, it would seem hoppers are formed into a recirculating sity remained constant at that level for that lobster populations could be fur- rearing unit, and each is stocked with over 10 weeks (Steneck et al.15). By ther augmented by adding cobble to 2,000 larvae that are hatched within a 1992, several of the blue lobsters had shallow coastal areas having good post- 2-day period. A second system contains larval supply but poor recruitment site ten 40 l fi berglass tanks similar to the 15 Steneck, R. S., R. Wahle, D. Dow, E. Blackmore, characteristics. Based upon fi eld mea- fi rst system, each stocked with 1,500 and B. F. Beal. 1990. Maine lobster enhancement surements of similar naturally occur- larvae (Beard and Wickins, 1992). program, phase 1 and 2. A proposal to the Maine ring densities of 3–6 early benthic phase Larvae are fed twice daily with frozen Aquaculture Inovation Center. Unpubl. rep., 19 p. 2 Avail. from Robert Steneck, Univ. Maine, Ira C. lobsters/m and about 0.5–1 adoles- mysid shrimp, Neomysis sp., and a sup- Darling Mar. Cent., Walpole, ME 04573. cent phase lobsters/m2, habitat enhance- plement of Artemia nauplii 3 times per

32 Marine Fisheries Review week. It takes 9–26 days for the larvae to reach Stage IV (postlarvae) with an average of 16 days at a temperature of 20oC (Beard et al., 1985). Once at Stage IV, the postlarvae are removed by a hand net, counted, and placed into individual compartments. The grow-out unit consists of a primary system of 6,212 l capacity with 5 rows each with 4 rearing troughs (2.94 m long × 0.51 m wide × 0.15 m high). The second system, which has a 12,116 l capacity, contains 4 rows, each with 4 troughs. Each trough contains 80 individual compartments (5 cm long × 5 cm long × 10 cm high) (Beard and Wickins, 1992). Postlarvae are fed twice daily with 2–6 pieces of mussel in the morning and 6–9 mysid shrimp in the afternoon. Reared at temperatures of 18°–21°C, Stages X to XII (about 11–15 mm CL) are reached in about 3 months (Beard et al., 1985). These juvenile lobsters are then tagged with a binary coded Figure 16.—Diagram of juvenile lobster showing position of microtag at the base of micro-tag (1 mm long × 0.25 mm diam- the fi fth walking leg. From Burton (1992). Used with permission. eter) which is inserted at the base of the fi fth walking leg (Fig. 16). Tag retention through several molts is about 90% or selected coarse substrates (stones sites between 1983 and 1988 (Anony- (Wickins et al., 1986). Marked lobsters 7–20 mm diameter) which offered suit- mous, 1995). Each lobster was tagged are detected by passing the whole lob- able crevices. In experimental releases, with a coded micro-tag and was released ster through an instrument which emits lobsters liberated at the surface took by MAFF divers onto various selected an audible sound if the tag is present, 5–10 minutes to reach the bottom (16 habitat patches in Bridlington Bay on and each year from 1988 to 1994, an m depth) (Howard, 1983). If released at the east coast of England (Bannister et average of 10,000 lobsters were tested the surface, these lobsters would be dis- al., 1991). Results from these releases for the presence of these tags. Full placed from the suitable site; however, have been very encouraging, with hun- details on experiments and techniques 90% of the lobsters released at a care- dreds of returns being documented: 26 used at MAFF are given by Richards fully selected site with a suitable bottom in 1988 (Bannister et al., 1989), 110 and Wickins (1979), Howard (1982, found shelter in 2 minutes (Howard, in 1989 (Bannister et al., 1990), 218 1988), Beard et al. (1985), Bannister 1982). in 1990 (Bannister et al., 1991), 115 and Howard (1991), Addison and Ban- Furthermore, water velocities on the in 1991 (Burton, 1993), 152 in 1992 nister (1994), and Bannister (1998). bottom signifi cantly affect the ability (Addison and Bannister, 1994), and 32 Beard and Wickins (1992) provide a of lobsters to move: large lobsters (15 in 1993 (Cook, 1995) for a total of comprehensive report on techniques cm TL) were exhausted by moderate 653 recaptures, ranging in age from 3 for mass culture, tagging procedures, currents of 30 cm/sec, while smaller to 9 years (Anonymous, 1995; Table transportation methods to release sites, lobsters (5 cm TL) could rest in the 8). Most were recaptured from areas release techniques, number of person- lee of small obstructions to avoid such clustered within 5 km of their initial nel needed, and time required for each currents (Howard and Nunny, 1983). release sites (Anonymous, 1995). A procedure. Because of this ability to avoid cur- large proportion of these recaptured During MAFF’s experimental lob- rents and the ability to fi nd shelter in lobsters were of legal size and ten were ster enhancement trials, several studies 2 minutes, Howard concluded that 80% egg-bearing females; thus, there was no assessed the reaction of hatchery-reared of hatchery-reared stock released into doubt that hatchery-reared stock were lobsters to substrates and water currents suitable substrates had as much chance contributing to the natural stock (Ban- in an attempt to determine the proper of survival as their wild counterparts nister et al., 1989, 1998). location for the release of lobsters. (Howard, 1982). Estimated returns on investment for Howard and Bennett (1979) reported Using the above methodology, MAFF stocked lobsters are encouraging. For that postlarval lobsters (8 mm CL) read- released about 49,000 hatchery-reared example, if mortality is assumed at 10% ily burrowed into fi ne cohesive mud Stage XII lobsters at 80 different reef per annum, 40–50% of those released

61(2), 1999 33 Table 8.—Recaptures of hatchery-reared lobsters released in the United Kingdom from MAFF and NWNW SFC hatcheries (source: Burton (1993), Cook (1995), and Bannister (1998)). In 1990 the ﬁ rst lobsters of commercial size were screened to determine Numbers recaptured whether the tag was present; 105 tagged Hatchery and Number lobsters were recovered. Of these, 33 2 2 year of release released 1988 1989 1990 1991 1992 1993 1994 were from the 1984 cohort and ranged MAFF3 in size from 87 to 102 mm CL (with a 1983 2,390 14 19 6 0 0 0 (65-82) (78-96) (88-96) mean size of 93.24 mm CL), while 67

1984 8,616 12 67 97 25 14 0 were from the 1985 cohort and ranged in (52-78) (54-95) (67-100) (85-101) (87-108) size from 84 to 99 mm CL (with a mean 1985 7,979 0 24 112 21 5 0 (70-92) (83-114) (81-106) (86-97) size of 89 mm CL). In addition, ﬁ ve 1986 11,562 0 0 1 28 39 5 lobsters were captured from the 1986 (88) (85-96) (85-100) 1987 12,629 0 0 2 41 72 20 cohort, ranging in size from 86 to 96 (76, 82) (70-102) (85-108) mm CL (mean of 90.2 mm CL) (Table 1998 5,952 0 0 0 22 7 (84-108) 8) (Cook, 1990). During the recapture NWNW/SFC4 experiments of 1991, some lobsters 1984 1,250 0 0 33 6 3 0 0 gave off a false positive response when (87-102) (87-100) (113)5 1985 3,750 5 0 67 74 31 11 0 passed through the machine. Dissec- (51-58) (84-99) (84-106) (84-102) (84-102) (96)5 tion revealed that these lobsters were 1986 2,438 0 3 5 28 26 2 0 (56-58) (86-96) (85-105) (85-108) (96)5 not tagged, but their tissues contained 19876 5,079 0 0 0 2 36 23 0 (88, 90) (83-96) (93)5 small particles of rusted metal which 1988 6,706 0 0 3 40 44 11 were picked up by the extremely sensi- (85-86) (84-101) (90)5 (95)5 tive detection machine. The source of 1 Numbers in parentheses represent size in m CL. these rust particles remains unknown 2 Data still being processed. (Cook, 1992). In total, 453 lobsters have 3 Ministry of Agriculture, Fisheries, and Food. 4 North Western and North Wales Sea Fisheries Committee. been recaptured, representing a recap- 5 Mean CL size of lobsters recaptured. ture rate of 2.4%. Of these lobsters, 6 Animals released in December 1986 had the same code as those released in 1987. 445 were of a legal, commercial size which varied from 85 mm to 103 mm could reach market size in about 5 by the 4.5 economic multiplier (which CL (Cook, 1995). The sex ratio of the years. From those, 70–80% would be represents the worth of the lobster as recaptures was skewed towards females available for capture by the ﬁ shery indi- it is sold and resold relative to the per- in approximately a two to one ratio (288 cating a possible tenfold increase in sonnel who are employed to hold, ship, females versus 157 males), but only value (Wickins, 1983). Furthermore, and prepare lobsters), a total economic 20 out of 195 females examined were with survivors contributing their own value of $4,097,700 would be realized. ovigerous. These were slightly larger progeny to the wild population, the than nonovigerous females (93 mm return on investment could be com- North Western and vs. 90 mm CL) (Cook, 1995). Burton pounded (Aiken and Waddy, 1989). At North Wales Sea (1993), Addison and Bannister (1994), 15 pence (about US$0.30) to produce Fisheries Committee Cook (1995), and Bannister (1998) a lobster the size of 3 cm, an operation Following the techniques used by provide comprehensive updates on the could break even with only a 3% sur- MAFF, a hatchery was built at the Uni- United Kingdom’s stock enhancement vival rate (Anonymous, 1982). versity College of North Wales, Menai program. Optimistically, if proper procedures Bridge, and lobsters are being released are used during rearing and release off Aberystwyth, Cardigan Bay, North Scotland operations, no less than 42% of those Wales (Bannister et al., 1989). Produc- Lobsters are also reared at the SFIA lobsters being released would reach tion began in 1984, with a maximum Marine Farming Unit, Ardtoe, Argyll, commercial size (Anonymous, 1982). rearing capacity of 3,072 postlarval lob- West Scotland. The Ardtoe hatchery With these predictions and the use sters. From 1984 through 1988, 19,237 was established to provide up to 10,000 of Syslo’s economic impact statement postlarvae were reared and released juvenile lobsters/year in two batches. (Table 4), the return on investment could (number adjusted after tag loss). Before Berried females are obtained from the be very high. For example, if 500,000 1988, the smallest lobster released was wild and held in troughs 3 m long × 1 postlarval lobsters were released, and 10 mm CL, with the mean size of each m wide × 0.6 m high (Fig. 17). Once the an assumed 42% (as noted above) sur- batch at 12–14 mm CL. In 1988, released eggs hatch, larvae are collected, counted, vived to reach commercial size, then lobsters ranged from 6 to 45 mm CL and transferred to a rearing system the 290,000 (42% of 500,000) animals (Cook et al., 1989). Five lobsters were where they are held in 80 l polypropyl- would, at a value of $3.14/pound, repre- recaptured in 1988, ranging in size from ene hoppers (Fig. 18, 19) and fed on sent a potential $910,600 to lobstermen. 51 to 58 mm CL, and all were from the frozen mysid shrimp and chopped mus- By multiplying this potential $910,600 1985 releases (Cook et al., 1989). sels. At 16°–17°C larvae reach Stage

34 Marine Fisheries Review Figure 17.—Broodstock fi berglass trough 3 m long × 1 m wide × 0.6 m high. PVC is used for all pipe and valve work. 2 mm mesh screens cover the 15 cm diameter PVC frames on the outfl ow and overfl ow pipes. Each broodstock trough houses 15 egg-bearing females. From Burton (1992). Used with permission.

Figure 18.—Larval kreisels made from 80 l polypropylene hoppers, originally designed for agricultural purposes. The diffuser plate is made of PVC with holes drilled around the perimeter to provide water agitation. The 15 mm diameter standpipe is screened with 2 mm mesh screening. From Burton (1992). Used with permission.

IV in 12–18 days, but survival is some- tems for rearing to Stage XII: one is a Both systems house the lobsters indi- what low (5–30%, averaging 14%) horizontal trough system, as described vidually (Burton, 1992; Fig. 21) and (Burton, 1992). At Stage IV, the post- above (Fig. 19) and the other is a verti- they are fed manually or with an auto- larvae are transferred to one of two sys- cal stack (Fig. 20). matic feeding system (Wickins et al.,

61(2), 1999 35 Figure 20.—Vertical stacking system made of frames of aluminum and trays 63 cm long × 44 cm wide × 13 cm high. Rollers are used to make the drawers holding the trays move backwards and forwards in the frame during feeding times. From Burton (1992). Used with permission.

ket-sized lobsters, ranging in size from 85 to 102 mm CL. Berried females and males with mature testes have been Figure 19.—Trough ongrowing system for juvenile lobsters. The fi berglass trough’s found, confi rming that hatchery-reared inner dimensions are 254 cm long × 57 cm wide × 15 cm high. The siphon produces stock will mature and reproduce in a 25 mm rise and fall of the water level every 15 minutes. From Burton (1992). Used the wild. Samplings indicate that these with permission. recaptures are from the 1984–85 cohorts (Table 9) (Anonymous, 1991b; Burton, 1987) which can dispense mysid shrimp sible condition and to minimize preda- 1991, 1992). or pelleted diets. During the growth tion during release. In 1990 releases of Stage V juve- process, the diet is similar to that of the SFIA began releasing lobsters at niles began using a smaller microtag larvae, but it may be supplemented with Ardtoe and Scapa Flow in 1984, and (0.5 mm long × 0.25 mm diameter). For Artemia (Burton, 1992). Survival from lobsters have been captured ever since the fi rst release, 1,200 Stage V–VI lob- Stage IV to Stage XII is about 80%. (Anonymous, 1991a). An extensive sters were tagged, with 1,170 surviving Two systems are used to release the study conducted by divers in 1985 recov- 3 days later (77.5%). Unfortunately, in lobsters: a stacked tray (Fig. 22 and ered 5 microtagged lobsters (Walker, subsequent batches a higher mortality 23) and a pipe release (Fig. 24). Both 1986). However, 1989 saw the fi rst was experienced in lobsters of less than methods are designed to convey the recoveries of market-sized lobsters with 6 mm CL; however, this mortality has microtagged Stage XII juveniles to the 6 having carapace lengths of up to 100 been subsequently stabilized (Burton, selected substrate (methodology after mm (Burton, 1992). The 1990 recover- 1991). Tag retention is 96% after ani- MAFF: Burton, 1992) in the best pos- ies were more successful with 105 mar- mals have molted at least once. No data

36 Marine Fisheries Review from these releases is available thus far (Cook, 1995). Beginning in the summer of 1995, a pilot-scale lobster hatchery was established on the Scalloway Islands, Shetland to assess the practicality of producing juvenile lobsters for stock enhancement (Watt and Arthur, 1996). Egg-bearing females, obtained from local fi shermen, were held fi rst communally in a large tank and then were separated just prior to their eggs hatching. The females were fed on whitefi sh, salmon, and crab meat 2–3 times weekly. Despite these holding conditions, 500 g females, known to produce ~700 eggs, only produced about 500 larvae, which were transferred to three different kinds of larval rearing systems: 1) an 80 l con- Figure 21.—Exploded diagram of tray construction for the stacking and transporta- ical tank with air stones at the bottom tion systems. The polystyrene trays, 540 mm long × 432 mm wide × 44 mm high, for water agitation, 2) an 80 l conical house 80 lobsters individually. In the stacking system, two trays are placed into each tank with a diffuser plate (similar to drawer, with the upper tray having a bottom made of 2 mm mesh. The trays are taped that in Fig. 18) for water agitation, and together. From Burton (1992). Used with permission. 3) individual trays suspended in water. Only 10% of the larvae in both conical tank arrangements survived; none of the larvae survived when housed individually in submerged trays (Watt and Arthur, 1996). Stage IV postlarvae were produced in 8–12 days and were then transferred to ongrowing, individual bins. They were reared only to Stage V, with a 77% survival rate, and then were released via a 3-inch fl exible pipe in a design similar to that diagrammed in Fig. 24. In 1995, 1,000 juveniles were released in this manner, but the hatchery was expected to expand within the next couple of years to produce over 30,000 juveniles per year (Watt and Arthur, 1996). Ireland Recently, Ireland has become interested in the hatching and release of Homarus gammarus juveniles. The fi rst prototype lobster hatchery in the country was constructed at the University College Galway’s Shellfi sh Laboratory in Carna, Galway (Grogan, 1997). Here they modelled their entire project after a Cutler, Maine hatchery program (Browne and Mercer, 1998). Female lobsters are maintained in barrels of circulating seawater. Larvae are then transferred to Figure 22.—Arrangement of trays in a frame. These stacks are taken to the special tanks at a density of 1,000 indi- sea by divers for release of tagged juveniles. From Burton (1992). Used with viduals per tank and are maintained at permission.

61(2), 1999 37 Figure 23.—Diagram of how lobsters are released by divers using the stack method. From Burton (1992). Used with permission.

Table 9.—Recaptures of hatchery-reared lobsters released from SFIA Ardtoe and Scapa Flow hatcheries (source: Burton (1993) and Anonymous (1995)). the lobsters reach Stage IV, they are released. Methods of release involve Numbers recaptured1 Hatchery and Number transporting the lobsters individually in year of release released 1985 1986 1987 1988 1989 1990 1991 1992 1993 plastic trays covered with tissue paper. SFIA2/Ardtoe Each tray is then lowered in a lobster pot 1984 451 2 2 3 9 8 1 0 usually on a rocky bottom. The tissue (31, 41) (45, 59) (53-57) (43-84) (71-102) (94) 1985 1,268 0 0 2 6 5 1 paper dissolves within 20 minutes after (54, 70) (56-76) (76-86) (76) 1986 513 0 0 0 0 0 it has entered the seawater, and the lob- 1987 553 0 1 4 8 4 1 sters can then escape into the rocks. The (56) (48-60) (55-63) (59-84) (65) 1990 259 0 13 pots keep predators from attacking the SFIA2/Scapa Flow lobsters during this time. Currently, the 1984 4,469 3 n.d. 18 3 16 84 68 233 13 hatchery can produce over 30,000 juve- (18-22) (52-56) (49-55) (56-100) (81-102) (84-118) 1985 3,800 0 0 0 1 10 3 123 13 nile lobsters per year and will continue (67) (82-102) (84-87) 1986 2,356 0 0 0 0 0 53 to do so for about seven years (Grogan, 1987 3,610 0 0 0 1 373 53 1997; Browne and Mercer, 1998). (85) 1988 2,260 0 0 0 83 43 1989 3,025 0 0 13 Norway

1 numbers in parentheses represent size in mm CL. During the early 1980’s the worlds 2 Sea Fish Industry Authority. largest “commercial” lobster hatchery 3 CL data not available. 4 n.d. No data available was constructed at Kyrksœterøra, south of Tronheim, Norway (Schjetne, 1987; temperatures of 21oC (Grogan, 1997). Carne while the brine shrimp eggs are Tveite and Grimsen, 1990), in response Larvae are fed a mixture of algae and imported from Great Salt Lakes in Utah, to the collapse of the Norwegian lobster brine shrimp. Algae are cultivated at USA. Approximately two months after ﬁ shery, whose landings fell from 500 t

38 Marine Fisheries Review Figure 24.—Diagram of how divers release lobsters via the pipe method. A header tank, incorporating a reservoir is mounted on the stern of the boat and is supplied by a pump. The rear portion of the header tank has a 75 mm diameter ﬁ tting to which an armored pump suction hose is attached. The lobsters are gently suctioned through the pipe and directly delivered to the bottom substrate. From Burton (1992). Used with permission. in the 1950’s to less than 30 t in the past have been described in Erenst (1985) this time, larvae are fed frozen Artemia few decades (van der Meeren, 1994). and Grimsen et al. (1987); however, 2–3 times/day (Grimsen et al., 1987). The concept of this operation differed the management of the hatchery has Mortality is very high during this phase from other hatcheries in that H. gam- changed and so have the rearing tech- of operation. Under the best of circum- marus lobsters were reared under opti- niques. Since 1989, Norway’s Institute stances it is 50%, but more often it is mal conditions to 1 year of age in a of Marine Research has managed this closer to 85% (Uglem17). Upon reach- large circular pool (Erenst, 1985; Sch- hatchery (van der Meeren and Nœss, ing Stage IV, the lobsters are transferred jetne, 1987), similar to that described in 1991). to separate boxes as described in Grim- Van Olst et al. (1977). They are released Female lobsters are obtained from sen et al. (1987). Here they are fed speciﬁ cally with the idea of replen- the wild for eggs. About 1,000–10,000 frozen Artemia 5–7 times/week at 5% ishing a depleted stock. The hatchery larvae are produced from each female, of their wet weight. Moist pellets and can produce 120,000 Stage XIII lob- depending upon their size. These larvae the computer-controlled feeding system sters, but rarely exceeds 30,000–50,000 are transferred to kreisels (350 l) where described in Grimsen et al. (1987) are (van der Meeren16). Culture techniques they spend 8–18 days, depending on no longer utilized (Uglem17). water temperature. Optimal tempera- 16 van der Meeren, G. I. Institute of Marine Re- ture for this growth process is about 17 search, Austevoll Aquaculture Research Station, o Uglem, I. 1991. Institute of Marine Research, N-5392, Storebo, Norway. Personal commun. 12 24 C which results in Stage IV lob- The Lobster Hatchery, P.O. Box 130, 720 Kyrk- July 1989 and 21 Feb. 1997. sters in 8–12 days (Uglem17). During saeterora, Norway. Personal commun., 22 Nov.

61(2), 1999 39 The cost of producing a Stage XIII coded micro-tags (van der Meeren et al., 1994). Since then, the lobsters have lobster is estimated at US$2.00, but with 1990). This release was the fi rst large- been placed in plastic cages which are current practices it may now be possible scale release experiment and, from 1991 then immersed for 30 minutes in a to reduce this cost as much as 30–40% to 1994, release of 15,000 to 30,000 basin of ambient seawater. Observations (Uglem17). From 1979 to 1989, 100,000 tagged juvenile lobsters/year occurred during release indicated that the lob- 1-year-old Stage XIII juveniles were (van der Meeren et al., 1990, 1998). sters sank slowly to the bottom, landing released (Grimsen et al., 1987; Addi- Actual releases have been less: 29,693 in a walking position; they then slowly son and Bannister, 1994) but were not juveniles were released in 1991, 29,919 moved into shelters within 20 minutes tagged. However, lobsters released into in 1992, 17,360 in 1993, and 27,414 (van der Meeren, 1991b). Other stud- open and/or enclosed areas with artifi - in 1994. In October 1992, the fi rst 19 ies are underway to assess the impact cial shelters and fed blue mussel fl esh legal-sized male lobsters were recap- that released lobsters may have on their provided some information on growth tured and ranged in size from 83 to wild counterparts and to determine pop- and how that affected the density at 91 mm CL. Their tags revealed that ulation dynamics in the same region which lobsters can live. These data sug- these lobsters were hatched in 1988 without the addition of released lob- gest a carrying capacity of one 2-year- and released in April of 1990 (van der sters. These studies will provide infor- old lobster/4–5 m2, based on releases Meeren and Naess, 1993), suggesting mation to assess whether sea ranching of 45 1-year-old lobsters into an area more rapid growth rates than previously is profi table and whether it represents of 50 m2 and a recapture of 12 lobsters expected. In fact, the cultured lobsters an effective lobster management tool, 2 months later, as well as subsequent seem to recruit to the fi shery within particularly in terms of its expense (van releases of 36 1-year-olds with a recap- 3–4 years and can support the fi shery der Meeren and Naess, 1993). ture of 11 lobsters after 10 months for at least 5 years. Recapture rates for (Tveite and Grimsen, 1990). single year classes range from 5 to 8%. France The 715 lobsters retrieved between Furthermore, ovigerous females from Prior to 1972, France attempted to 1983 and 1988 were distinguishable hatchery origins are now of equal pro- build up a reserve of adult lobsters, from wild counterparts by morphologi- portion in the population as wild oviger- particularly berried females, by estab- cal traits (such as double seizer/cutter ous females and are thus contributing to lishing sanctuary zones where all fi sh- claws). They ranged in size from <22 the overall reproductive effort in these ing was prohibited. Sixteen zones were cm to >25 cm and included berried release sites (van der Meeren16). established in 1963 and all saw lobster females. In 1989, recaptured, hatchery- Behavioral traits of hatchery-reared increases (Lorec, 1987; Latrouite and reared lobsters accounted for more than lobsters have also been examined, and Lorec, 1991). Beginning in 1961, berried 50% of the fi shermen’s catch (Tveite they appear to exhibit the same behav- females were also stocked in tanks until and Grimsen, 1990). ior as their wild counterparts. They can their larvae hatched, grew, and metamor- More recently, a pilot-scale experi- adjust their behavior according to shel- phosed into Stage VI postlarvae. They ment at Norway’s Austevoll Aquacul- ter and predation risk, light intensity, were then released into the wild. ture Station involved rearing hatchery and individual distance to conspecifi cs Because neither of those techniques lobsters to 1- and 2-year-old juveniles. (van der Meeren, 1990). However, van produced noticeable improvement in A total of 9,800 lobsters were marked der Meeren (1991a) found that behav- landings, two hatcheries were estab- by branding a spot on either the fi rst ioral responses depended on the kind lished in 1972—one on the Isle of Yeu joint of the tail or the center of the cara- of treatment given to lobsters prior to and the other on the Isle of Houat; a pace; these lobsters were then released release. Several stresses were intro- third was established later at the Isle (van der Meeren and Naess, 1991). duced in her study: sudden exposure to of Sein (Lorec, 1987). The hatcheries These marks are capable of withstand- light, pressure, and water loss. “Pelagic were designed according to the Japa- ing several molts and are still identifi - rushes” (upward swimming) decreased nese system for raising shrimp, rather able once the lobster is recaptured (van when the number of stresses were than the techniques of Hughes et al. der Meeren, 1990). None of these lob- reduced. The lowest proportion of rush- (1974). From 1972 to 1977, these hatch- sters were recaptured in 1989; how- ing and highest proportion of walking eries concentrated on increasing the sur- ever, marked lobsters were recaptured appeared in treatments where no stress vival of larvae to Stage V juveniles. They in 1990. These lobsters had increased was applied (van der Meeren, 1991a). achieved survival rates of 90% to Stage about 26.5 times in weight and 40% in The best transportation methods to III by feeding larvae to excess on living carapace length (van der Meeren and release sites were also investigated. food, particularly spider crab larvae and Naess, 1991). First, lobsters were transported to the Artemia salina (which were raised on In 1990, a large-scale release pro- release site in thermal boxes fi lled with algal cultures of Tetraselmis and Phaeo- gram began using 13,500 1.5-year-old wet, chilled newspapers and then placed dactylum). Survival rates to Stage IV and lobsters (21.1 mm CL, 1988 year class) on the bottom directly. However, these V were about 75–80% (Audouin, 1974). and 7,700 6-month-old lobsters (12.1 animals were very sluggish, slow to Up to 250,000 Stage V juveniles were mm CL, 1989 year class). These lob- gain shelter and, as a result, experi- released and 15,000 1-year-old lobsters sters were tagged with l mm binary enced high mortality (van der Meeren, were released; however, no improve-

40 Marine Fisheries Review ment in landings occurred and it was sters remained near or in the site of their only other option for selling the suble- concluded that these releases were not release. However, two of the hatcher- gal-sized lobsters is to the less lucra- enhancing stocks (Henocque, 1983; Le ies were closed before these releases tive canner business (Aiken and Waddy, Gall et al., 1983). (Lorec, 1987), and, despite the prom- 1995). Beginning in 1978 and continuing ising initial results, no further juvenile Eyestalk ablation, which was fi rst through 1983, the hatcheries began lobster releases were made after 1987 recognized to accelerate molting and to focus on the impact that released (Lorec, 1987). Since then, restocking growth by Zeleny (1905), was fully Stage V lobsters might have on the fi sh- efforts and monitoring of recaptures investigated. Mechanisms controlling ery. Juvenile lobster marking methods were discontinued since the low number molting were studied and reviewed were examined, and hybrids of Homa- of recaptures was seen as demonstrat- by Passano (1960) with such infor- rus americanus and H. gammarus were ing that enhancement did not occur mation creating a baseline for exper- used prior to the development of the (Latrouite, 1998; Latrouite4). iments with ablated lobsters. Initial Bergman- Jefferts microwire tag (Lorec, work, however, produced confl icting 1987; Latrouite and Lorec, 1991). Since Product Enhancement results. Flint (1972) reported that bilat- the only distinctive mark was a extra ros- Product enhancement involves hold- eral eyestalk ablation on American lob- tral spine, this method of phenotypically ing low-valued lobsters until they sters increased, rather than decreased, marking released animals was discon- become marketable. Lobsters of lower the time between molts. In contrast, tinued, but not until after 1,300 1-year- value may be soft-shelled, missing one others succeeded in accelerating the old hybrids were released. Recaptures or both claws, or one molt away from molt cycle with ablated lobsters but at were expected in 1980 (Audouin, 1981), legal size. Such aquaculture started as the cost of lower survival (Rao et al., but none were reported. Fishery sta- early as 1872 in Massachusetts (Anony- 1973; Sochasky et al., 1973). Stewart tistics were also used to monitor the mous, 1873, 1874) with large quantities and Castell (1976) suggested that poor success of the hatchery efforts, but, as (about 40,000) of low-valued lobsters survival in ablated lobsters was related in other countries, natural fl uctuations placed into enclosed basins. The lob- to diet and nutrition, as was observed in in landings overshadowed any impacts sters were fed during the summer crayfi sh by Smith (1940). that released lobsters may have made. months only (Anonymous, 1873; Rath- Further experiments demonstrated Efforts were then directed toward bun, 1886). The basins were natural that eyestalk-ablated lobsters fed an ade- raising 1-year-old lobsters, and in 1984 enclosures formed by land or rocks, quate and balanced diet were capable these 1-year-olds (12 mm CL) were with one end constructed as a dike of increased growth and high survival microwire-tagged with 1 mm magneti- (Anonymous, 1873). Results were sat- (Mauviot and Castell, 1976; Bishop and cally coded tags. At the same time, a isfactory (Anonymous, 1873; Rathbun, Castell, 1978); consistent weight gains program was established to determine 1886), but depended on the infl uence of of 70% were achieved (Bishop and Cas- how juveniles released in natural envi- the ambient water temperatures. tell, 1978). More recently, Coulombe ronments would respond (Latrouite and Because a signifi cant proportion of and Motnikar (1989) experimented with Lorec, 1991). Juveniles were found to the Canadian lobster catch is rather two methods of unilateral eyestalk abla- prefer the same types of habitats as small—usually one molt away from tion: removal by excision and cauter- postlarval lobsters, particularly algal- legal size (Pringle et al., 1983)—a ization and removal by strangulation. covered small rocks with many intersti- 5-year Canadian study was undertaken Lobsters 76–81 mm CL were kept in tial spaces between them (Bertran et al., in 1963 to determine the effects of vari- an open seawater system with ambient 1985). Optimal densities of juveniles in ous combinations of temperature, light, water temperatures and natural photo- naturalistic environments were less than lobster density, shelter, diet, sex, size, periods. All lobsters were fed to satia- 1/m2, and the hatchery-reared juveniles and maturity on accelerating the growth tion with thawed herring and shrimp. were less hardy than their wild counter- rates of sublegal lobsters. Females tend However, neither of these two methods parts (Latrouite and Lorec, 1991). to skip molts in order to spawn, so had a signifi cant effect on the molt rate Between 1984 and 1987, the hatch- only males were used. Gains in lobster (Coulombe and Motnikar, 1989). eries released 25,480 tagged juveniles weights were not greater than losses Owing to the variable eyestalk abla- directly onto sites believed favorable from mortalities and mutilation, and, in tion results, cage culture methods were for their survival. Researchers expected most cases, the timing of molting was reassessed, and in 1983 a suspended that 4–5 years would elapse before later than what had been predicted. Con- fl oating system yielded encouraging legal-sized lobsters would be recap- sequently, this project was terminated in results (Fradette, 1984a, b; Fradette tured (Lorec, 1987); however, despite 1968 (McCleese, 1969). Wilder (1971, et al., 1987). Lobsters ranging from low rates of recaptures in 1988 (1 lob- 1972) suggested that with the existing 76 to 90 mm CL were maintained on ster), 1988 (7 lobsters), and 1989 (14 state of knowledge, such rearing was crab, Cancer irroratus, or pelleted diets lobsters), nearly all were larger than not economically feasible. Nonetheless, described in Gagnon et al. (1984). Cost legal size, indicating a faster growth rate this kind of culture has been reeval- studies, with respect to time, personnel, than anticipated (Latrouite and Lorec, uated several times, because the eco- and marketing needs yielded the fi rst 1991). Furthermore, the recaptured lob- nomic concept is sound, given that the economic feasibility study for a fi rm

61(2), 1999 41 engaged in semi-intensive lobster culti- not fed; however, food may have been Communal rearing systems were also vation (Fradette et al., 1987). obtained from particles in the water or designed which were less complex, less More recently in the Bay of Fundy, from animals growing inside the cars expensive, and required less space. Young-Lai and Aiken (1989) grew sub- such as mussels, oysters, and marine Although these rearing systems experi- legal (75.2 to 80.9 mm CL) lobsters in worms. enced some success, low survival and cages through one molt to commercial The fi rst growing season produced nonuniform-sized lobsters were major size. These lobsters were fed on a diet encouraging results with lobsters rang- fl aws (Van Olst et al., 1976a; Sastry of commercial salmon ration and raw ing in size from 106 to 159 mm TL, with and Zeitlin-Hale, 1977; Carlberg et al., herring, and the biological feasibility of a mean of 122 mm. While these experi- 1979). These problems were partially culturing lobsters in cages through one iments were exploratory and incon- alleviated by providing a variety of sub- molt was demonstrated. However, there clusive, they did demonstrate great strates in the communal tanks (e.g. oyster were no data on the venture’s economic variability in growth rates between com- shells), as well as by sorting out individ- feasibility. munally reared lobsters, with the great- uals according to their size and removing In Scotland, a different approach was est growth rates occurring in those their chelipeds (Aiken and Young-Lai, used by Futcher beginning in 1968 lobsters reared at lower densities (Mead, 1981; Aiken and Waddy, 1988; Waddy, (Mundey, 1969). Immature or market- 1902; Mead and Williams, 1903). This 1988; Waddy et al., 1988). able lobsters were placed in “cages” (12 variation in growth rate has been con- Likewise, many experimental sys- m wide × 12 m long × 2.4 m deep), which fi rmed by more recent researchers in tems have been developed and ana- were anchored in sheltered coves at suf- California and Canada (reviewed by lyzed for the complete grow-out phase fi cient depths to avoid freshwater runoff Van Olst and Carlberg, 1979; Waddy, from juvenile to market size (Schuur from the shore. Lobsters were main- 1988; D’Abramo and Conklin, 1985). et al., 1974; Sastry, 1975; Hand et al., tained on “trash” fi sh (from local fi sh- Growth data from earlier studies at 1977; Van Olst et al., 1977; Mickelsen ermen) and crabs, but they also fed on Wickford also provided some insight et al., 1978; Richards and Wickins, organisms growing on the cage material. on estimating lobsters’ growth rate in 1979; Conklin et al., 1981; Beard et al., These lobsters were held until the market nature (Hadley, 1906a). John Hughes at 1985; Ingram, 1985). Of these newer price was high or until immature lob- the Massachusetts State Lobster Hatch- systems, the fl ushing tray designed by sters grew to marketable size (Mundey, ery retained individual Stage IV lob- researchers at the St. Andrew’s Bio- 1969; Bowbeer, 1971). Despite its suc- sters from each years’ hatch to study logical Station, Can., and the Univer- cess (Bowbeer, 1971), operations ceased growth rates and held some animals for sity of California, San Diego, appeared shortly after 1971, due to a disagree- as long as 10 years (Hughes and Mat- to yield the best results (Van Olst et ment between landowners (Burton18). thiessen, 1962, 1967). Lobsters were al., 1976b). Other advancements have maintained at ambient seawater temper- been made with automatic feeding sys- Full Grow-out atures and fed on fresh fi sh and shellfi sh. tems developed for larval (Serfl ing et Interest in culturing lobsters from Records were kept on molting frequen- al., 1974b; Fig. 13) and juvenile rear- egg to maturity arose in 1900 at Wick- cies, growth rates, food requirements, ing (Grimsen et al., 1987; Wickins et ford, R.I. (Mead, 1902; Mead and Wil- and mating behavior (Hughes and Mat- al., 1987). Several systems have also liams, 1903). After hatching and rearing thiessen, 1962, 1967). These lobsters been developed to rear juvenile lobsters larvae to Stage IV, the lobsters were were hatched and reared to legal size individually (Chanley and Terry, 1974; placed communally into “cars.” These (about 450 g) in a little over 3 years, as Lang, 1975; Conklin et al., 1981; Beard cars were constructed with galvanized opposed to the 6–10 years required in and Wickins, 1992; Burton, 1992). A iron screen sides which permitted free nature. complete description of these culturing circulation of water. Each car was pro- Given these advantages, Hughes spec- techniques can be found in Aiken and vided with sand, gravel, seaweed, etc., ulated that commercial lobster farming Waddy (1995). to simulate natural habitats. During the was possible (Hughes, 1968b). By rear- Experimental pilot-scale operations summer months, these cars were sus- ing lobsters in optimal levels of salinity for commercial lobster production were pended from a fl oating houseboat to and oxygen, providing them with proper attempted by some in the 1970’s, uti- a depth of 45 cm. The lobsters were food, and keeping the temperature at lizing much of the knowledge gained maintained at ambient seawater temper- a constant 20°C, market-sized lobsters from work by Hughes (1968b), Hughes atures and fed various foods. In the fall, could be produced in less than 2 years and Matthiessen (1962, 1967) and from the cars were lowered to about 2.4–3 m (Hughes et al., 1972). A further reduc- the communal rearing studies of Van deep and maintained there until spring. tion in this time was obtained by pheno- Olst et al. (1976a), Sastry and Zeit- During the winter months, lobsters were typic selection of fast growing lobsters lin-Hale (1977), and Carlberg et al. (Hughes et al., 1972), and owing to (1979). Unfortunately, economic rea- these successes, several experimental sons and lack of biological informa- 18 Burton, C. A. 1991. Sea Fish Industry Author- lobster farms arose (Shleser, 1971; Shle- tion brought an end to these projects. ity, Marine Farming Unit, Ardtoe, Acharacle, Argyll PH36 4LD Scotland. Personal commun., ser and Tchobanoglous, 1974; Van Olst For example, in the mid-to-late 1970’s, 24 Apr. and Carlberg, 1979). a “mini” lobster farm was created at

42 Marine Fisheries Review the Long Island Lighting Company’s and Aquaculture Enterprises of Califor- useful as a supplement (Burton, 1991, power generating station in Northport, nia (Hall, 1979; Anonymous, 1987a). 1992). Further studies are underway to N.Y. Together, Frederick B. Wishner of Similarly, the Department of Fisheries develop a better pelleted diet (Burton, New York, Anthony D’Agostino from and Oceans in Canada also supported 1992). These studies have mainly served the Marine Science Institute at Mon- such research (Ford and Van Olst, 1975; to show the superiority of one type of tauk, N.Y., and Christopher Gross, a Van Olst et al., 1980; D’Abramo and diet over another in terms of cost, sur- biologist from Long Island Lighting Conklin, 1985), and the United King- vival, and larval growth. Company in Hicksville, N.Y, success- dom promoted programs culturing the The studies on artifi cial diets, in par- fully reared lobsters in cooling waters European lobster, H. gammarus, to ticular, have shown that long-chain poly- from the power generating station in marketable size (Richards and Wick- unsaturated fatty acids are extremely a canal (Portersfi eld, 1982). Unfortu- ins, 1979; Anonymous, 1980; Richards, important for the survival of both the nately, this culture effort lasted only 1981; Beard et al., 1985). larvae and postlarvae. Because of its about 6 months (Gross19). Another Nutritional requirements were also low cost and ease of use, Artemia salina attempt was made by A. Gmeiner in examined to help formulate diets for is an excellent food source for rearing Woodside, N.Y., in the early 1970’s cost-effective growth. Diets composed larvae from hatching through the fi rst 6 using a closed water system in the base- of material that lobster larvae would months of growth (Aiken and Waddy, ment of his home. However, his lob- never encounter in nature have been 1989); however, different types of brine sters contracted a disease, caused by extensively examined. These include shrimp vary in fatty acid content and Fusarium sp., which resulted in heavy feeds such as living Artemia cysts or thus in quality (Fujita et al., 1980). mortality prior to molting (Lightner and adults (Conklin et al., 1975; Carlberg While optimal feeding schedules for Fontaine, 1975), and this farm was also and Van Olst, 1976; Conklin et al., live and frozen A. salina have been discontinued (Fontaine20). 1978; Rosemark, 1978; Capuzzo and determined (Carlberg and Van Olst, These early failures did not discour- Lancaster, 1979; Bordner et al., 1986; 1976; Aiken and Waddy, 1989), the age further experimental studies, but the MacKenzie, 1987), frozen adult brine lack of development of a cost-effec- lobster was fully assessed to determine shrimp (Hughes, 1968b; Rosemark, tive, adequate diet continues to be a if it met such essential requirements 1978; Good et al., 1982; Eagles et al., major hindrance to commercial lobster for commercial aquaculture as con- 1984, 1986), ground beef and beef liver culture today (Waddy, 1988). Recent sumer demand, profi t potential, ability (Herrick, 1895; Emmel, 1908), shred- and extensive bibliographies on crus- to reproduce in captivity, simple larval ded or crushed fi sh and crab tissues tacean nutrition (Castell and Boston, development, high food conversion effi - (Herrick, 1895; Emmel, 1908; Smith, 1990; Conklin, 1995) will be of great ciency, and resistance to disease (Gates 1933 with H. gammarus; Templeman, assistance to those interested in devel- et al., 1974; Cobb, 1976). Because it 1936), chopped soft-shelled clam (Mead oping artifi cial diets. did meet several of the requirements, it and Williams, 1903; Barnes, 1906a), Broodstock management techniques was selected by the National Oceano- or artifi cially prepared foods (Castell have been developed using preoviger- graphic and Atmospheric Administra- and Budson, 1974; Conklin et al., 1975, ous wild female lobsters (Waddy and tion as one of the four marine animals 1977, 1978; Rosemark, 1978; Capuzzo Aiken, 1984a). By using the indices of having a “high priority” for aquacul- and Lancaster, 1979; Bowser and Rose- Perkins (1972) and Hepper and Gough ture (Glude, 1977; Van Olst and Carl- mark, 1981; D’Abramo et al., 1981; (1978) for embryonic developmental berg, 1979; Van Olst et al., 1980). The Bordner et al., 1986). rates, researchers have been able to major U.S. researchers involved in such Purifi ed diets—HFX CRD 84 con- calculate and control the time of hatch- research were based at San Diego State sisting of crab protein concentrate, ing for H. americanus and H. gamma- University, Bodega Bay Marine Lab- wheat gluten, corn starch, celufi l, cod rus, respectively (Schuur et al., 1976; oratory, the University of California liver oil, corn oil, minerals, and vita- Richards and Wickins, 1979; Beard et at Davis, Woods Hole Oceanographic mins (Boghen et al., 1982) or BML 81S al., 1985; Beard and Wickins, 1992; Institution, the University of Rhode consisting of casein, egg white, wheat Burton, 1992). Furthermore, year-round Island, the University of Maine, and the gluten, corn starch, celufi l, cod liver production schedules for eggs (Waddy Massachusetts State Lobster Hatchery oil, corn oil, soy lecithin, minerals and and Aiken, 1992) and larvae (Waddy (Van Olst and Carlberg, 1979). Private- vitamins (Conklin et al., 1980)—show and Aiken, 1984a, b; Aiken and Waddy, sector research was also conducted by good growth and high survival, while 1985) are now available. Sanders Associates of New Hampshire pelleted diets have had variable suc- Private-sector research has also process. However, pelleted diets incorpo- vided useful information for aquaculture.

19 rating natural food items, such as mysid In 1974, when an ex-lobsterman, Emile Gross, C. 1991. Long Island Lighting Com- 14 pany, 175 East Old Country Road, Hicksville, shrimp, crab, and crangon shrimp have Plante (Chapman ), with a novel pat- NY 11801. Personal commun., 6 Dec. shown some promising results (Cook ented initial habitat design approached 20 Fontaine, C. T. 1991. National Marine Fish- and Worsley, 1986). In contrast, Arte- Sanders Associates of New Hampshire, eries Service, NOAA, Galveston Laboratory, 4700 Avenue U, Galveston, TX 77550. Personal mia ﬂ ake formula does not seem to pro- the company elected to initiate a re- commun., 6 Dec. mote high survivability, but it may be search and developmental program to

61(2), 1999 43 explore the feasibility of lobster cul- cal power generating stations, as orig- commercial production was still con- ture (Anonymous, 1979). Lobsters were inally recommended by Dow (1969), sidered economically unjustifi able. reared for 5 years at Kittery, Maine, Klopfenstein and Klopfenstein (1974), Nevertheless, because of the poten- at accelerated temperatures on special and Shleser and Schuur (1975). tials involved, mathematical models of diets (Hall, 1979). A similar, 2-year pro- Researchers found that market-sized lobster culture facilities were developed gram was conducted in Nashua, N.H. lobsters originally raised from eggs (Rauch et al., 1975; Botsford, 1977). (Chapman, 1983; Chapman et al., 1988) could be grown in 2 years by using These models have been used to proj- to confi rm experimental assumptions thermal effl uent (Van Olst, 1975; Ford ect culture costs, determine accuracy and concepts, develop prototype equip- et al, 1976; Van Olst et al., 1976b; of projections through sensitivity anal- ment, and show the viability of com- Van Olst and Carlberg, 1978). How- ysis, and to determine optimal culture mercial lobster production (Hall, 1979). ever, concerns arose about toxic chemi- methods (e.g. temperature, container Those experimental and pilot programs cals being present in this type of water. size, and fl ow rates) (Johnston, 1976; helped develop a licensable technology Becker and Thatcher (1973) described Botsford et al., 1977, 1978; Johnston package for the American lobster cul- a large number of elements and chemi- and Botsford, 1980). Costs of space, ture (Chapman et al., 1988). cals (e.g. copper, zinc, cadmium, cobalt, land, buildings, tanks, structure, and Unfortunately, commercial produc- chlorine, chromium, lead, arsenic, and trays (Allen and Johnston, 1976), as tion did not proceed for several rea- acids) found in effl uents and their pos- well as waste treatment costs based on sons, including, but not limited to, high sible effects on aquatic life, particu- fl ow rates (Tchobanoglous and Shle- oil prices (which affect construction larly those on growth and fecundity ser, 1974), were assessed using ther- costs of the equipment made of plastic), (Bowen, 1966; Sprague, 1969). Both mal effl uent for aquaculture (Fig. 25) increased wild lobster landings (Anony- compounds and temperature regimes and culminated in a book on the subject mous, 1979), the absence of an artifi cial associated with thermal effl uent were (Allen et al., 1984). diet, and the expense to heat seawater to examined using various life history Others have studied the feasibility of optimal levels (Fig. 25). Nonetheless, stages of the lobster (Dorband, 1975; producing 1,000,000 1-pound lobsters several hundred lobsters were raised to Dorband et al., 1976; Ford et al., 1976, annually, with a general description of marketable size (Chapman14). Methods 1979; Johnson, 1977; Felix, 1978) and a computerized facility (Coffelt and to conserve these costs were investi- were found to be nontoxic and unim- Wickman-Coffelt, 1985). Their projec- gated in the United States and Europe portant in lobster culture. Pilot facilities tions include capital costs in excess of in the form of recirculating culture sys- were then proposed and evaluated for $31 million with an annual operating tems (Hand et al., 1977), solar pow- the commercial culture of lobsters with cost of over $3 million. Thus, pilot-scale ered systems (Portersfi eld, 1982), and effl uent water (Wright, 1976; Turner et projects were suggested and started up the use of cooling waters from electri- al., 1979). Despite these pilot studies, in Provo, Utah (more than 800 miles from the ocean), on the remote Car- ribean Island of Anguilla, in California, and in Hawaii. In Provo, Utah, two Brigham Young University graduate students, Rex Infranger and Roger Mickelsen, developed a program involving the use of artifi cial seawater, solar power, and a system of cages (Mickelsen et al., 1978). They reported successfully raising 1-pound lobsters in 21–30 months by using special diets and temperatures of 22°C (Portersfi eld, 1982). A brief description of their experimental lobster farm and their claim of producing 1,000,000 lobsters a year is presented in Hemming (1981). Currently, an upgraded cage system designed and built based on the original version exists (Mickelsen et al., 1978), but, due to the proprietary nature of this project, information has been limited. However, they operate a functional pilot plant facility Figure 25.—Culture costs using three sources of fuel: fossil fuel, thermal effl uent with everything from hatching to grow- and fossil fuel, thermal effl uent. (Adapted from Botsford et al., 1978). out capability with expectations of mar-

44 Marine Fisheries Review keting their product in the near future Kona Cold Lobster, Ltd. in Hawaii 1966) to Fatty Basin on the west coast of (Infranger21). is currently farming a unique blue lob- Vancouver Island. This was followed by The commercial lobster farm on ster. By mating the H. americanus with the introduction of an additional 5,000 Anguilla, West Indies, applied culture an H. gammarus, the result is a hybrid adults between 1965 and 1966 (Ghe- techniques similar to those of the Uni- lobster that is bright blue in color. Their lardi, 1967). Meanwhile, construction versity of California and the Massachu- product is being marketed to the aquar- began in 1967 to establish a hatchery at setts State Lobster Hatchery, and their ium trade and as a garnish for seafood Fatty Basin, and by May, 123 large “ber- accomplishments are reviewed in Bel- platters in up-scale restaurants (HAAC, ried” females for broodstock were air- leville (1981). 1996). shipped from George’s Bank (Ghelardi Aquaculture Enterprises (AE) spent and Shoop, 1972). These transplanta- 12 years in California experimenting Transplantation Programs tions demonstrated that lobsters would with lobster domestication. Most of their Homarid lobsters are generally lim- grow, survive, reproduce, and behave technical support came from the Sea ited in distribution to the north Atlantic normally in Pacifi c waters (Ghelardi Grant College Programs of the Univer- Ocean. Homarus americanus inhabits and Shoop, 1968, 1972). Barber (1983) sity of California, Davis, the Bodega Bay the northwest Atlantic coast from North reviewed these earlier experiments and Laboratory, the University of Maine, and Carolina to Labrador, while H. gamma- concluded that, with the present knowl- from St. Andrew’s Biological Station in rus inhabits the northeast Atlantic coast edge of the physical parameters (tem- New Brunswick, Can., (Loupe, 1991). from Norway to Morocco (Cooper and perature, dissolved oxygen, etc.) of the AE has taken the concept of “lobster Uzmann, 1980). Prior to heavy com- Masset system, transplanting of lob- farming” from a biological potential to mercial exploitation, H. gammarus also sters to Masset Inlet may be feasible. a demonstrated, economic reality. The occurred in parts of the western Med- During the early 1970’s, California AE cost-effi cient growing system pro- iterranean (Williams, 1988; Holthius, once again attempted to develop an Amer- duces cultured 1-pound lobsters which 1991). Because of these cold-water ican lobster fi shery along its coast (Ford are price- and taste-competitive with wild limitations, transplantation programs and Schuman, 1971; Ford and Krekorian, lobsters. Domestication involved a mul- beginning in the late 19th century were 1972, 1973). However, due to the compet- tidisciplinary approach involving water attempted as a way to increase yields itive interactions between H. americanus chemistry, physiology, pathology, nutri- from the lobster fi shery. This involved and Panulirus interruptus, which showed tion, systems engineering, and repro- transplanting homarid lobsters to the that H. americanus would displace P. duction (Wilson, 1980). AE’s husbandry Pacifi c Ocean, where suitable environ- interruptus, release of wild American and bioengineered technology, combined mental conditions exist (Van Olst et lobsters was not recommended (Kreko- with the discharged sea water from south- al., 1980). Such efforts date to 1873 rian et al., 1974; Lester, 1975). ern California’s Edison Coastal Power and to 1889 for the States of Califor- More recently, Canadian workers Plant, resulted in a 3-year marketable nia and Washington, respectively (Rath- transplanted 2,174 males (81–114 mm lobster, a 70% survival rate, a reproduc- bun, 1892). Nothing resulted from those CL) and 2,310 nonovigerous females tion rate 60% that of wild lobsters, and early attempts, but there are a number of (81–112 mm CL) to St. Michaels Bay in a food conversion ratio of 4.5:1—all for historical reviews on the attempts that Labrador (Boothroyd and Ennis, 1992). less than $3.50 a pound production cost followed (Perrin, 1876; Stone, 1882; While these lobsters were capable of (in 1986 prices) (Anonymous, 1987a). Ryder, 1886a; Smith, 1896), including molting, mating, and reproducing, most In 1988 AE relocated to Hawaii a full account compiled by Rathbun of the female lobsters resorbed their where they now raise lobsters by mixing (1890). In Canada, transplantation of eggs, and those few that did extrude the warm surface waters off Hawaii with lobsters to the east coast of Vancouver were not capable of generating and sup- the deep colder waters, thereby achiev- Island, B.C., was attempted fi rst in 1896 porting a fi shery. ing a 22oC temperature in which the and again in 1905 and 1908 (Fraser, Japan also tried to introduce Ameri- lobsters thrive. AE has also developed 1916). However, since there was no can lobsters into its waters as early as special husbandry techniques incorpo- controlled observation of these animals 1915. Those earlier introductions were rating a plastic grid of their own design after transplantation, no information is not successful, but Jiro Kittaka more (Anonymous, 1990) and their own feed- available on the fate of these lobsters. recently attempted to transplant both H. ing techniques using local fi shing waste These transplantations are reviewed by americanus and H. gammarus (Kittaka, (Loupe, 1991). In 1990, AE had some Butler (1964). 1980; Kittaka et al., 1983). Initial exper- 5,000 animals in residence (Anony- In 1965, several thousand Stage I and iments began with egg-bearing females mous, 1990). Currently, AE is working Stage IV larvae were experimentally of both species from the Massachu- on broodstock strains, testing feeding transplanted from the Massachusetts setts State Lobster Hatchery, the Centre rations, and experimenting with proto- State Lobster Hatchery (Anonymous, Oceanologique de Bretagne and Associ- type production equipment (Wilson22). ation Peche Aquaculture Sud Bretagne in France, and the University College in 22 Wilson, P. L., III. 1990. Aquaculature Enter- 21 Infranger, R. 1992. Sea Inc., 560 South 100 prises, P.O. Box 3314, Kailua-Kona, HI 96745. Galway, Ireland (Kittaka, 1990). Larval West, Provo, Utah. Personal commun., 10 May. Personal commun., 26 Nov. rearing was fi rst attempted with the

61(2), 1999 45 planktonkreisel, but later an outdoor crabs and crayfi sh have become a gour- animals have upon naturally recruiting octagonal tank (1.8 m deep) was used met food in many restaurants, the lob- populations. (Kittaka, 1980, 1990). Stage IV H. amer- ster too can be marketed in this way Great Britain, Norway, and Scotland icanus lobsters were released at Koshiki (Wear, 1990). Research and develop- have demonstrated that hatchery-reared Islands near Kyuski in 1981. This was ment is needed to determine market lobsters do survive after release, are followed by another release in 1982 of demand, system design, control of molt- capable of reproduction, and can 156 Stage IV and V lobsters and 97 ing, and pilot operations, which could enhance existing wild populations. In Stage XI–XV lobsters. These experi- eventually lead to commercial viability. fact, if hatchery-reared juveniles are ments were carried out on artifi cial reefs Furthermore, if lobsters are accepted as released onto carefully selected sub- (cement blocks) and in cages (3 m wide × a soft-shelled product, the possibility of strates, they are capable of adding to the 3m long × 0.3 m high) designed for marketing them at smaller sizes simi- commercial stock within 4 to 6 years monitoring (Kittaka et al., 1983). lar to that of the crayfi sh exists (Wear, following release. Observations revealed that both size 1990). Also, if a market is established These results are encouraging and groups exhibited burrowing behavior for a legal-sized, soft-shelled product, should result in the reassessment of how similar to conspecifi cs in the Atlantic an outlet would then be open for lob- hatcheries can be used in the future— (Kittaka et al., 1983). All H. america- stermen to sell their soft-shelled catch particularly as hatchery functions have nus lobsters dispersed from the release during molting season. shifted from mostly research-oriented site, but 30% of H. gammarus lobsters practices to mostly stock-enhancement remained under their shelters. Further- Conclusions practices. For the future we hope that more, both species successfully repro- Although economics will decide the hatcheries will again pursue rigorous duced in cages and in large pools eventual fate of lobster hatcheries and research programs in addition to their (Kittaka, 1984a). Grow-out was con- stock enhancement programs, much bio- stock enhancements programs, as they ducted in outdoor tanks covered with logical information has already been have a proven research track record and a 10 cm layer of oyster shells on the gained from them. While some hatcher- great potential for further investigations. bottom (Kittaka, 1984a). Lobsters were ies still haphazardly release postlarvae fed natural foods with supplements of into coastal waters without regard for Literature Cited crushed mussels and/or shrimp pellets possible impacts on naturally recruiting Able, K. W., K. L. Heck, Jr., M. P. Fahay, and C. (Kittaka, 1988) and held communally at postlarvae (such as displacement), others T. Roman. 1988. Use of salt-marsh peat reefs by small juvenile lobsters on Cape Cod, Mas- densities of 5–710, with a survival rate are currently providing important infor- sachusetts. Estuaries 11(2):83–96. estimated at 82–89% (Kittaka, 1984a). mation on the survival of recent benthic Acheson, J. M. 1997. The politics of managing Homarus gammarus grew faster than recruits and how long they take to enter the Maine lobster industry: 1860 to the present. Hum. Ecol. 25(1):3–27. H. americanus up to Stage XVII, but H. the fi shery. Such information will not ______and R. S. Steneck. 1997. Bust and then gammarus was smaller in body weight only be useful in predicting year-to-year boom in the Maine lobster industry: perspec- after Stage XII and in total length after fl uctuations in landings, via manage- tives of fi shers and biologists. N. Am. J. Fish. Manage. 17:826–847. Stage XVIII (Kittaka, 1984a). Repro- ment models, but also will provide better Addison, J. T., and R. C. A. Bannister. 1994. ductive size, 540 g, was attained in 4–5 assessments of the value of “seeding” Re-stocking and enhancement of clawed lobster stocks: A review. Crustaceana (Leiden) years (Kittaka, 1984b) and these lob- waters with postlarvae and/or juvenile 67(2):131–155. ster species are currently breeding in lobsters. Acquisition of these data, as ______and M. Fogarty. 1992. Juvenile lobster the local waters of Sanriku. Because of well as the supplying of stock for biologi- habitat limitation: What can landings tell us? Lobster Newsl. 5(2):10–12. the slower growth rates of the European cal experiments, requires the operation of Aiken, D. E. 1980. Molting and growth. In J. S. lobster, it is believed that the Ameri- a hatchery in any district, state, or coun- Cobb and B. F. Phillips (Editors), The biology can lobster will be a more suitable can- try where a commercially important fi sh- and management of lobsters, vol. I, p. 91–163. Acad. Press, N.Y. didate for mass introduction (Kittaka, ery exists. Although hatcheries should ______and S. L. Waddy. 1980. Reproductive 1988). Furthermore, H. americanus is not be used to replace current manage- biology. In J. S. Cobb and B. F. Phillips (Edi- tors), The biology and management of lob- expected to pose no threat to Japanese ment strategies for fi shable wild stocks, sters, vol. I, p. 215–276. Acad. Press, N.Y. littoral organisms, but they may affect they can be used as a stock enhancement ______and ______. 1985. Production of the native shrimp (Kittaka, 1988). research tool, wherever deemed neces- seed stock for lobster culture. Aquaculture 44: 103–114. sary for the health of the fi shery. ______and ______. 1986. Environmental Soft-shelled Product However, any stock enhancement infl uences on recruitment of American lobster The concept of aquaculture is not must be conducted in a responsible (Homarus americanus): A perspective. Can. J. Fish. Aquat. Sci. 43(11):2258–2270. new and it is believed by Wear (1990) manner which will provide the best ______and ______. 1988. Strategies for that methods like those which produce possible outcome for the survival of maximizing growth of communally reared juvenile American lobsters. World Aquacult. soft-shelled crabs may be applied to the stocked and wild lobsters. This may 19(3):61–63. lobster. In the late 1800’s, soft-shelled mean employing and stocking artifi cial ______and ______. 1989. Culture of the lobsters were said to be an excellent reefs in areas currently unsuitable for American lobster Homarus americanus. In A. D. Boghen (Editor), Cold water aquaculture food (Mather, 1894). Recently, it has new benthic recruits, as well as deter- in Atlantic Canada, p. 79–122. Can. Inst. Res. been suggested that just as soft-shelled mining the effect that hatchery-stocked Reg. Develop., Moncton, N.B., Can.

46 Marine Fisheries Review ______and ______. 1995. Aquaculture. In culture venture. Fact Sheet, Sanders Assoc., Atema, J., S. Jacobson, E. Karnofsky, S. Oleszko, J. R. Factor (Editor), The biology of the lob- Nashua, N.H., 2 p. S. Szuts, and L. Stein. 1979. Pair formation in ster, Homarus americanus, p. 153–175. Acad. ______. 1980. Intensive lobster cultivation. the lobster, Homarus americanus: Behavioral Press, N.Y. World Fishing 29(6):61, 63, 69, 71, 73. development, phermones and mating. Mar. ______and W. W. Young-Lai. 1981. Dacty- ______. 1982. New techniques look set to Behav. Physiol. 6:2277–2296. lotomy, chelotomy and dactylostasis methods boost production in lobster fi sheries. World Audouin, M. J. 1974. L’Ecloserie de homards for enhancing survival and growth of small Fishing 31(11):37. de Isle d’Yeu. Int. Counc. Explor. Sea CM lobsters in communal conditions. Aquaculture ______. 1983a. Final environmental impact 1974/K:39, 6 p. 22:45–52. statement and regulatory impact review for ______. 1981. Aspects techniques des eclo- Allen, P. G., and W. E. Johnston. 1976. Research the American lobster, Homarus americanus, series de homards: Production des post-larves direction and economic feasibility: An exam- fi shery management plan. New Engl. Fish. et des juveniles. In Publ. Cent. Natl. Explor. ple of systems analysis for lobster culture. Manage. Counc., Saugus, Mass., 140 p. Oceans, Actes Collogues 12:79–85. Aquaculture 9:144–180. ______. 1983b. American lobster fi shery Bannister, R. C. A. 1998. Lobster (Homarus gam- ______, L. W. Botsford, A. M. Schuur, and management plan. New Engl. Fish. Manage. marus) stock assessment in the United King- W. E. Johnston. 1984. Bioeconomics of aqua- Counc., Saugus, Mass., 48 p. dom: hatchery-reared juveniles do survive in culture. Develop. Aquacult. Fish. Sci., vol. 13, ______. 1986. Final amendment no. 1. Fishery the wild, but can they contribute signifi cantly Elsevier Sci. Publ., Amst., 351 p. management plan for American lobsters. New to ranching, enhancement, and management Anderson, A. W., and C. E. Peterson. 1953. Fish- Engl. Fish. Manage. Counc., Saugus, Mass., of lobster stocks? In L. Gendron (Editor), ery statistics of the United States: 1950. U.S. 70 p. Proceedings of a workshop on lobster stock Dep. Int., Fish Wildl. Serv., Sect. 2:56–98; ______. 1987a. From concept to consumer: enhancement held in the Magdalen Islands Sect 3:146–176. The technical evaluation of lobster farming. (Quebec) from Oct 29–31, 1997, p. 23–32. Anonymous. 1873. Culture du homard en Ame- Aquacult. Enterprises, Kailua- Kona, Hawaii, Can. Ind. Rep. Fish. Aquat. Sci. 244. rique. Bull. Soc. d’Acclimattation 10:957– 10 p. ______and A. E Howard. 1991. A large 960. ______. 1987b. Final amendment no. 2 to the scale experiment to enhance a stock of lob- ______. 1874. Notes. Nature 10:214. American lobster fi shery management plan. sters, Homarus gammarus L., on the English ______. 1892. Lobsters. 20th Annu. Rep. New Engl. Fish. Manage. Counc., Saugus, east coast. Int. Counc. Explor. Sea, Mar. Sci. Comm. Fish., State N.Y., p. 69–73. Mass., 30 p. Symp. 192:99–107. ______. 1895. Annual Report of the New- ______. 1988. Guide to Cutler Marine Hatch- ______, ______, J. F. Wickins, T. W. foundland Department of Fisheries for the ery. Hatchery Pamphl., Cutler, Maine, 2 p. Beard, C. A. Burton, and W. Cook. 1989. A year 1894, 134 p. ______. 1989. Final amendment no. 3 to the brief progress report on experiments to evalu- ______. 1899. The lobster. 4th Annu. Rep. American lobster fi shery management plan. ate the potential of enhancing stocks of lob- Comm. Fish., Game, For., State N.Y., p. New Engl. Fish. Manage. Counc., Saugus, sters, Homarus gammarus L., in the U.K. Int. 290–293. Mass., 21 p. Counc. Explor. Sea. CM 1989/K:30:1–8. ______. 1906. 6th Biennial report, State Com- ______. 1990. Aquaculture report. Seafood ______, B. M. Thomas, J. T. Addison, S. missioners of Fisheries and Game for the Leader 10(5):119. J. Lovewell, and A. E. Howard. 1990. The years 1905–1906. State Conn., Public Doc. ______. 1991a. Research and development 1989 results from a lobster stock enhancement 19:21–34. program for 1991–1992. Sea Fish Ind. Author- experiment on the east coast of England. Int. ______. 1909. Lobsters. Annu. Rep. For., ity, Mar. Farm. Unit, Ardtoe, Scotl., 4 p. Counc. Explor. Sea. CM 1990/K:13:1–8. Fish, Game Comm. State N.Y. 1907–1908– ______. 1991b. Lobster stock enhancement. ______, B. M. Thompson, J. T. Addison, S. 1909:295–298. Sea Fish Ind. Authority, Mar. Farm. Unit, R. J. Lovewell, and A. R. Lawler. 1991. The ______. 1912. Annual report of the fi sh cul- Ardtoe, Scotl., Inform. Sheet, 4 p. 1990 results from a lobster stock enhancement turist. 1st Annu. Rep. Conserv. Comm., State ______. 1991c. Do lobster hatcheries really experiment on the east coast of England. Int. N.Y., p. 168, 172, 175. work? Lobster Bull. 4(1):2. Counc. Explor. Sea CM 1991/K:33, 5 p. ______. 1919. New method in lobster culture. ______. 1991d. Final amendment no. 4 to Barber, F. G. 1983. Lobster transplant to Masset 8th Annu. Rep. Conserv. Comm. 1918. State the American lobster fi shery management plan Inlet. Can. Tech. Rep. Fish. Aquat. Sci. 1181, N.Y., Legisl. Doc. 54:84–85. incorporating an environmental assessment 6 p. ______. 1920. Cold Spring Harbor hatchery. and regulatory impact review. New Engl. Fish. Bardach, J. E., J. H. Ryther, and W. O. McLar- 9th Annu. Rep. Conserv. Comm. 1919. State Manage. Counc., Saugus, Mass., 22 p. ney. 1972. Lobster culture. In Aquaculture: N.Y., Legisl. Doc. 83:77. ______. 1994. Amendment #5 to the Amer- the farming and husbandry of freshwater and ______. 1930. 18th Biennial report, State ican Lobster Management Plan: Incorporat- marine organisms, p. 630–649. Wiley Inter- Board of Fisheries and Game for the year ing a draft supplemental environmental impact sci., N.Y. 1930. State Conn., Public Doc. 19:24–29. statement. Vol. 1, New England Fishery Man- Barnes, E. W. 1906a. Methods of protecting and ______. 1936. 9th Biennial report. Maine Dep. agement Council, Saugus, Mass., 129 p. propagating the lobster, with a brief outline Sea Shore Fish., p. 29–33. ______. 1995. Lobster stocking: progress and of its natural history. Annu. Rep. R.I. Comm. ______. 1955. Biennial report, State Board of potential. Signifi cant results from UK restock- Inland Fish. 32:120–152. Fisheries and Game for the years 1954–1955. ing studies 1982–1995. MAFF Directorate ______. 1906b. The propagation of lobster State Conn., Public Doc. 19:214. Fish. Res. Lowestoft, 12 p. fry for the purpose of increasing the supply ______. 1963. Division of Marine Fisheries ______. 1996. Finding new homes for lob- of lobsters in the waters of the state. Methods annual report for 1962–1963. Commonwealth sters. Nor’Easter vol. 8, no. 1, Spring/Summer of artifi cial propagation and cultivation. R.I. Mass., Dep. Nat. Resour., Div. Mar. Fish., 1996. Comm. Inland Fish. Annu. Rep. 36:111–119. Annu. Rep. 500-10-63-355:17–28. ______. 1997. Right whales: lawmakers fi ght ______. 1907. Lobster culture at Rhode Island ______. 1964. Division of Marine Fisheries new regs on lobster fi shers. Bangor Daily in 1906. R.I. Comm. Inland Fish., Annu. Rep. annual report for 1963–1964. Commonwealth News, Assoc. Press, 10 April 1997. 37:88–94. Mass., Dep. Nat. Resour., Div. Mar. Fish., ______. 1998. Draft environmental impact ______. 1939. An analysis of the objectives Annu. Rep. 500-10-64-402:11, 19–33. statement (DEIS) and regulatory impact of lobster rearing and problems of reinvigo- ______. 1965. Division of Marine Fisheries review. Federal lobster management in the rating the lobter industry. In Special report annual report for 1964–1965. Commonwealth Exclusive Economic Zone. National Marine of the development of conservation relative Mass., Dep. Nat. Resour., Div. Mar. Fish., Fisheries Service, Northeast Region, 104 p. to the feasibility and cost of propagation of Annu. Rep. 500-1-65-499:8, 18–30. Appelløf, A. 1909a. Undersokelser over humme- lobsters by the Commonwealth of Massachu- ______. 1966. Division of Marine Fisheries ren (Homarus vulgaris) med saerkskilt, hensyn setts, Appendix A, p. 10–32. Mass. House annual report for 1965–1966. Commonwealth til dens optraeden ved Norges kyster. Aars- Doc. 2051, State Library, Boston, Mass. Mass., Dep. Nat. Resour., Div. Mar. Fish., beretning Vedkommende Norges Fisk.:1-185. Barshaw, D. E. 1989. 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61(2), 1999 47 with other substrates. Mar. Ecol. Prog. Ser. (Homarus gammarus). Univ. Rennes, France, ______, ______, and ______. 1992. 106:203–206. These, 250 p. Infl uence of a thermocline on vertical ______and D. R. Bryant-Rich. 1988. A long- ______, J. Y. Gauthier, and J. Lorec. 1985. distribution and settlement of postlarvae of term study on the behavior and survival of early Variations de la duree de la phase pelagique the American lobster Homarus americanus juvenile American lobster, Homarus america- des post-larvaes de homard Europeen, Homa- Milne- Edwards. J. Exper. Mar. Biol. Ecol. nus, in three naturalistic substrates: eelgrass, rus gammarus, en function de la nature des 62(1):35–59. mud and rocks. Fish. Bull. 86:789–796. fond. Int. Counc. Explor. Sea CM 1985/K:22, Bowbeer, A. 1971. The underwater lobster farm— ______and K. L. Lavalli. 1988. Predation upon 15 p. another man’s approach to live lobster supply. postlarval lobster, Homarus americanus, by Bishop, F. J., and J. 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