LOBSTERSERIES COVER PHOTO: Unidentified fisherman, with taken from a topentry woodenslat trap, Over 90%%uo of the spiny lobster landings are taken in thismanner. Picture from Gary E. Daois.f

TABLE OF CONTENTS

Foreword Gary E. Davis 27 Florida'sSpiny Lobster Fishery: An Historical Perspective RonaldLabiskv, Douglas R. Gregory,Jr., and Joseph A, Conti 28 Review of the Horida Spiny Lobster Resource David C. Simmons 37 The LarvalRecruitment Problem of theSpiny Lobster RobertA. Menziesand J. MichaelKemgan 42 The PostlarvalStage of ScyllarideanLobsters WilliamG, Lyons 47 LobsterFishery Management under the Fishery Conservation and ManagementAct James R. Zuboy, A. C. Jones, and T. J. Costello 50 EconomicConsiderations in the Managementof the Horida Spiny Lobster Fishery Fred J. Prochaska and James C. Cato 53 JuvenileSpiny Lobster Gary E. Davis 57 Spiny LobsterCulture: An Alternativeto Natural Stock Management George R. Tamm 59

American FisheriesSociety 5410 Grosvenor Lane Bethesda, Maryland 20014

FLORIDA'S SPINY LOBSTER FISHERY: AN HISTORICAL PERSPECTIVE

Ronald F. Labisky, Douglas R. Gregory, Jr., and Joseph A. Conti

ABSTRACT

The Fiorida-basedfishery accounts for 98 percent of the spiny landed cornmerdaiiy in the ; this catch is comprised almost exclusively of the Horida spiny lobster, Panu1irusargus. The commercial lobster industry began in the lower Florida Keys during the early 1800's, principally as a bait fishery that supported the local finfish industry. Subsequent devel- opments in the lobster fishery have essentially mirrored the prosperity of Key West. In 1908, landings totaled about 53,000 pounds,valued at $3,600.The fisher beganto exhibitprogressive expansion in the 1940's,a trendthat continuedthrough the 1970's. A sport diver! fishery for tobsters, which began in the 1950's, has also gained prominence. Coinmercial landings in Florida, induding catchesfrom international waters,peaked in 1972 at 11.4 million pounds, The peak in ex-vessellandings value, $13.4million, was attained in 1974.The strong international fishery, which developed during the 1960's,was severely curtailed in 1975 by the dosure of the Bahamian fishing grounds to foreign fishing. Domestic catchesfrom Florida waters dechned slightly during the 1970's, despite a substantialincrease in fishing effort. Retrospectively,the spiny lobster has not only strongly impacted the culture, sociology, and economics of south Fiorida, but has also emerged to rank second only to as Florid's leading fishing industry.

The Florida spiny lobster is a decapod crustaceanbelonging to the family Palinuridae, which contains 8 extant genera with 48 speciesthat are principally tropical and subtropical in distri- bution George and Main 1967!. These palinurids support major fisheriesworldwide the five top-ranked being Australia, South Africa, Brazil, Cuba, and the United States, respectively NMFS 1974; Streeter and Weidner 1976!. In the latter three western-centralAtlantic fisheries,P. argus, an inhabitant of the shallow continental shelf waters from North Carolina to Brazfi Williams 1965!, is the mainstay lobster species,accounting for 90 percent of the commercial catch Chekunova 1972!. In the United States, 98 percent of the commercial catch of spiny lobsters is taken by Florida-based fishermen Williams and Ronald Labisky Douglas R. Gregory, Jr. Prochaska 1977!. And the bulk of this catch, comprised exclu- sively of P. argus, comes from habitats associated with the species of , Florida Keys Prochaskaand Cato 1974! Fig. 1!. A ! growing to the In terms of ex-vesselvalue, the spiny lobster fishery in Florida weight of five pounds a piece, is second only to the shrimp industry Prochaska 1976a!. To live in great numbers in the date, the high in lobster landings, 11.4 million pounds, was holes of the coral rocks, on the reached in 1972, and the high in ex-vesselvalue, $13.4 million, mangrove islands: these in 1974. Even though these figures include lobsters taken by shrimps are by west-Indians Florida fishermen from international waters, they are in marked improperly called lobsters, al- contrast to the 53,000 pounds of lobsters valued at $3,600 though they have not the two landed in Florida in 1908 Crawford and De Smidt 1922!. claws, as lobsters: they are Man's discriminatory taste buds have vaulted the lobster to beautifully spotted with red, the lofty status of an epicurean delight. Consequentiy, the ac- yellow, blue, qreen, grey, and celerated popuhrity of the lobster has increased exploitation a little black; but theV all JosephA pressures on the world's lobster stocks. Nowhere is this more change into one red colour by true than in Florida. Technological advances and increased boiling..., " capitalizationwithin the spiny industry of FIorida, So descriptively, but mistakenly, wrote William Gerard De as in the fishery for the , Homarus americanus, Brahm 772, p. 11!, who had been appointed surveyor gen- in the North Atlantic Cobb 1976!, have placed heavy pressures eral of the British colony of East Florida, which had been re- on the resource in recent decades. In addition, these pressures cently acquired from the French and Spanishas a prizeof war in on the domesticstock have been magnified by the closure of the the Peace of Paris pact of 1763. The speciesreferred to by De previously available Bahamian fishing grounds. Now, declining Brahm was undoubtedly the spiny lobster, argus, landings have not only reduced the profitability of spiny lobster commonly called the Caribbean or Florida spiny lobster or, fishing but also created concern as to the future stability of the among the Florida lobstermen, crawfish. domesticfishery itself. This paper tracesthe development of the

Fisheries, Vol. 5, No, 4 market for fresh fish inaintained a strong finfish industry in the ing lobster fishing was enacted by the State of Florida in 1919; rich watersof the Florida Keys, and, consequently,a lobster bait its intended purpose, as would be expected, was to conserve fishery. the supply of lobsters.Consequently, a closedseason was estab- Cast nets, gill nets, haul seines, and grains - or 3-tined lished between March 1 and June 1, but with provisions that barbed or barblessspears! were the most frequently used gear allowed the taking of lobstersfor fish bait, research,or propaga- for catching lobsters in the 1800's Brice 1898; Crawford and tion throughout the year Schroeder 1924!. In 1921 the closed De Smidt 1922!, Cast nets, constructed of woven twine with a season was changed to the period March 21 to June 21. diameter of 12-16 feet and fringed with lead weights, consti- Spiny lobster production increasedslowly during the 1920's tuted a popular and effective gear-type. Gill nets and seines Table 1!, presumably in response to the growing retail trade, were used primarily to take lobsters while they were scattered The fishery, which until this time had been supported by during feeding forays on the shallow grassflats. lobstersinhabiting shallow waters immediately surrounding Key The profitability of this early lobster fishery in Florida is con- West, was promoted to expand geographically and technologi- jectural. In 1895, lobsterssold at $0.02 per pound Brice 1898!; cally.By 1922the principalfishing grounds had beenextended thus, a daily catch of 100-200 pounds may have produced a to a 25-mile linear zone that encompassedthe southern shores handsome income for skilled lobstermen, who had but little of the lower Florida Keys and the shallow Atlantic reef areas capital invested in fishing gear and boats. both eastand west of Key West Crawford and De Smidt 1922!. The late 1880's chronicled an abrupt decline in the prosperity The geographicrange of the fisherywas probablydictated by of Key West. Sponge die-offs in 1879, coupled with apparent the limited capabilitiesof the then common sail-powered fishing overfishing, signaled a decline of the sponge fishery in the boats Fig. 2!. Those few fishermenwith gasoline-powered Florida Keys Collins 1887!. The cigar industry became flnan- boats were able to venture somewhat faither to Bahia Honda ciallv plagued by labor disputes Browne 1912!; its relocation to in the east and to Marquesas and the Dry Tortugas in the Tampa was underway by 1886 Thurston 1972!. And strained west but the overall small size of the boats reshicted their use political relations between the United Statesand Cuba precipi- to periods of favorable weather Crawford and De Smidt 1922!. tated Cuba's enactment in 1868 of a prohibitary duty on fresh Fishing technology was also changing. Bully nets Fig. 3!, fish imported from the United States. This enactment had by which had largely replaced the seines,nets, and grains of earlier 1882 taken the vigor out of Florida's finfish industry and its years, accounted for about 50 percent of the catch of spiny supporting spiny lobster bait fishery Thurston 1972!, Thus. the lobsters in the 1920's Crawford and De Smidt 1922!. This 1800's saw the birth, but little substantivegrowth, of the spiny change in fishing technology was precipitated by two factors: lobster fishery in Florida, first, increasingexploitation had reduced the number of lobsters that could be found in the shallow waters where seines and nets had been previously used effectively; and second, lobsters THE EARLY l900's speared with grains spoiled rapidly, and therefore had a lower The early 1900's marked a continuation of the decline in Key market value Crawford and De Smidt 1922!. Thus, the bully West's economy and population. The relocation of the sponge net became king. To deplov the bully net, a fisherman stood in fisheryto TarponSprings and the cigarindustry to Tampahad the bow of an open skiff that he pushed slowly with the long deprived Key West of its two principal industries Tebeau handle of the net until he spied a lobster or group of lobsters, 1971!. Although spiny lobsters maintained soine importance in whereupon he inverted the net and scooped up his catch. The the local retail markets, landings during the first decade of the bully net was effective to depths of about 2 fathoms Smith centuryremained low becausethe fisherywas still largelyde- 1948!. pendent on the finfish fishery, at this time principally an opera- Fish traps were the second most popular gear-type for taking tion that supplied only Key West. Other concurrent devel- spinylobsters during the 1920's,accounting for about 20 per- opments, however, were beginning to stimulate the spiny lobs- cent of the catch Crawford and De Smidt 1922!. These traps, ter fishery.The constructionof the OverseasRailroad in 1912 constructed of wire, measured about 36 x 24 x 20 inches. enabled dealers to experiment with shipment of spiny lobsters Trap fishermen usually employed only about 15 traps and to northern restaurants and hotels. At first, lobsters were checkedthem daily, The obvious advantagesof traps over bully shipped alive in barrels among alternating layers of sponge nets were that traps could be fished in deeper water and during clippings and ice. This inethod proved cumbersome and moderately bad weather, whereas bully nets required calm and costly; careful handling was required from the time of capture shallow water for visual sighting of the lobsters Schroeder until the lobsters reached their destination. The shipping of 1924!. cooked lobster tails, packed in ice, proved inore successful; That the spiny lobster fishery did not show progressive although the cooking processwas more labor intensive, rates of strength throughout the 1920's and 1930's was probably due to spoilage were much less for cooked tails than live lobsters the persistent economic turmoil that plagued Kev West, Al- Schroeder 1924!. By the late 1910's, the estimated annual though the early 1920's saw the entrance of fast-inoney enter- catch of lobsters was about 350,000 pounds; 40 percent of this prises in south Florida both a land boom and a spectacular catch was shipped, 40 percent was consuined locally, and 20 increasein rum-smuggling as a result of prohibition in 1920 percent was used as fish bait Schroeder 1924!. The commer- - Key West's economy was relatively unaffected and the island cial retail trade for spiny lobsterscontinued to expand and was city's population actuallv declined Tebeau 1971!. By the end firmly establishedbv 1920, as indicated by the construction of a of the 1920's spiny lobster production had increased to only large ice-making and cold storage facility in Key West, Markets about 900,000 pounds Table 1!. Florida's lobster regulations for spiny lobsters had been establishedin Miami, Jacksonville, were amended in 1929; the closedseason was lengthened from Atlanta, Philadelphia,New York, and Boston Schroeder 1924!. 3 to 4 months, March 21 to July 21; and for the first time, a The growing lobster fisheiy apparently stimulated concern for minimum legal sizelimit of 1 pound was established Prochaska the protection of the resource.The first legislationdirectly affect- and Baarda 1975!.

30 Fisheries Vol 5 No 4 Table 1, Landings,ex-vessel economic value, and fishing elort in the spiny lobster fPanuiltrusargus! fishery in Rorida, 1925-79.'

Pounds Price per Number Number Traps per landed Value pound of boats of traps boat or Date ,000's! $1,000's! $! or vessels ,000's! vessel

1925-26 88 $7 $ .08 1927-28 817 65 .08 1929-30 873 79 .09 1931-32 1,342 107 .08 1933-34 266 13 .05 1935-36 832 50 .06 1937-38 643 45 .07 1939 405 28 ,07

1940 400 28 .07 1941 2,087 104 .05 1942 1,673 117 .07 1943 1,911 229 ,12 1944 1,176 188 .16 1945 794 159 .20 1946 1,586 317 .20 1947 4,891 978 .20 1948 3,954 830 .21 1949 3,581 788 .22

1950 1,560 281 .18 376 12 31 1951 3,097 589 .19 319 17 53 1952 1,612 403 .25 102 15 146 1953 1,995 399 .20 147 26 175 1954 1.947 428 .22 157 30 194 1955 2,295 528 .23 166 39 235 1956 3,113 825 .27 128 33 256 1957 4,040 1,124 .28 161 37 228 1958 2,954 837 .28 187 34 184 1959 3,181 955 .30 254 52 204

1960 2,849 1,100 ,39 221 74 333 1961 2,803 969 .35 195 52 268 1962 3,107 1,187 .38 248 75 300 1963 3,583 1,408 ,39 246 80 326 1964 3,6318!' 1,562 341 114 333 1965 5,7147! 3,219 .56 332 139 418 1966 5,3501! 2,469 ,46 488 151 309 1967 4,4147! 2,733 .62 628 186 352 1968 6,1553! 4,409 .72 452 168 373 1969 7,5816! 5,258 .69 440 165 374

1970 9,8692! 5,918 .60 492 219 262 1971 8,2063! 7,057 .86 520 226 434 1972 11,4178! 11,771 1.03 599 248 415 1973 11, 1723! 11,661 1.04 671 280 417 1974 10,8749! 13,382 1.23 690 371 538 19754 7,4087! 9,863 1.33 823 520 632 1976 5,316 9! 8,586 1.60 1977 6,3448!' 10,425 1,64 1978 4,5855!' 9,610 2. 10 1979 5,9502!' 11,705 1.97

'Catchand effortstatistics derived from the followingsources; Dawson and Idyll 1951;NMFS 1974; and unpublishedstatistics from NMFS, Southeast FisheriesCenter, Miami, Florida. 'Statistics not available. 'Numbers within parenthesesrepresent the percentage of landings comprised of lobsters caught in international waters. Bahamian continental shelf dosed to foreign fishing in 1975. 'Prehminary estimates. Spiny lobster production in south Florida actually declined during the 1930's, averaging only about 700,000 pounds an- nually Table 1!, History reveals that this slowing of the spiny lobster fishery was inevitable. First, the Great Depression se- verelycurbed the retailmarket for the spinylobster, which al- readyhad the statusof a luxuryfood in the North.Second, the city of Key West was in the throes of economic disaster;in 1934, the Federal Emergency Relief Administration a New Deal agency! intervened for purposes of rehabilitating the commu- nity Tebeau1971!. Third, the OverseasRailroad, Key West's only direct link to the Floridainainland, was irreparablyde- stroyed by the Labor Day humcane of 1935. The latter catas- trophe provided the impetus for the construction of a new roadway, the Overseas Highway, in 1938 Tebeau 1971!, Thus, a direct transportation link between the Florida mainland and Key West had again been established,and with better eco- Figure 3. Bully-netting for spiny lobsters. The bully net, fished nomic times ahead, a new opportunity for the spiny lobster froina smallskiff in shallowwater, was the predominantgear type industry. used to catch lobsters during the formative stages of Florida's fishery. THE 1940's: AN EVOLVING INDUSTRY The commercial flshing grounds for spiny lobsters during the 1940's expanded markedly extending from Palm Beach on World War II revitalizedKey West. The Navy reactivatedand Florida's east coast, throughout the Keys, and southwestward expanded its military base; more significantly, it constructed a toward the Dry Tortugas Dawson and Idyll 1951!. Within the water pipeline along the new OverseasHighway between the Horida Kevs, still the major grounds, fishing was concentrated Horida mainland and Key West to provide the city with a de- principally along the paralleling offshore reef in the Atlantic. pendable supply of fresh water Tebeau 1971}. Population and The commercial flshing gear used most cornmonlv in the prosperitv were reborn in the United States' southernmost city! 1940's,in orderof popularity,included wooden slat-traps, bully The spinv lobster fishery of south Horida evolved into a via- nets, and ice-can traps. The wooden slat-trap Fig. 4!, which has ble industry during the 1940's. The decade saw the catch of becomethe mainstaycommercial gear-type, was probably in- !obster double and the ex-vesselprice triple; the high point of troduced into the Keys fishery about 1940 Irvin Knowles 1980, the decade occurred in 1947 with a catch of nearly 5 million personal communication!. The slat-trap, 1 5 x 2 x 4 feet in pounds, valued at nearly $1 million Table 1!. Undoubtedly, the dimension. was constructed of wooden laths spaced about 1.5 lobster fishery benefited from the flourishing national economy inches apart; the entrance, guarded bv downward projecting of both the war and post-war years. laths. was in the top of the trap, Each trap was weighted with

Figure2. The earlyinshore fishing fleet at KeyWest fromSchroeder 1924!. These sailboats, called smackees, and skils were used in fishing for sponges,spiny lobsters,and finfish,

32 Fisheries, Vo!. 5, No. 4 stonesor concrete. Baited with fish heads,the traps were gener- afly set in 3-7 fathoms of water; each trap was deployed individ- uallv, and marked by line and float. A crew of two or three men, using conventional gasoline-powered boats up to 30 feet in length could fish 100-400 traps in total, but could work the traps at a rate of only about 100 traps per day becausethe pulling of traps had to be done manually. A good day's catch totaled 400-500 lobsters, or about 600-700 pounds Smith 1948!. The bully net, the principal gear of the early 1900's, was restncted in use to lobstermen who either fished in the shallow waters of Florida Bay or could not afford the capital outlay needed for slat-tmp fishing or both. Bully netters, although catching only 100-200 pounds of lobstersper day, did not have the vessel,bait, and trap expensesassociated with slat-trap fish- ing Smith 1948!. The ice-can trap emerged as an interesting gear-type in Horida during the 1940's Smith 1948!. These traps were con- structed from discardedrectangular steel tanks, about 30 x 9 x 48 inches in size, that had been used in ice-making. The single opening of the rectangularcan was collapsedin such a way as to leave two triangular entmncesfor lobsters.Lobsters entered the cans for the shelter they offered; consequently, these traps did not require bait. Generally worked bv lobstermen at 1- to 2-week intervals, can traps occasionally yielded a catch of as many as 100 lobsters. The weight of these can traps, however, limited their use to shallow waters -2 fathoms!. Each type of fishing gear was used for a specific purpose. Baited slat-trapswere employed primarily in the deeper waters associatedwith the offshore reef on the Atlantic side of the Keys; bully nets were used in the shallow waters of Florida Bay; and unbaited ice cans were used in shallow inshore waters. Traps baited with fish heads were ineffective in Florida Bay because Figure 4. The wooden slat-trap. First introducedinto south Florida the fish heads attracted stone Menippe mercenania!, in the 1940's, the slat-trap has become the dominant fishing gear which, in turn, destroyed or maimed any lobsters captured in for spiny lobsters in Florida. the traps Irvin Knowles and Troy Baker 1980, personal com- lobster fishing as an occupation Trov Baker 1980, personal municaiion!. communication!. The market for spiny lobsterscontinued to expand, enhanced The increased fishing effort for spiny lobsters in the 1950's greatly by the advent of deep-freeze processing techniques stimulated further changes in harvest regulations. In 1953 the Smith 1948!. Delivered live to the fish buyers, lobsters in the Florida legislature shifted the chronology of the closed season best condition were sold to local retail markets and restaurants from the period March 21 to July 21 to the period April 15 to in the Keys. The bulk of the catch, however, was boiled, and August15, andredefined the legalsize limit from a minimumof then frozen for subsequentshipment. The 1940's also marked a 1 pound to a minimum tail measurementof 6 inches Prochaska surge in the foreign import of spiny lobster tails into the United and Baarda 1975!. The purpose of this size change was to States principally from the Caribbean, South African, and enable fishermen and law enforcement personnel to identify Australian fisheries Chace and Dumont 1949!, an event that legal-sizedlobsters more easily,Again, in 1955, the closed sea- signaled economic competition. Further expansion of the son was changed to the period March 31 to August 1. The Florida lobster fishery was on its way. rationale for the numerous changesin the timing of the 4-month closedseason must have been predicatedon politics rather than THE 1950's: GROWTH biology becausethe protectionof the breedingstock would have not changed appreciably within the outside limits of these The population of the Florida Keys Monroe County!, which calendar dates. had doubled during the preceding decade 14,000 in 1940 Throughout the 1950's, wooden slat-traps, baited with fish and 30,000 in 1950 reached 48,000 by 1960 Mathis et al. heads, canned sardines,or pet food, continued to be the most 1979!. This rapid growth was probably due to the large number prevalent commercialgear Cope 1959!. Ice-can traps were still of military familiesstationed in Key West druing World War II, to used, though Siebenaler 952! mentions that these traps were post-war prosperity, and to an expanding tourism industry. illegalbecause they capturedmany sublegal-sized lobsters. The The spiny lobster fishery experienced similar growth during bully net continued to decline in use during the 1950's the 1950's. The number of boats and vesselsdeployed in the Siebenaler 1955; Cope 1959!. However, a new lobster pre- fishery more than doubled, from 102 in 1952 to 254 in 1959, dator appeared on the Florida scene during the 1950's skin and the number of traps fished more than tripled, from about and scuba divers equipped with gigs and spears Cope 1959!, 17,000 in 1951 to about 52,000 in 1959 Table 1!. The catch which were then legal. This event marked the beginning of the and ex-vesselprice of lobsters also increased. The combination underwater sport fishery that today competeswith the commer- of a warm climate and an attmctive lobster market encouraged cial fishery for the lobster resource Beardsley et al. 1975!. many Navy retirees to remain in the Florida Keys and to adopt The major fishing grounds in the 1950's were still centeredon

July August 1980 33 the southern or reefsideof the Florida Keys in depths from 3-15 cowhide was lessexpensive, longer lasting, and easierto store. fathoms Cope 1959!. An experienced crew of two men was Furthermore, slat-traps baited with cowhide attracted lobsters, able to work as many as 150 traps per day Cope 1959!; the but not predatory stone crabs,and thus could be fished success- traps were usually pulled and rebaited every 2-12 days fully for the first time in Horida Bay Troy Baker and Irvin Siebenaler 1955; Cope 1959!. When resettingthe trap, one or Knowles 1980, personal communication!. With the inclusion of more live lobsters were commonly left in the trap as a lure; in Florida Bay, the fishable area for spiny lobsters in south Horida addition to the standard bait Cope 1959!. As a rule of thumb, a nearly doubled in size. catch of 3 pounds of legal-sizedlobsters per slat-trap per week Both the number of boats/vessels and the number of traps in was considered satisfactory. the fishery essentiallydoubled during the 1960's Table 1!. Not The boats used in the lobster fishery during the 1950's only were traps added to the fishery by incoming fishermen,but rangedfrom 14-footskiffs, usually propefied by outboarden- the number of traps used by each fisherman also increased. gines,to 36-foot motor launches Cope 1959!, The typical Concurrently, both the catch and the ex-vesselprice of lobsters commercial boat was wooden hulled, 26-36 feet in length and also doubled. The catch in 1969 totaled 7.5 million pounds of shallow draft!, and powered by diesel, marine gasoline, or valued in excess of $5 million Table 1!. Interestingly, the converted automobile engines. Although diesel engines, be- poundage of lobsters taken per fished trap reflected no consis- causeof lower fuel and maintenancecosts, were preferred, their tent upward or downward trend. high initial purchaseprice steeredmost fishermento gasoline Throughout the 1960's, fishermen were switching to larger engines. craft from boats lessthan 5 tons to vesselsgreater than 5 tons! The slat-trapswere commonly pulled by hand or hand-lifted at a rapid pace Williams and Prochaska1976!, This increasein with davits. However, Cope 959! reported that some fisher- .boat sizeappeared to be associatedwith the development of the men used a gypsy head driven by a power takeoff from the commercial fiberglassboat-building industry in the lower Rorida main engine or by a separatesmall gasoline engine to haul their Keys Troy Baker 1980, personal communication!. The modern traps aboard the boat; the latter, of course, greatly increasedthe lobster fishing vesselis of fiberglassconstruction, 30-55 feet in number of traps that could be fished per boat. length, and diesel powered Fig. 5!. Williams and Prochaska The 1950's were a period of growth for the spiny lobster 976! postulated that the demand for larger and faster vessels fishery,stimulated primarily by favorablemarket prices and an was influenced by the availability of lobsters on offshore fishing influx of additional fishermen, including many Navy retirees. grounds probably referring to the Bahamas!. By 1967, only Looking ahead, the 1960's were to become a decade of con- about 43 percent of the lobsters landed in Florida were domes- tinued growth, characterized by additional fishermen, ticallv harvested,the remainder coming from Bahamian waters. technologicalprogress, explosive expansion of fishing grounds, Again, the increase in fishing effort precipitated changes in and development of a south Florida-basedinternational fishery. lobster fishing regulations, In 1965 legislation was enacted to define the minimum size limit further, to protect egg-bearing females,and to regulate gear. The minimum sizelimit was rede- THE 1960's: MORE GROWTH fined as a carapacelength measurementof 3 inches and a tail The decade of the 1960's was a period of radical change in length of 5.5 inches. The taking, or stripping, of egg-bearing the south Florida spiny lobster fishery. Thousands of Cuban females was prohibited. Traps and buoys were required to be immigrants entered the fishery; the average ex-vesselprice per not only permanenfly marked with the owner's lobster permit pound nearly doubled; technologicalimprovements occurred in number, which was obtained at no cost, but also color coded; boat design, engine systems, electronic navigation, depth re- this effort appeared to be aimed at discouragingtrap theft and corders, and trap-pulling systems; domestic fishing areas ex- poaching. Also, an import permit was established to allow panded; and an international fishery developed. Florida fisherman to land lobsters captured in international Some 300,000 Cuban immigrants, fleeing Fidel Castro's waters. newly established government, setfled in Florida during the Thus the lobster fishery of the 1960's was characterizedovert- 1960's Tebeau 1971!. Many of those immigrants who had ly by growth and prosperity. The upcoming 1970's, however, been fishermen in their home country tumed to fishing once would chronicle more troubled times for this crustaceanfishery, they were in Horida. The United States government provided subsidiesand other services,via the Cuban Alien Act, to expe- THE 1970's: UNCERTAIN TIMES dite the settlement of the thousands of new resident aliens. One importantaspect of thiqaid was the exemption for Cubans from In 1971, it became legal to land spiny lobsters from intema- the restrictionson fishing-vesselownership by aliens Holmsen tional waters in Florida during the state's closed season 1972!, Many Miami-basedimmigrant fishermen tumed to lobs- Prochaska and Cato 1974!. International catches by Florida ter fishing, and primarily concentrated their efforts on the al- lobstermen comprised nearly half of the lobsters landed during ready familiar Bahamian fishing areas Holmsen 1972; Sim- the early 1970's, despite a loss in the availability of some tradi- mons 1974!. tional foreign fishing grounds. Bahama's expansion of its As the ex-vesselprice of lobsterscontinued to increase,more sovereignoffshore fishing zone from 3 to 12 miles in 1969 lobstermen were able to obtain diesel engines,larger boats, and forced many Rorida fishermento turn to the more distant electronic equipment. During the early part of the decade, hy- Caribbean fishing grounds, By 1972 as many as 50 Florida- draulic systems were developed for pulling traps Troy Baker based freezer boats were fishing for lobsters in the Caribbean 1980, personal communication, This mechanical innovation, Holmsen 1972!. Lobster landings in Florida, including intema- prevalent by the late 1960's, facilitated a markedincrease in the tional catches,peaked in 1972 at 11.4 million pounds, valued at number of traps that could be worked per boat per day. A!so, approximately$13 million Table 1!, Total landingsremained the use of salted cowhide as lobster bait was introduced during high until 1975, when they declined to 7.4 million pounds a the mid-1960's; compared to fish heads and canned sardines, decreaseof 35 percent from 1972. The decfine in Florida land-

34 Fisheries, Vol. 5, No. 4

Beardsleyet al. 1975; Wise 1976!. A tremendousillegal market tributed data on ihe recreational catch of spiny lobsters in for sublegal-sizedlobsters shorts! also developed and persists Florida. Many of the statisticson landings and fishing effort were Johnson 1974; Warner et al. 1977!. In fact, trap robbing, pre- obtained through the courtesy of Statistics Division, TIMS, cipitated by declining catches and the illegal trade in shorts, Southeast FisheriesCenter, NMFS. Reviews of the manuscript becameso prevalent that various fishmen groups resorted to the were provided by Drs. F. G. Walton Smith, International renting of private patrol planes to protect their traps from OceanographicFoundation, Miami, Florida; Clarence P. Idyll, poachers.Collectively, theseincreasing harvest pressureson the NOAA, International Fisheries,Washington, D.C.; Dr. Albert C. resource provided impetus for the initiation of researchon the Jones, Southeast Fisheries Center, NMFS, Miami; and Wm. biology and management of the spiny lobster by federal, state, Seaman, Jr., Florida Sea Grant College, University of Florida, and university scientists Seaman and Aska 1974!. The upshot Gainesville.This study was supported by NOAA F!orida Sea of this effort prompted the American Institute of Fishery Re- Grant Contract Nos. 0<7-158-4406 and 04-8-M01-76!, the search Biologists to sponsor the publication of a position paper U.S. Fish and Wildlife Service Florida Cooperative Fish and on the Horida spiny lobster fishery Beardsley et al. 1975!. Wildlife Research Unit Contract No, 14-16-009-79-061!, and A tangential problem, specifica!iyillegal narcoticssmuggling, the University of FIorida. also cast shadowson the south Florida lobster fishery during the This paper is a contribution Journal Series 2406! of the 1970's, The smuggling of narcoticsinto south Florida is preva- Florida Agricultural Experiment Station, Gainesville, Florida. lent, hard to police and very lucrative. That smuggling affects the lobster fishery is certain, but its short- and long-term impact LITERATURE CITED can only be surmised.To illustrate, lobster fishing is a hard way Beardeley,G.L, T.d. Costello, G. E.Davis, A. C. donee, and D, C. Simmons. 1975. The to make a modest living, whereassmuggling is an easy, though Roridaspiny lobster 8sheryi A whitepaper. Fla. Scf. 38!:144 149 risky, way to make megadollars;consequently, some fishermen Brice.J. d. 1898. 6. The 8shand lishedes ofthe coastal waters of RorldaPages 263 342in UnitedStates Commission of Fish and Fisheries. Part XXII. Report of theCommissioner for have tumed their attention to narcotics smuggling because of theyear ending June 30, 1896. U.S. Government Printing Ofgce, Washington, D.C., USA. the lure of big money. Smuggling activities, therefore, lnay be Browne,d. B. 1973912!. KeyWest The ald and the new A facsimilereproduction of the 1912edition. BicentennialRondlana Facsimile Series!. Univ. Ra. Press, Gainesville, Fla. actually providing a reduction in the fishing effort for lobster; USA. such a reduction, of course, would not be evident in the re- Chace,F. A.,and W. H. Dumont. 1949. Spiny lobsters Idennnratian, world distribution, and ported statistics on effort. Substantial overcapitalization could U.S.trade. Comm. Rsh. Rev. 11!:1 12. Chekunova.W.I. 1972.Geagragcheskoe rasproslranenle kolyuchlkh langustov i ekologiches- also be occurring becausesmuggling requires larger, faster, and kiefaktory, apredefyayushchis ikh pramyslovye skopleniya. Geographical distribution of more expensivevessels than lobster fishing; these vessels,after spiny!obstrns and ecological factors determining their commercial concentmtions. ! English the heyday of smuggling has passed,will undoubtedly be redi- translationNMFS Foreign Fishery Translahons Tr 73-5 X511 pp. ! BonitetMiravogo Okenana,Trudy Vses. Nauchno-Issled Inst. Morsk Rybn Khoz.Okeanogr. VNIRO! rected toward fishing. And, finally, no industry or community 771110 119. can be infiltrated by illegal activities and still remain unaffected. Cobb,S. d. 1976.The Amencan lobster: The biology of Homarus emericanus. University of RhodeIs!and/NOAA Sea Grant,!dar. Tech. Rep. 49:1 32. And a new uncertainty for the spiny lobster fishery has liter- Coglns,J. W.1887. X!V. Reporton the discovery and investigation offishing grounds, made ally swept in from the sea. The recent 980! massiveimmigra- bythe Fish Comnessian steamer Alhatrass dudng a cmisealong the Atlantic Comt and in the tion of Cuban refugees into Key West and south Florida will Gulfol iMestco;with notes on the Gug 8shenes. Pages 217 311in UnitedStates Camrrus- sional Fishand Rsherfes. Pari Xlf .Report of theCommissioner for 1885. United States very likely impact Florida's lobster fishery through the addition GovernmentPrinting Oglce, Washington, D.C., USA. of more Cuban lobstermen to the already crowded fishing Cope,C. E. 1959.Spiny lobster gear and Fishing methods U S Fish Wildl Serv.Leaf ranks just as it did in the 1960's. 487:1 17. Crawford,D. R., and W. J. J. DeSmidt 1922. The spiny lobster, Panulirus mgus, of southern The Fishery Conservationand ManagementAct of 1976 has Florida:its natural history and utllisatlon. Bull. Bur. Fish. 38: 281 310.United States Gov- had significant impact on many United States fisheries,includ- emmentPanting Ofhce, Washington, D.C. Dawson,C. E., andC. P. Idyll.1951. Invesagatlons on the Florida spiny lobster. Panuhrus ing that for the spiny lobster. Importantly, P, argus is a species argre Latreille!Ra. Board Conserv. Tech. Serv. 2:1 39. targeted for Fishery Management Plans by the South Atlantic, DeBrahm, W. G. 1974 772!. The At anticpiot. A facsimilereproducnan ofthe 1772 edition Gulf of , and Caribbean Fishery Management Councils. BicentennialFloddiana Facsimile Series!. Univ. Presses Ra Gainesville,Ra. USA George,R. W.,and A. R.Main. 1967. The evolunon ol spiny lobsters Pallnuridae I:A studyof The direction and impact of these federal regulations on the evolutionin themarine environment. Eva!ution 21!:803 820. resource is as yet unknown. That new management strategies Hotmsen,A. 1972.Hmvesnng spiny lobsters off Florida.Madtlmes 1686 7. are needed to insure the maintenance of a maximum sustain- Johnson,L E. 1974,Stansecal trends in thespiny lobster 8shery. Pages 15 18in W.Seaman, Jr.,and D. Y. Asks,eds. Conference proceedings: Research and information needs of the able yield in the south Florida spiny lobster fishery is, however, Floridaspiny lobster 8shery. Ra. Sea Grant Pragr. SUSF-SG-74-201. fully recognized, Mathls,K., J. C.Cato, R. L. Degner, P. D. Landrum,and F, d, Prochaska. 1979 Commer- cial8shing acnvity and faoity needs in Rortda:Dade and Monroe Counties Univ. Ra., Agnc History reflects that the spiny lobster has not been a passive MarketRes. Cent. Ind. Rep 79-3:1 68. force in the settlement and development of south Florida. Nationalblur!ac Fisheries Service 1974. Amencan and spiny lobsters 1947 73: Basic Rather this marine crustaceanhas vividly impacted the culture, econamicindicators. Naaonal Manne Rshedes Service, NOAA, Carr Fish. Stat. 6272: 1 58. Prachaeka,F.d. 1976a.Ronde commerdal manne 8sheries: Growth, re!ative importance, and sociology, and economicsof the F!orida Keys, a processthat is inputtrends. Ra. Sea Grant Prost. Rep. 11:1 50, destined to continue in the future. De Brahm's non-clawed Prochaska,F.d. 1976b.An economicanalysis ol effort end yield in theFlorida spiny lobster "shrimps" have indeed done well! indushywith management considerations. Pages 661 674in B.F. Phillips, B. F.Cobb, ill, andA. B. Stockton,compilers. Proceedings of the first annual tropical and subtropical 8shenestechnotagtml conference. Texas A&M Univ. Sea Grant TAh U-SG-77-105. Prochaska,F.J., andd. R, Beards.1975. Rartda's 8sheries management programs: Their ACKNOWLEDGMENTS development,administration, andcurrent status. Untv. Fla., Agri. Exp. Sta. Bull. 768:1 64 Prochaska,F.d., and J. C. Cato. 1974. Landings, values, and prices in commercial8sheries John D. Newton, Jr., Department of Anthropology, and Wal- forthe Florida Keys region. Ra, Grant Progr. SUSF-SG-74- Xii 2 !. ter R. Keithley, Jr,, Department of Food and ResourceEconom- Schroeder.W.C. 1924.Rsherles of KeyWest and the dam industry of southernRorida AppenrkxXII to the Report of the United States Commissioner of Fishenes for 1923.Bureau ics, University of Florida, Gainesville,contributed resource ma- ol FisheriesDocument 962. United States Government Printing Ofgce, Washington, D.C. terials and ideas to the paper. Irvin Knowles and Troy Baker, Seaman.W., Jr., and D. Y, Aska. eds. 1974. Conference proceedings: Research and informa- veteran Key West lobstermen, generouslyshared their observa- tion needso! the Rondespiny lobster 8shery. Ra. SeaGrant progr. SUSF-SG-74- 201:1 64. tions on the development of the spiny lobster fishery. Jalnes R. Slehenaler,d,B, 1955.Commerdal fishing gear and 8shing methods in Rarlda.Ra. Board Zuboy, SoutheastFisheries Center, NMFS, Miami, Florid, can- Conserv.Tech. Ser. 13:1 47

36 Fisheries, Vol. 5, No. 4 Simmons,D.C. 1974Studies on the U.S spiny lobster fishery in theBahamas region Pages Williams,A. a. I'!hs 'Ihedecapod o the Carolinas U S.ash Wild! Sent Fish. 37 41in W.Seaman Jr andD Askseds Conference proceedings Research anJ inf

REVIEW OF THE FLORIDA SPINY LOBSTER RESOURCE

David C. Simmons

ABSTRACT The spinylobster resource !n Floridahas developed into an importantcommercial and recreationalFishery resource. Fishing pressureon thisresource has increased dramatically over the last fifteen years without a correspondingincrease in landings.Some important research has been completed on this resource. but before a sound managementplan can be developed some critical inforroation gaps need to be fiiied.

THE RESOURCE occasionally captured by Horida fishermen are the scvllarids. These lobsters have several common names including slipper he Florida commercial lobsters, shovelnose lobsters, bulldozers, and cockroaches. T spiny lobster fishery has Thev are not known to be in commercial abundance. in recent years ranked second For the remainder of this review all references to spiny only to Rorida's shrimp fishery lobsters will refer only to . On several occasions in economic importance. The referencewill be made to carapacelength. This measurementis fishery is concentrated in the used to set minimum legal sizeand is taken on the dorsal surface southern part of the state from from the groove between the horns directly above the eyes to around Fort Pierce southward the rear edge of the carapaceor "head" region Fig. 2!. Fisher- and to the Dry Tortugas to the men usually use a go-no-go gauge to determine if a lobsteris of west. It extends into Florida Bay legal size. and out to about 50 fathoms along the edge of the Gulf LIFE HISTORY FEATURES Stream Fig. 1!. There is some David C. Simmons* additional recreational harvest Adult spiny lobsters are found on coral reefs and rocky sub- in the northern Gulf of Mexico. strate from Brazil, through the Caribbean and Gulf of Mexico, to In 1977, 92 percent of the Florida domestic lobster production North Carolina, with a disjunct population in Bermuda Wil- was landed on the west coast of Florida, primarily in the Keys. liams 1965!. They inhabit shallow patch reefs,particularly in the Roughly one-third of the Florida landings are taken within three miles of the Florida coast. The fishery is essentiallva single speciesfishery directed to- ward the common spiny lobster or crawfish Panulirus argus, althoughtwo other species of spinv lobster occur in Florida waters. Panulirus laeui cauda, the smoothtailed spiny lobster, is of commercial importance in Brazilian waters, but is rarely cap- tured this far north. It is easily distinguished from the other spedes by having several small white spots confined onlv to the margins of the tail.Panulirus guttafus, the spotted spiny lobster, is fairly abundant in Florida waters, but it remains in rocky or coral reef areas and therefore is not likely to encounter lobster traps. The averagesize of the spotted lobster is smaller than the other spedes Caillouet et al. 1971! and it is easily distinguished from them by havingnumerous small white spots covering its entire body. Panulirus argus, the speciesthat forms the basisof Florida's lobster fishery, is distinguished from the other two spiny lobsters by having large round white spots on the dorsal surface of the tai! segments,Another group of lobsters that are Figure 1. Checkered area representsapproximate location of the *See page 56 for biography. Florida spiny lobsterfishery.

July August 1980 37 i~ CARAPACEEERGru ~l During mating, the male depositssperm packets tar spots! on I I the underside of the female. A short time after the mating act, the female extrudes the eggs through openings at the basesof the third walking legs, while the fourth pair of legs is used to scratch the surface of the sperm packet. The newly released sperm fertilizesthe eggsas they pass back and stick to fine hairs on the swimmerettes on the underside of the female's tail. The eggsare carried by the "bemed" female for about 4 weeks until they hatch. Some reproductive activity occurs throughout the year in Rorida waters but is concentrated during March through July. Figure 2. A go-no-go gauge used to determine if a spiny lobster Femalesmigrate into areaswhere there are resident adult males is of legal size. for mating and then move to the edges of reefs or coastal shelvesto incubate and releaselarvae Sutcliffe 1952; Cooper late summer and early fall Davis 1977a!, and are caught to et al. 1975; Buesa 1965!. During this time, nearshore reefs depths of 100 fathoms Robert Felik, National Fisheries,Inc., often show a predominance of males 0-80 percent! Sutcliffe Miami, personal communication!. 1952; Olsen et al. 1975; Davis 1977 a!. Females generally re- Juvenile and adult lobsters are gregarious during daylight tiirn to inshore reefs after hatching their eggs. Most female P hours, when they are found congregated in holes, caves, and argus reach maturity between 3 and 3'/2 inches in carapace under ledges associatedwith hard corals and rocky outcrop- length, and males at probably a somewhat larger size Morgan pings. individua! lobstersleave these day-time sheltersnightly to 1977!. Females reach peak egg production between 4 and 5 forage on adjacent reef areas and grass flats Herrnkind et al. inches in carapace length Kanciruk and Herrnkind 1976a; 1975!, Creaser 1950!. The number of eggsproduced at each spawning Most migrations take place as the lobsters progress from one increaseswith increasinglobster size. A lobster with a carapace life cycle stage to the next, or in responseto seasonalenviron- length of 3 inches will produce about '/z million eggs and a mental changes. The larvae, called phy/fosomos [or phyf- lobster with a carapacelength of 5 incheswill produce about 1.5 fosomesj, are flat, transparent greatly modified for a million eggs, planktonic existence.At hatching they are about 1/16 of an inch The diet of juvenile and adult spiny lobsters includes mol- long and after undergoing about 11 developmental stagesreach lusks, crustaceans,algae, echinoderms,coelenterates, annelids, a length of about an inch and a half Lewis 1951!. The duration and sponges Barroso Fernandes 1969; Peacock 1974; of the larval life is unknown. Estimatesrange from 3 months to 1 Hermkind et al. 1975; David 1977a!. Food items collected dur- year. This planktonic stage provides for long distance dispersal ing nocturnal foraging may be carried back to dens or traps! for potential, but controlled vertical movements in the water col- consumption during the day Craig 1974; Gary E. Davis, Na- umn may allow the larvae to remain in the area of hatching via tional Park Service,personal communication!. Medusaemay be eddies, layered counter currents, and other irregularitiesin the important in the diet of older larvae Provenzano 1969!. movements of water masses. Larvae that survive their Growth of spiny lobsters takes place as a result of molting. planktonic existencemetamorphose into postlaivae that resem- Just before and during molting, the water that is taken up causes ble adults in form but are transparent. The postlaivae swim the body tissuesto distend, thereby increasingin size. Legs and toward shore bv night and buirow in the bottom by day until antennae lost or broken since the last molt can be regenerated, they reach inshore seagrassnurseries, where they gradually be- but growth rate may suffer significantly because energy nor- come pigmented Johnson 1974; Serfling and Ford 1975; Phil- mally devoted to increasesin size is spent in tissue repair and lips 1977; William G. Lyons, Florida Department of Natural replacement. injuries may also hinder effective foraging and Resources, personal communication!. defense, further reducing growth rates. After shedding the exo- Postlarvaeare at first solitary, but as they develop they find skejeton, a new soft shell is exposed that takes several days to protection in numbers to ward off and avoid predators. They harden, depending on ambient water temperature and the age seek shelter in sponges,under rocks and crevices,in algal mats, of the lobster, The componentsof growth have been studied by and among the spines of sea urchins. Soon they require larger utilizing marked animals released and recaptured after various shelter and congregatein submerged root systemsof red man- time periods of freedom in the wild. These studies have shown groves, in rocky outcrops, and in shallow patch reefs. After that very young juveniles molt about every month for the first spending up to 3 years in such nursery areas, the juveniles several months of demersal existence. Once they reach a mature and move to offshore reefs, Offshore, they tend to be- carapacelength of about 2 inches, molting takes place at 12- to come resident with seasonal migrations between shallow and 14-week intetvajs. Carapacelength increasesabout V4inch dur- deep reefs in conjunction with reproductive activity and in re- ing each molt, until they reach maturity when significant dif- sponse to the onset of winter conditions, particularly in sub- ferences in growth rates between sexes become apparent, tropic and temperate regions Hermkind 1977; Davis 1977a!. Males continue adding about 1 inch per year, while the annual High concentrationsof lobsterstend to build up in shallow reefs growth of females,which expend greater quantities of energy in in the fall, following the return of egg-bearingfemales. In areas egg production and care, is reduced to about '/2 inch Davis with limited or widely dispersedfood and shelter, nomadism is 1978; Munro 1974; Peacock 1974; Sweat 1968; Gary E. Davis, frequently a common movement pattern for both adult and National Park Service, personal communication!. immature lobsters Hermkind 1977!. Estimatesof age from postlarvaeand sizeindicate that a lobs- Resident reef dwelling adult lobsters range up to 300 yards ter with a carapacelength of 3 incheswould be about 2'/z years from the dens out onto adjacent vegetated areas nightly, Typi- old Munro 1974; Olsen and Koblick 1975!. cally gregarious by day, nocturnal foraging is done solitarily During their planktonic phase,larval lobstersare preyed upon Hennkind et al. 1975!. by pelagic fishes, including tunas Baisre 1963!. Natural

Rsheries, Vol. 5, No. 4 enemiesof juveniles and adults are various speciesof groupers, snappers,, octopi, turtles, and skates Buesa 1965; FAO E 000 1968; Randall 1967!. A fatal and apparently infectious disease RVl is causedbv microsporidianprotozoan infection Edwin lversen, 0 I,'Io RosenstielSchool of Marine and Atmospheric Science,personal N communication!. Affected individuals become disoriented and ILO R their muscle tissue is "milky" in appearance, similar to what is L known as "cotton" shrimp Sindermann and Rosenfield 1967!. I,ao Man accounts for the majority of lobster mortalities in areas Nss IOO where there is a directed fishery for lobsters. In a heavily fished R area off Jamaica, total annual mortality was estimated at 78 percent with 7 percent attributed to natural mortality and 71 percent to fishing mortality Munro 1974!. Becauseof the small 700 laaa sz 'Is ao sr aa 'as aa Io 74 '70 70 size distribution of lobsters captured off the Florida Keys 'IS 'II ' I7 0'I 'a'I a'S 'I.'7 ao LOBSTERSISNINO SEISON Warner et al. 1977!, it appearsnearly every lobster that attains 'TESRst legal size during a fishing seasonis captured. Figure 3. Number of commercial spiny lobster fishing permits is- sued each seasonby the Florida Department of Natural Resources THE FISHERY 954-55 to 1977-78!. The Florida spiny lobster fishery began around the turn of the with wooden lath. The traps are about 3 x 2 x 2 feet in size; century. During the early beginnings of this fishery, lobsters slatsare spaced1'/z inchesapart, with an openingin the top were used extensivelyas bait for catchingother fish as well as an guarded by wooden teeth, and weighted with cement. These important food for man. In the 1920's lobsters had a market traps are buoyed with styrofoam fioats and set individually or in value of $1.00 per dozen, but 4 or 5 dozen when used for bait strings of about 10 traps. The most popular baits used are fish could net a fisherman $40 to $50 worth of fish Crawford and heads or cow hide. It is a common practice in Florida to leave De Srnidt 1922!. small lobstersin traps to attract larger, legal size, lobsters, As exploitation increased,regulations were imposed to pro- Fishermenwork about700 trapseach and pufi about 170traps tect the resource. An act passed in 1919 was the first dealing per day. In the early 1950's,fishermen could make adequate with lobster fishing in Florida and established a 3-month closed catchesby workingonly 100 to 300 traps Dawsonand Idyll season from March 1 to June 1. Excluded from the closed 1951!. In 1952 there was an estimated15,000 traps in the season were lobsters taken for bait purposes. In 1929 the first fishery. This increasedto 165,000 in 1969, In 1970 when the size restriction was enacted and was set at 1 pound. Although $50permit fee was charged, the numberof trapsdropped to both closed season and minimum size have changed over the 130,000.Since 1970,there hasbeen a steadyincrease in the years, these were the only lobster fishery regulations imposed numberof traps,reaching over '/2 millionin 1975 Fig.4!. At a until 1965. Legislation introduced in 1965 emphasized gear costof about $12 per trap Williamsand Prochaska1977!, the regulations and required commercial fishermen to obtain a total number of traps in the fishery representsa considerable permit to fish for lobsters. The 1965 act also prohibited the investment. harvest of egg-bearingfemale lobsters.In addition to protecting egg-bearing females, present regulations include minimum carapacelength of more than 3 inches, and a closed seasonof April 1 through July 25. To fish lobsterscommercially, a person must purchasea permit from the State at a cost of $50 and the permit number must be displayed on all traps, floats, and boats. In addition, traps and boats must be color coded. Fishermen without a lobster fishing permit are limited to 24 per day and O O0 770 they are not allowed to use traps. It is unlawful for any fisher- 70 man to take lobsterswith spears,hooks, or grabs. An exclusive aooo ~S. 0 seasonfor sports fishermen was establishedon July 20 and 21 I of each year. Fishermen are restricted to 6 lobsters per day OOI ssa Ih O during this 2-day season Prochaskaand Baarda 1975; F!orida O Sao 0 Statutes, Chapter 370.14!. 0 The number of commercial lobster fishing permits issued by Sl0 IR the State each year increased from 255 during the 1954 55 s seasonto a high of 2,719 during the 1969 70 season.In 1970 SR170 a $50 fee was chargedto obtain a permit and the number issued dropped to 961, There was a steady increase in the number permits from the 1970 71 season to the 1975 76 season when 1,824 permits were issued. The number of permits have remained about the same since the 1975 76 season Fig. 3!. The commercial lobster fishery is almost exclusively a trap fishery, although small amounts are taken by diving and by 'I0 'aa 'sa 70 'll 70 bully netting, and as an incidental catch in shrimp trawis. In YSRR 1974, 96 percent of lobster landings were harvested in lobster Figure 4, Number of spinylobster traps used in Florida each year traps. The most popular used in Florida is made 952 to 1975!.

July August 1980 39 There is little standardizationin spiny lobster fishing craft de- sign. Typically, however, they are about 35 feet long, with a wooden hull, and powered with a dieselengine. There is usually a small cabin forward with the aft deck open work area. The total number of fishing craft in the lobster fishery and average sizeare increasing.The nuinber of craft with a carryingcapacity of 5 5 tons or greateris increasingat a greaterrate than smaller craft. In 1952 there were about 100 craft in the fishery that had a carrying capacity less than 5 tons. In 1971 the number had increasedto 250. In 1952 there was no craft in the fishery with a carrying capacity of 5 tons or greater. In 1971 there was 270 of these larger craft. During the period 1959 through 1961 the averagecarrying capacity of the large craft was 10 tons. During Figure 6. Number of poundsof spiny lobsters landed in Florida the period 1969 through 1971 this averagehad increasedto 25 each year 951 to 1977!, divided into domestic production solid line! and productionfrom international waters dashed line!. tons Williams and Prochaska 1976!, Most of the lobster fishermen working in Florida waters begin pulling their traps in the early morning and return to port in the ings have been kept separatelysince 1964 Fig. 6!. Until 1975 late afternoon for unloading. The catch is usually kept alive in a landings from iniemational waters were almost entirely from container on deck. Some fishermen make extended trips lasting fishermen working the Bahama Banks. In 1975 the Govem- a couple of weeks to areassuch as the Dry Tortugas. On these ment of the Bahamas followed the lead of the United States longer trips lobsters are usually tailed and iced down or frozen. with the American lobster and declared the spiny lobster a crea- There is usually a one- or two-man crew and the most common ture of their continental shelf thus excluding all foreign cornmer- way of paying helpers is a lump sum for the day Noetzel and cial lobster fishing. This resulted in a drop in landings from 4,3 Wojnowski 1975!. The number of full-time lobster fishermen' million pounds in 1974 to 2 million pounds in 1975 and a has increasedfrom about 170 in 1952 to 1,600 in 1974 Fig. 5!. further reduction in 1976 to ys million pounds. In 1977 there Some of these fishermen, however, fished in international wat- was an increase in landings from international waters to 1.1 ers, primarily the Bahamas and the western Caribbean. million pounds. Since the early 1950's, Florida domestic landings of spiny I5 OD lobstershave fluctuated between a low of 1.6 miHionpounds in 1952 to a high of 6.8 million pounds in 1970. In recent years K4 I the fisher< has been based on lobsters that become legal size D during a fishingseason and this available yield can be harvested l,ODO with less than the present level of effort Prochaska 1976; Wil- liams 1976; Williams and Prochaska 1977!. Amounts of lobsters harvested in recent years therefore depend on the degree of s 0 successof larval recruitment about 3 years earlier and the de- gree of survival of juveniles. 5DO Because of increasing dock-side price per pound, there has been a trend of increasing value of domestic production even though the total weight of landings has fluctuated widely Fig. ZlO5 7!. The highestvalue of domesticlandings was reachedin 1974, 1'555 55 55 55 M 55 5 ~ 55 OO 55 when they were valued at $8.1 million. In 1977 the value was Figure5. Numberof full-timespiny lobster fishermen working in reduced to $6.7 million, a 17 percent decrease from 1974. Rorkla each year 952 to 1974!. During this same period the average price per pound increased from $1.23 in 1974 to $1.69 in 1977. This increasein price per An economic survey of 25 Rorida Keys lobster fishermen was pound helped offset a 39 percent reduction in pounds landed. conductedduring the 1973 74 fishingseason Prochaska and Williams 1976!. This survey showed that the average lobster fishingoperation produced $14,000 worth of lobstersper sea- son with a net return above total costs to the boat owner of $5,000.Fishing craft 16 to 22 feetin lengthhad an averagenet returnof $3,000and craft31 to 36 feet in lengthhad an aver- age net return of $7,000,This income,however, was usually supplementedby switchingto othertypes of fishingsuch as for stone crabs and mackerel. Some fishing operations are able to maintaina fairlyconstant income throughout the year by switch- ing to otherfisheries as lobstersbecome less available as the seasonprogresses Noetzel and Wojnowski1975!, Total reportedFlorida landings of spiny lobstersare corn- bined domesticproduction and productionfrom international waters. Amounts of these two components of the Rorida land-

'An individual who spends 50 percent or more of the working year in Figure 7. Value of spinylobsters landed in Rorida each year 951 commercial lobster fishing activities, inctudtng port activity such as ves- to 1977! divided into domesticproduction solid line! and produc- sel repair and re-rigging. tion from international waters dashed line!.

40 Fisheries, Vol. 5, No. 4 The dock-sidevalue of landingsfroin international waters was the process of preparing reports on their research that when at its highest in 1972, amounting to $6.8 million. Since then, complete will constitute a valuable contribution for managing landings have been reduced so the total value of landings froin the lobster resource, The Department of Natural Resources international waters has also been reduced. conducted similar research in the Middle and Upper Keys, in- The overall trend in the domesticcommercial lobster fishery is cluding a tagging study in 1978 79. increasing effort. The number of fishermen entering the fishery A managementstrategy that has recently gained some public each year is increasing,the number and the size of the fishing attention is the establishmentof spiny lobster sanctuaries.It has craft are becoming greater, and more traps are being used each been stated that closing certain nursery grounds and adult year, That the number of lobsters available each year is not habitat for spiny lobsters increasesbiomass, reduces mortality, decreasing means that the catch per unit of fishing effort is improves growth, and supplementsrecruitment to surrounding decreasing. Because of increasing dock-side price per pound areas. Closed areas also allow for undisturbed natural popula- the economic return to fishermen has shown a gradual upward tions, which provide a comparativedata base for management, trend Prochaska 1976; Williams and Prochaska 1976, 1977!. recreational non-harvesting use underwater observers!, and a The recreationalfisherman is another important user of Flori- reserve genetic pool Caribbean Fishery Management Council da's spiny lobster resource.The most popular recreational fish- 1978!. Fort Jefferson National Monument can already be con- ing method is skin or and catching lobsters with sidered a sanctuaryfor adult lobsters,and the State of Florida in gloved hands. A few recreational fishermen use a bully net to June 1979 passed a law establishing a spiny lobster nursery catch lobsters. This net consists of a mesh bag fastened to a sanctuary in Biscayne Bay and Card Sound, It has been esti- hoop that is attached at a right angle to a long pole, Bully- mated that 95 percent of the lobsters in the Bay are between netting requires clear shallow water and is usually done at night about 2 and 3 inches in carapacelength with 33 to 50 percent when lobsters are foraging for food. A light is directed toward having injuries in the form of missingantennae or legs. These the bottom and a glassbottom bucket is used to reduce glare. injuries are largely due to handling by fishermen. Injured When a lobsteris spotted, the net is thrust over it and the lobsterstake an additional year to attain legal sizebecause they entanglesitself in the webbing Cope 1959!. put energies into repairing the missing limbs instead of into An experimental sport harvest was camed out in Fort Jeffer- increasingcarapace length. Tagging studies have shown that as son National Monument at the Dry Tortugas. An 8-month-long lobsters mature they move out of the sanctuary where they are driver harvest, limited by a daily bag limit of two lobsters, dis- accessibleto fishermen Gary E. Davis, National Park Service, persed the density of the previously unfished population to 42 personnal communication!, percent of the pre-harvest density Davis 1977a, 1977b!. In addition to the points already raised, there are several The magnitude of the recreational harvest and its impact on other topics that will have to be considered in developing a the economy is unknown but is believed to be substantial. In management plan for Florida's spiny lobsters. For example, 1975 during the 2-day recreational lobster season it was esti- should the economic yield to commercial fiisherrnenbe im- mated that 1,300 pleasure craft with 4,100 active divers cap- proved? If so, some thought will have to be given to limiting the tured 10,700 legal-sizelobsters in Dade County. Unfortunately number of fishermen and units of gear. What is the effect of these types of isolated observations are not sufficient to make leaving short lobstersin traps to attract larger,! egal sizelobsters? seasonal catch and effort estimates Austin 1976!. The mag- ls it desirable to require that escape hatches be constructed in nitude of Florida's recreationallobster fishery is one of the most, traps so that when it is retrieved the shorts wi!I fall out? Is a if not the most, critical information needs for managing the 3-inch caparace length the proper minimum size?The Carib- Florida lobster resource. bean Fishery Management Coundl is considering a minimum carapacelength of 3' inches to assure at least one spawning INFORMATION NEEDS before capture and to take advantage of the rapid growth of smaller animals. There are several other critical information needs for manag- ing the Floridalobster resource. One importantquestion and COOPERATIVE RESEARCH probably the hardest one to answer is what is the origin of the postlarvae that enter Florida waters?It has been suggestedthat There is excellent cooperation among the lobster researchers Florida's spiny lobster population may be recruited from larval in Florida. These researchersand rneinbers from industry have lobsters carried into Florida waters from the Yucatan Channel held four annual workshops where information needs have Ingle et al. 1963; Sims and Ingle 1966!. Oceanographiccondi- been identified and progress on research projects given. The tions off Florida, however, are favorable for retaining larvae by proceedingsfrom three of these conferenceshave been pub- utilizing eddies and counter currents Brooks and Niiler 1975; lished Seaman and Aska 1974; Seaman and Jones 1975; Lee and Mooers 1977!. Dr. Robert Menziesand his associates Warner 1978!. Based on the impetus of the first two confer- at Nova University are examining this problem by studying the ences, a white paper was prepared on the Florida spiny lobster genetic makeup of lobsters taken from various locations in the fishery Beardsley et al. 1975!. This paper pointed out critical Caribbean area Menzies et al. 1977!. In a related study, the information needs and management actions were suggested. Department of Natural Resourcesis looking at the recruitment Through the cooperation of spiny lobster researchers of postlarvae to the Florida Keys, The postlarvae are being throughout the world, a bibliographyon spiny lobstershas been captured in specially designedtraps and quantitatively sampled prepared by the Florid Sea Grant Program Kandruk and to determine seasonal recruitment patterns. Hermkind 1976b!. The life history of the Florida spiny lobster needs to be better understood. The Florida Sea Grant Program in Key West re- cently completed studies on the life history of lobsters in the LITERATURE CITED lower keys including migration, reproduction, growth, and mor- Austin,C. B. 1976. Rscreaiional boaiimi in Bhcayne8ay. Univ. Miami sea Grant, spec. Rep. tality Warner et al. 1977!. These researchersare current!y in 5:247 253.

July August 1980 41 Baisre.J. A. 1964Sobre los estadios asde la langosta comun Punuiirus argus. Contrib. 1978National Park Service juvenile/sub-adult population studies Pages 8 12in CientInvmt. Pesq., Bauta. Cuba. 19 37 pp. SpinyLobster Res Rev Conf..Key West, Hoitda, December 16. 1976Ha. Sea Grant Barroso-Femandes,L.M. 1969. Sabre a alimentacaode langosta Punulirus argus Lah ! Program 1804,Crustacea Reptaniia I-estagto juvenal. Biol. Estud. Peace 9!:21 33 Dawson,C.E., Jr., and C. P. IdylL 1951. Invesrigattons onthe F!onda spiny lobster, Panuliius Beards ey,G.L, T. J. Costa!!o,G.E. Davis, A. C. Jones, and D. C. Simmons.1975. The argus Latreille! Fla BoardConserv., Tech Serv. 2. 39pp Floridaspiny lobster fishery. A whitepaper Fla Sci.38! 144 149 FAO.1968 Report to theGovernment of British Honduras Belize! on investigationsinto Brooks,I. H., and P. P, Niiler. 1975 The F!orida Current at KeyWest Summer 1972 J Mar marinefishery management, research and development policy for spinylobster fisheries Res.33!:83 . 92. Basedon the work of W.H. L. AllsoppRep FAO/UNDP TA!, 481! 95pp Buesa,R. J. 1965.Biology and Rshtng of spmylobster Pnnuiirus argus Latreille!. Pages Herrnklnd,W. F. 1977Movement patterns in pallnurid lobsters. Pages 209 244in B.F. 62 77 in 5 Boganov,ed. Soviet-Cuban fishery research. Translated fram the Russian by Phillipsand J. S Cobb coordlnators! Workshop onlobster and rock lobster ecology and IsraelProgram for Scientific Translations, Jerusalem 1969 Tt69-59016!. physiology.CSIRO Melbourne, Div. Fish. Oceanogr., Cir.7. Cai!louet,C.W.. JrG, L, Beardsley,and N. Chitty.1971. Notes on size, sex ratio, and Herrnkind,W.F., J. A. Van Derwalker, and L Barr.1975. Population dynamics, ecology and spawningolthe spiny lobster Ponulirus guttutus Latreille !,near Miami Beach, FIa BullMar behaviorofspmy lobsters, Punu/irus argus, ofSt. John, U.S V.lz IV!Habitahon, patterns of Sd.21!:944 951. movementand general behavior. Bull. Nat. Hzst. Mus. Los Angeles Co. 20:31 45. CaribbeanFishery Management Councg. 1978. Fishery management plan for spiny lobsters Ingle,R. M.. B. E!dred,IL W. Slms,and E. A. Bdred.1963. On the possible Canbbean PuertoRico and U.S VirginIslands!. February I drah prepared by SoutheastFtshenes originof F orida'sspiny lobster populahons. Fla. Board Conserv Tech. Ser 49. 12 pp Center,NMFS, Miami, Fla. 186 pp. Johnson,M. W. 1956 The larval development of the spiny lobster, Panufirus inter Coupon,R,A,. R, E its,and S. Serging. 1975 Populations dynamics, ecology and behavior of mptus Randall!,with notes on Ponulirusgrocilis Street!. Proc. California Acad Sci., spinylobsters. Panurrus argus. of St. John. U.S V I. 0 l! Populahoneshmarion and turnover 29! I 19 Bu!LNat. HLs Mus . Los Angeles Co 20:23 30 1974.On thedispersal ol' lobsterlarvae into the EastPaogc Banter , Cope.C. E. 1959.Spiny lobster gear and gshing methods Li S.Fish Wild Serv.,. Fish Leafl. Palinundae!.Fish. Bull . U.S.72!:639 647. 487.17 pp. Kanciruk,P., and W. F. Herrnktnd.1976a. Autumnal reproduction m Panulirus argus at Craig,A. K. 1974New developments in the spiny lobster fishery of southeasternFlorida. Bimini,Bahamas Bul!. Mar Sci. 262417 432 Pages131 142in Proc.Gug Canbb. Fish inst., 26th Anno. Sess. Kanciruk,P.,and W. F. Herrnklnd. eds. 1976a. An indexed bibliography ofthe spiny lobsters, Crsudozd,D.R., and W. J. J. De Smidt. 1922. The spiny lobster, Punufirus argus, of southern familyPalinuridae Fla Sea Grant Progr. Rep. 9 101pp. FIorida:Its natural history and utilization Bull U S.Bur. Fish 38:281 310. Lee.T, iMandC, N, K. Mooers. 1977 Near-bottom temperature and cunent variability over Creaser,E.P. 1950Repetition of egg-layingand number of eggsof theBermuda spiny theMiami slope and terrace. Bull. Mar. So. 27!:758 775. lobster.Pages 30 31in Proc.Gulf Caribb. Fish inst., 2nd Annu Sess Lewis,J. B. 1951.The phyllosoma larvae of thespiny lobster Panuiirus argus. Bull. Mar. Sci Davis.G. E. 1977a.Effects oi recreationalharvest on a spinylobster. Pun u ruei argus, popula- GulfCanbb 1!:89 102. fion.Bull. Mar. Sci. 27!:223 236. Morgan,G. R. 1977.Review of thepopulation dynamics of the Palinurldae Pages 245 258 1977bFishery harvest in an underwaterpark. Proc 3rd Int.Coral Reef Symp.. in B F Phillipsand J 5 Cobb,coordinators Workshop on lobster and rock lobsterecology R.S.M.A.S.,Mmnt, F!a. !:605 608. andphysiology CSIRO, Melbourne, Div F shOceanogr., Circ 7

THE LARVAL RECRUITMENT PROBLEM OF THE SPINY LOBSTER

Robert A. Menzies and J. Michael Kerrigan

ith any renewable natu- regiODSwhere the genuSPanufifus iS COmmOn,majOr fiSherieS ral resource, if various exist for P. argus Fig. 1! in Florida, the Bahamas, most Carib- parameters such as mortality, bean islands, the east coast of Central America, and the Carib- fecundity, and recruitment are bean and Atlantic coasts of South America southward to below known, resource managerscan the Amazon River. P. /aeuicauda is Fig. 2! also a major con- project harvest rates so as to tributor to this fishery principally in and around Brazil and P. sustain reasonable yields Lar- gutt IIUS Fig. 3! iSa minOrCOntributOr thrOughOut the range. On kin 1977!. Assessment of these the Pacific Coast of the Americas P. inteITupthfsat one time parameters is sometimes dif- supported a large fishery in California, but presently the Baja ficult because of peculiarities in California Mexico! area is the main producer. Farther south the life cycle or behavior of the P. grcfcilisand P. infkitus make up the majOr fiShery. resourcespecies, In many cases Throughout the smaller PacihcIslands P. penicillatus is wide- the most difficult parameter to spread, although P. marginatus is the major spedes in the determine is identification of the Robert A, Menzies Hawaiian Islands. Farther west one of the world's largest rock management unit. This is par- lobster fisheries P. cygnus, Fig. 4J is centered in Western Aus- ticularly acute in the case of spiny lobsters because of their tralia. A variety of other species contributes to fisheries long-lived pelagic, planktonic stage. However, becauseof their throughout the Padfic and Indian Oceansfrom to South worldwide economic importance, considerable effort has gone Africa. into research on their life cycle, The main difficulty in identification of spiny lobster manage- Spiny lobsters or rock lobsters as they are sometimes ment units, as was mentioned earlier, is their long pelagic called represent a respectablepiece of the fisheriesportion of planktonic phase. In general, the life history of all spiny lobsters the economy of many countries in tropical, subtropical, and is very similar Eig. 5!, In the case of P. argus, breeding and temperate zones. Taxonomically the group is placed in the de- spawning usually take place on the reef tracts that parallel the capod crustaceanfamily Palinuridae and is comprised of eight shores throughout its range. The larvae released,called phyllo- genera George and Main 1967!. The two most important gen- somes,are shortly at the mercy of oceancurrents and rarely find era economically speaking are casus species! and Panulirus their way into nearshore or bay waters. 9 species!,Jasvs, which abounds in the cooler subtropical and The pelagic planktonic existence lasts from six months to a temperate regions, supports major fisheries in South Africa, year, depending upon the species.For some speciessuch asP. Southern Austraha, Tasmania, and New Zealand, In the warmer argus the length of this phase is not fixed with certainty Lewis

42 Fisheries, Vok 5, No. 4 Munro,J. L. 1974.The biology, ecology, exploitaaon and managementofCaribbean reef RandaB.J. E. 1967Food habits of reefAshes of theWest indies Stud Trop Ocesnogr . Inst Ashes,Part Vl Thebinlogy. eco!ogy and biometrics of Caribbean reef fishes: Crustaceans Mar.Sci, Univ Miami 5 665 847 spinylobsters and crabs!. Res. Rep from Zool Dep., Univ West Indies. No 3. 57pp Seaman,W., Jr., and D. Y. Aska.eds. 1974 Research and information needs of theFlonda Menzies.R.A.. J. M. Kerrigan,and P. Kanciruk, 1977 A biochemical-geneecapproach to spinylobster Ashery Ha SeaGrant Proqr, Rep. 1 64pp thelarval recruitment problem of thespiny lobster Penuirus argus Page 38 in B F Phillips, Seaman,W., Jr., andA..C. Jones, eds. 1975. Review of Horidaspiny lobster resotrch, andJ S.Cobb, coorchnators Workshop onlobster and rock lobster ecology and physiology. includingresults and recommendations oi a colloquium held October I and 2 I 4 in CSIROMelbourne Div Fish Oceanogr . Cir 7 MiamiHonda: Summaries of current lobster research pro!ects in Flonda State Univ Syst NoetzeLB. G.,and M. G, Ivlojnowski. 1975 Costs and earnings inthe spiny lobster fishery, Ha SeaGrant Progr. Gainesville, and NMFS. Southeast Fish. Cent Mimi 54pp HondaKeys. Mar. Fish Rev. 37!:25 31 Serfling,S. A.,and R F. Ford.1975 Ecoilogical studirn ol' the puerulus larva stage of tlie Olsen,D, A, W.F. Hermkind,and R. A. Cooper, 1975 Population dynamics, ecology and Californiaspeiy lobster,Ponuiirus interruptus. Fish. Bull, U S73! 360 377 behaviorof spinylobsters Ponvjrvs argus, of St John.U SV I: I ! Introducnonandqeneral Sims,H. W ..and R. M. Ingle. 1967. Canbbean recruitment of Flonda's spiny lobster popula- populahoncharacteristics Bull Nat.Hist Mus. Los Angeles Co. 20 11 16 tion Q J Fla Aced.So 29! 207 242 Olsen,D. A., and I. G. Koblick.1975 Popo!anon dynamics, ecology and behavior of spmy Sindermann,C.J., and A. Rosenteld.1967. Principal diseases of commercially important lobsters,Pano!irus argus. of St. John. U.S.V.I 1! Growthand mortahty Bull Nat Hist mannebivalve Mullusca and Ciustacea. Fish. Bull.. LI S 66! 335 385 Mus.Los Angeles Co. c0 17 21 Sutcliffe,W. H., Jr. 1952.Some observa!tons of the breeding and migration of theBi. rmuda Peacock.N. A. 1974A studyof thespiny lobster hshery of Antiguaand Barbuda. Pages spinylobster Panukiur argus Paqes 64 69in ProcGult Canbb Fish Inst 4thAnnu Sess 117 130in ProcGulf Caribb Fish Inst, 26th Annu Sess Sweat,D. E. 1968Growth and tagging studies on Ponulirus argus Laueille! in theFlonila Phillips,B. F. 1977.A reviewof thelarval ecology of rocklobsters Pages 174 185in B. F KeysHa BoardConserv. Mar. Res Lab. Ser. 57. 30 pp Phillipsand J 5 Cobb,coordinators. Workshop on lobsterand rock lobster ecology and Warner.E. E.. C.L Combs,and D. R. Gregory,Jr 1977Biological studies of thespiny physiologyCSIRP Melbourne, Dtv. Fish, Oceanogr Cir 7 lobsterPunvkius argus Decapoda Palinundae! in South Horida Paqes 166- 183inProc Prochaska,F.J. 1976An economicanalysis of esortand yield in theFlonda spiny lobster GulfCanbb Fish Inst.. 29th Anno Sess industrywith management considemtions. Pages 660 674in B F.Phrilips J. S.Cobb III. Warner.R. D., ed.1978 Spiny lobster research review Proceedmgs of a conferenceheld andA B Stockton.compilers. Proc. of the Arst tropical and subtropical hshenes technological December16 1976,in Key West, Flonda. Ha SeaGrant Progr. Tech Pap No 4 56pp conference.Texas A&M Univ. Sea Cirant Coll TAMU SG-77-105. Williams,A. B.1965 Marine decapod crustaceans ofthe Carolinas Fish Bull U S.65! 2'!8 Prochaska,F.J., andJ. R. Baarda.1975. Florida's Asheries management programs: Their PP developmentadmimstraaon, anticurrent status Agnc Exper Sta., Inst Food Agr Sci.. Williams,J.S. 1976An economic analysn ofalternative management strategies lorthe spur, Llniv.Fla Bull 76864 pp lobsterindustry A dissertationpresented tothe graduate counal of Unw fla 164pp Prochaska,F.J., and J. S, Williams.1976 Economic analysis ol cost and returns in thespiny Williams,J. S., and F. J. Prochaska. 1976. The Florida spiny lobster fishery Landings, prices lobsterhshery by boat and vessel size. Fla Sea Cirant Mar Adv Bull SUSF-SG 76.004 18 andresource producnvity Ha SeaGrant Progr. Rep. 12. 50 pp PP . 1977Maximum economic yield and resource allocation in the spiny inbsti r mdustni Provenzano,A.J., Jr.1969 Recent experiments onthe laboratory reanng of tropicallobster South,l AgncEcon. Jy:145' 149. larvae.Pages 152 157in ProcGulf Caribb Fish Inst. 21st Annu Sess

Figure 1. Dorsal view of Panulirus argus. Figure 3. Dorsal view of P. guttatus.

4

+x

Figure 2. From top to bottom; P, argus, P. guttatus, and P. Figure4. Dorsalview of P, cygnus,the westernAustralian rock laevicauda. All are mature adults. Note the smaller, more slender lobster. stature of P. guttatus and the more stout, heavier-bodiedstature of P. laeuicauda. The antennae of these two are broken or missing.

July August 1980 43 Figure5. Schematicrepresentation of the life cycleof P. argus.Adult lobsters{1! usually largerthan 75%0 cm Cl inhabitthe reeftracts paralleling coastal islands and mainland. Breeding takes place here and female lobsters retain larvaefor 1-2 weeksbefore releasing them to opensea currents. ! The larvae,or phyllosomes,spend approximately 6 monthsas planktonwhile proceeding through 10-12 developmental stages to thepuerulus stage. ! Thepuerulus stage is a transparentflattened replica of an adult lobster, approximately 1 inch long. These forms move in to shallowwaters to begintheir benthicexistence, After a numberof moltsand developmentof pigmentation,! the postlarvaemove into "nurseryareas" such as shallowcoastal waters and bays.They inhabitthese areas for abouttwo yearsmore before moving out to the reef tracts. 1951!. During this period the larvae develop through 10 12 relative recruitment at the posilarval stage Litfle 1978!. Follow- reasonablywell-deflned stages,completing the ing the design of Witham et al. 968!, they have devised a to the puerulus stage, which marks the beginning of a benthic "habitat" to collect larvae as they settle out in shallow waters. existence.The phyllosomesare typically transparent, flattened, Figure 6 shows the range of forms found on these habitats or and long-legged,their shapepresumably maximizingbuoyancy. collectors. The puerulus on the other hand, although still transparent and Interestingly the observationsof the DNR researchersin the slighily flattened, appears as a miniature lobster. This appear- lower Keys and those of our group in the upper Keys over the ance is further enhanced as the postlarvae develop pigmenta- past two years reflect maximum posflarval influx eight to ten tion within several days after settling out Fig 6!. months after rnaxirnum spawning activity has occurred in these The rest of the life cycle involves approximately a three-year locations. Either the phyllosome life of six months tentatively stay in bay or nearshore "nursery" areasuntil they reach a size suggestedby Lewis 951! is wrong or the bulk of incoming of approximately 75 85 rnm legal harvest size in Horida is 76 posflarvaeare from populations having intense breeding activity mm! in carapacelength CL! or about 340 g /4 lb!, After the in October and November. "nursery" phase, the animals generally move out to the There is some evidence in the P. argus flshery that recruit- offshore reef tracts, where they spend most of the rest of their ment of larvae to Florida may be from non-Florida populations, lives. Breeding and spawning in the Horida Straits begins in Sims and Ingle 967! conducted a number of drift botfle exper- spring, reaching a maximum in June and July and tapering off iments and surveyed the density of various stages of phyllo- in early fall Davis 1975; Warner et al. 1977; Kanciruk and somes in the Florida Straits and Yucatan Channel, They Hermkind 1977!. Elsewhere in the range, breeding activity is suggestedthat most of the larvae recruited to Florida may have maximal in the summer, although in areas to the south some been spawned in locations south of the Yucatan Channel. breeding activity is seen all year. Peak influx of postlatvae to These areas may have been as distant as the lesser Antilles or Florida does not occur until the spring of the next year, i.e., even Brazil Fig. 7!. A number of other workers Austin 1972; February, March, April, and into May. This pattern has been Hammer 1974! have conducted studies supplementing the observed by Florida Department of Natural Resources DNR! work of Sims and Ingle 967!. However, these studies do not researcherswho have devised a quantitative method to assess rule out the possibility of countercurrents or gyres acting to

Fisherfes, Vol. 5, No. 4 retardloss of larvaefrom theirspawning viciniiy. This hasbeen j jj'g'::: '~~QG~1 !'4l,$4;4:g"P:L'4!f;,4x;t:' ':» suggestedby Johnson 971, 1974!, who has studied the distri- r « bution of phyllosomesand pueruli of severalPacific species. Althougha large percentageof larvaemay be lost from the vicinityof the ,a significantnumber are retained by gyres and countercurrentscontributing perhaps predominantlv to localrecruitment. There are physical oceanographic data from a numberof areasindicating that this might be a viablehypothesis for at least some P. argus populations. In fact, the work of Brooksand Niijer 975! on the directionand velocity of the HoridaCurrent in the vicinityof the lower HoridaKeys during the summerdemonstrated a majorcounter current. Thus, lar- vae spawnedin the lower Horida Keys miqht be transported westwardinto the Gulf of Mexico.If they enteredthe eastern portionof the Gulf of MexicoLoop Current,it mightprovide a longenough path to allowtime for developmentto the puerulus '<~"m« oui"l"l-"'l"'v stageand a returnroute to the FloridaKeys. Fromthe foregoingit is clearthat the scopeof management 1 strategiescalled for would dependupon the patternof recruit- ment.At one extreme,management poficv would be strictlya local matter; at the other extreme, policy would be Pan- Caribbeaninvolving cooperahon of manynations. Further, dif- ferentgenetic consequences may also arise, depending upon thepatterns of larvaldispersal. If recruitment islargely a compo- Figure 6, Range of postlarval forms found on Witham habitats. site of many "upstream"populations, the geneticstructure- The upper four are the most common, Proceeding counter clockwise,the first is a clearpuerulus, a slightly pigmentedpost- i.e., allele frequencies of each population in the absence of larva, a darker brown postlarva, and finally an almost black larva. severelocal selectionpressures effect within the generation! The remaining two are generally classified juveniles and ace more shoulddiffer very little. On the other hand,if recruitmentfrom rare on Witham habitats, distantpopulations is minimal,gene frequencies in a particular

Figure7, The upperpart of the rangeof P. argus.Solid lines represent major current patterns while dotted lines depicta fewpossible local gyres. Lower case abbreviations arelocations ofanimal collections; b,Belize; im, Isla Mujeres; cs, Cay Sal Bank;vi, VirginIslands; dt, Dry Tortugas;kw, Key West;ek, ElliottKey; br, BocaRaton; bu, Bermuda.

July August 1980 45 In all the picture that seems to be emerging is that at least TH/ ~Q@S: ~g A 5kittztiesregelvel a 8:S.:." some population centers, e.g. Belize. may be set ofi' as distinct andyi! +S. 1n, btileqp inQQ@ M 1%/!@&t4tttIJn@ ': management units because of local hydrographic conditions. versiyc!f 'FIo're!i In 6aing~ii, '.tAf~ hii'i ~steit'e gaea On the other hand, areas such as the upper Hortda Kevs mav sm~ ~ conc~ted+IQ gleenmvernen$ Qfra@tttacttve " receive contributions of larvae from a variety of places depend- nuc!jete!.ough hajj. ' tory work scheduledfor this year and next will reveal the degree ' atnovi IJniyertlitIf.",Thispregrgm$k f&riariIy cog~ of contribution of these locales to Florida as well as the role of ' giOi.tIie appli~4ff, af bietdtyryg41'.and~+NL ap ~- the lower Cartbbean. When this study is completed we antici- 'f Mexico Proc Nat ShelthshAssoc 62 26 SO Brooks.f. H..anC P. O. Ntf let. tP7SThe Florida Curhater tanae inta the east pacific berne< toeeapoha. of detecting frequency differences between populations. Thus Pal about3 percent!have a proteinvaffant not presentin Hortda fernsot' larval rectumnent tnthe spiny lobster. Punvfuus atputa- t

Fisheries, Vol. 5, No. 4 and backwardswimming motions using abdominal pleopods, eluded that the puerulus has an average duration of 2'/2 and alsouse the rapid abdomenflex avoidancemotions com- months. This seems excessivefor several reasons. Nearshore monto juvenileand adultlobsters. Serfling and Ford975! studies have demonstrated regular, monthly arrivals of new re- clockedCalifornian Panulirus interruptus posfiarvae at 6 to 9 cruitment classesto the south Florida coast Little 1977!. Many cm/sec,and Phillips and Olsen 975! reported 30 cm/sec newlv arrived postlaivae from these studies have been main- maximuin and 15 cm/sec mean speeds for postlaivae of the tainedin aquariaunder conditions not favorablefor settlement, WesternAustralian P. longipescygnus. Swimming by Carib- yet all have progressedto the first juvenile stagewithin two beanP. arguspostiarvae has also been studied Calinski un- weeks Witham et al. 1968; Sweat 1968!, well before the amval pub.!.Swimming by all threespecies is directional,steady, and of the next recruitment set. Since late stagelarvae passrelatively not necessarilyrelated to currentdirection. Nearly all swimming near the south Florida coast via the Gulf Stream, it woulct be postlarvaehave been found in the top severalcentimeters of expectedthat some captivepostlaivae would remain in that surfacewaters, perhaps explaining the inabilityof planktonnets stagelonger than two weeksif their normalperiod was more and trawlsto samplethis stage adequately. Palinurid posflaivae than two months. Furthermore, there is no evidence indicating usuallyswim forward, extending the antennaestraight ahead that feedingtakes place in the postlarvalstage, despite gut with tips nearlytouching and folding the tail fan to minimize examinationsand aquarium observations, Two or threemonths waterresistance. Encountering abrupt light change shadow!or seems a very long period to expect such busy, energy- touchingobjects with the antennaetriggers an alarmresponse in consumingorganisms to operatewithout refueling, The two- to whichpostlarvae spread their antennaeto an angleof approxi- three-weekperiod suggested by Sheard949! for postlarvalP, mately60; expandtheir tail fan, and remain motionless or sink longipes cygnus seems more likely. slowlydownward to asdeep as 20 cmbefore resuming surface Whatever the sources of mechanisms,there are large quan- swimming.Postlarvae of severalspecial of Scyilaridae,including titiesof postlarvaerecruited to shallownursery areas adjacent to two speciesof Scy/landes Lyons 1970!, have been collected majorspiny lobster fisheries throughout the world. Most likely, swimmingin surfacewaters, but no observationsof swimming the shallows and shorelines act to concentrate the relatively techniquewere reported. Based upon specimens in sparselydistributed postlarvae immigrating from the oceans. tows,Lyons 970! rejectedRobertson's 968! contentionthat Oncesettled, postfarvae soon take on the pigmentationof the postlarvaeof Scyllarusamencanus did notswim. developingjuvenile and the concealmentstrategy ln whatmay amount to an oversimplification,the oldest,inost changesfrom invisibilityto camouflage.Curiously, no major developedlarvae are usually found the greatest distances from aggregationsof settled postlarvae have yet been shore in the oceaniccirculation systems off westernCentral found. American and Western Australia. Postlarvae of many scyllari- Thereare many reportsof scyllarideanpostlarvae collected dean specieshave been collectedoccasionally from oceanic from fouling assemblageson suspendedobjects such as trap plankton Lyons 1970; Johnson 1974; Serfling and Ford 1975!, lines,buoys, boat bottoms, and mangroveprop roots Lewiset but most, perplexingly,have been found near coastalareas. al. 1952; Lyons 1970!. "Floating habitat" collectingdevices This causedJohnson 971! to speculatethat mostlarvae off haveapproximated such situations, in Floridasimulating and/or westernCentral America may exist in deep,near-coastal bottom servingas substratumfor fouling communities Witham et al. waters as yet unsampled!,from which the postlarvaelater 1968; Sweat 1968; Littie 1977!, in southern California consist- emerge,or the larvaemay be caughtup in a systemof coastal ingof naturalsurf grass in cages Serfling and Ford 1975!, and eddies and countercurrents far more prevalent and complex in Western Australia consisting of "artificial seaweed" Phillips thanimplied by general studies of thehydrography. Presence of 1977!. These collectingdevices have enabled repetitive, quanti- significant quantitiesof postlarvaein stomachsof pelagic flablesampling of posflarvalrecruitment, resulting in recognition oceanicfishes such as yellowfintuna Michel 1968!, however, of several similar behavioral traits and some apparently signifi- do little to support these theories. cant differences between species. Phillipset al. 978! notedthat at theend of theplanktonic Bemusement by or attraction to underwater lights by surface existence,the water circulationof the easternIndian Ocean swirnrningpostlaival spiny lobstershas been noted in California transportslarvae of P, longipescygnus back toward the coast of Serfling and Ford 1975! and Western Australia Phillips and WesternAustralia. Upon encounteringcontinental shelf waters, Olsen 1975!. ! made similar observations during comparable the larvae metamorphoseinto pueruli, which swim 40 km studies in the Lesser Antilles during 1977 Calinski unpub- across the shelf to settle in inshore shallows. Fewer postlaivae lished!, and must conclude that, although postlarvae were re- are found in surfacewaters than at lower depths over the shelf, vealed by the light, they showed no particular attraction to it; butthe postlarvae rise to thesurface just prior to settlement.In instead, they passedlinearly through the cirde of light without south Rorida, washedby severalcurrents from the Gulf of varying course or circling. However, both Californianand Mexico and , larval transport may be more com- Western Australian studies have shown that suspendedcollect- plex.A theoryof predominanflyCaribbean recruitment of lar- ing devicesproduce far better catcheswhen lighted Phillips vae to Florida via the Yucatan Channel, based upon high inci- 1975a; Serfling and Ford 1975!. denceof larvaemoving northward in that system ingle et al. Virtual!y all settleinent of postlarvae on collectors occurs at 1963;Sims and Ingle 1966!,has recentlybeen challenged by night, with recruitinentoccumng during the dark moon new indicationsthat Rorida stocksmay differ geneticaflyfrom those in moon, first quarter! phases Lyons 1970; Little 1977; Phillips the Caribbean see Menziesand Kemgan, this issue!. 1977!, Serfling and Ford found no lunar periodicity in California Like manyother aspects of postlarvallife, the durationof the posflaivae, but their catches were quite small compared to postiaivalstage has been subject to conjecture.Serfling and Florida and Western Australian figures. According to Phillips Ford 975! calculatedmean swimming speed of P, interruptus 975b!, postlarvaecease settling on collectorswhen moonlight postlarvae,related that to distantareas where Johnson 960! intensity rises above about 10 percent full moonlight. Little found most late stageP. interruptusphyllosomes, and con- 977! also noted that no recruitment occurs during early night

Fisheries, Vok 5, No. 4 hours of the latter part of the full moon phase, even though ACKNOWLEDGMENTS complete darknessmay prevail. These observationssuggest that light ifltensity alone is not a factor in encouraging postlalval I am especially grateful to Michael Calinski, Kev West, settlement, particularly considering the enhanced success of Rorida, who allowed me to observed his studies of swimming lighted collectors. Little's 977! suggestionthat lunar influence postlarvalPanulirus argus, thereby granting me many new in- sights to the behavior of these fascinatingcreatures. I have also on tides and current patterns could affect settlement rates may merit further investigation. Again, however, Phillips 977! drawn to a limited extent from unpublished data recently col- found no relationship between the volume of water flowing into lected by E. J. Little, Jr., of the Florid Department of Natural a collectingarea and the catcheson collectors, either as a cvclic ResourcesMarine ResearchLaboratory. flow related to moon phase or from night to night at the time of setfiement. No relationships have been found between densities of P. long pcs cygnus postiarvae and surface water temperature of salinity Phfilips 1977!. In both the Californian and Western Australian popualtions, peak postlarval settlement periods have been confined to spring and summer Phillips 1977!. In Florida, where year-round recruitment occurs, seasonal recruitment LITERATURE CITED peaks may vary between collecting areas and between years Berry,P. E. 1974.Palinurid and scyUand lobster larvae of theNatal coast, South Africa. OceanogrRes Inst. IDurbanl Invest Rep 34 44pp Little 1977!, but are usually characterizedbv greatest settle- Chittleborough,R.G., and B. F. Phigips.1975. Fluctuations af yearwlass strength and re- ment during spring, with slack summer influxes followed by a cruitmentrn thewestern rock lobster Ponuiirus iangipes IMilne-Fdwards! Aust J Mar. fall increasenot so great as the spring setfiement. Lewis 951! FreshwaterRes 26 317 328 Ingle,R. M., B. Eldred,N. W. Sims, Jr., and E. Eldred.1963. On the possible Caribbean reported peak recruitment to south Florida during January originai Rorida'ssptny lobster populations Fla Bd.Consenr. Tech. Ser. Na. 40 12pp. through March, and extraordinarily great numbers of postlarvae Johnson,M. W. 1960Production and distribution of larvae ai thespiny lobster, Ponuii rue setiied on collectors in the Florida Keys immediately following tntenvptus Randagl. with records on P. grndksStmets. Bug ScrippsInst. Oceanogr 716k413462 the severe cold period of January 1977 Little unpublished!. 1971The palrnurid and scygarid lobster larvae of thetropical eastern Pactgc and their However, a similar cold period farther north along the Florida distnbuttonasrelated ta the prevathng hydrography. Bull. Scripps Inst. Oceanagr. 19:1 36. 1974On the dispersalaf lobsterhrvae rnto the EastPacific Bamer IDecapada. coast killed juvenile lobsters and was followed by decreased Palinuridae!.Fish. Bull. 72I3! 639 647. settlement of postlarvae during 1966 Lyons unpublished!. As Kanciruk,P.. and W. F. Herrnktnd, eds. 1976. An indexed bibliography of the spiny lobsters. Little 977! observed, greatly reduced salinities because of familyPaltnundae. Ra. Sea Grant Rep. No. 8 101pp. Lewis,J. B. 1951.The phygosoma larvae ai thespiny lobster, Panuieus argus Bull. Mar. Sd. seasonally related rainfall drainage have disrupted postlarval GulfCaribb. I I2!:89 103. recruitment in some south Florida estuaries. Otherwise, Little Lewis,J. B.,N. B. Moore,and W. Bahts. 1952 The pasUarval stages of thespiny lobate~, also found no relationship between recruitment magnitudes, Panuiirusargus. Bull. Mar Sci. Gulf Caribb. 2 I:324 337 Little,E. J., Jr. 1977Observations onrecruitment afpostlarval spiny lobsters, Panuiirus argus, and salinity and temperature. ta thesouth Ftarida coast. Fla. Mar. Res. Publ. No 29.35 pp. Frequent and widespread attraction of postlarvae to sus- Lyons,W. G. 1970 Scygand lobsters ICrustacea, Decapoda I. Mem. Hourglass Crursru, Val. I, Pt IV 74pp pended collectors in inshore waters suggeststhat these devices Mensies,R,A,. and J. M.Kerrigan. 1980 The larval recruitment problem oi thespiny lobster. may be usefultools for assessingcertain aspectsof lobster popu- Fishenes5I4!:42 46 lation dynamics, particularly those related to predicting fishery Michel,A. 1968Les iarves phyllasomes etIa post-larva deScyiiarides squnmosus H. iMilne- Edwards!Scyraridae lcrustaces decapodesl. Cah 0 RS T0 M, Ser.Oceanogr. 6. success Serfling and Ford 1975; Little 1977!. Postlarvae are 4k47 53. certainly more available and quantifiable than are larvae. Phil- Phillips.B. F. 1972A semi-quantitativerogector of thepuerulus larvae ai thewestern rock lobsterPano/irus iongtpes cygnus George IDecapoda, Palinuridae! Crustaceans lips and Hall 978!, summarizingobservations from collectors 22:147 154 monitored over periods as great as nine years, concluded that 1975aEffect of nocturnalillumination oncatches ofthe puerulus Imvae of the western all collectors at a site indicated similar trends in the level of rocklobster by collectorscomposed of artifidalseaweed. Aust. J. Mar.Freshw Res 26 411 414 monthly and yearly settlement, and that consistencyin indica- 1975bThe etfect of watercurrents and the intensity of moonlighton catches of the tions of settlement fluctuations were also obtainable between pueruluslarval stage of the western rock lobster. CSIRO Aust. Div Fish.Oceanogr Rep. Na. sites. These consistent fluctuations agreed with other data on 63 9pp 1977A reviewof thelarval ecology of rocklobsters. Pages 174 185in B.F. Phillips periodicity of late stage larvae off the WesternAustralian shelf. andJ. S. Cobb,eds. Workshop onlobster and rock lobster ecalogy and phystalogy CSIRO Three years of field studiesjust concluded in the Florida Keys Div Fish.Oceanogr Circ. No 7 Litfle unpublished! indicate similar collector reliability for post- Phillips,B. F.,and N. G.Hall. 1978. Catches otpuerulus larvae on collectors asa measureof naturalsettlement of the urestem rock lobster Panuiirus cygnus George. CSIRO Div Fish larval setflement of Panulirus argus. However, Chittleborough OceanogrRep No 98 18pp and Phillips 975! concluded that, owing to restricted holding phillips.B, Fand L Olsen,1975 Swimming behavior ofthe puerulus larvae of the western rocklobster Aust. J Mar Freshw.Res 26415 417 capadty of shallow nursery reefs, mainly due to limited food PhtUipa,B. F., D.W. Rimmer.and D. D, Retd.1978 Ecological investigations oi the Iate- supply, density-dependent mortality reduces most year-class stagephygosoma and puerulus larvae ot thewestern rock lobster Panuiirus longipes cygnus strengthsto similar levels by the time lobsters approach matur- MarBiol 45347 357. Robertson,P.B. 1968.The complete larval development of the sandlobster, Scyiiarus ity. Phillips and Hall 978! were also unable to detect a rela- amenmnusiSmith!, Decapoda, Scyraridae! in the laboratory, with notes on larvae hom the tionship between levels of catches of postlarvae on collectors plankton.Bull Mar.Sci 18I2!294 342. and subsequentjuvenile densities,speculating that the relation- Serfling,S.A., and R. F. Ford.1975. Ecological sturkes oi thepuerulus larval stage of the Cahfomiaspiny lobster, Pano liras interruptus.Fish. Bull. 73!:360 377. ship may have been maskedby density-dependentnatural mor- Sheard,K. 1949The manne craylishes spiny Iobsterst, family Paltnundae, oi Western Aus- tality. Is there a direct relationship between numbers of settling tralia,with particular reference to the Ushery on theWestern Australian crayllsb Ponuiirus iongipesi.CSIRO Bull. 247. 45 pp. posfiarvae and later size of the fishery population, or is some Sims,M. W., and R. M. Ingle. 1966. Caribbean recruitment of Florida's spiny lobster popula- other "biological filter" at work farther along in the fife cycle? tion Q J Ra Amd Sci29! 207 242. This question must be solved before a predictive monitoring Sweat,D. E. 196S.Growth and tagging studies on Panuir'rus argus fLatregle! in the Florida KeysFla Bd.Conserv Tech Serv No. 57. 30 pp system for fishery management based upon postlarval setfie- Witham,R,. R. M, Ingle.and E. A. Joyce.Jr, 1968Physiological andecological studies of ment can be inaugurated. Panuitrvsargus from the St. Luae estuary. Ra Bd ConsenrTech. Ser. No 53.31 pp.

July August 1980 49 LOBSTER FISHERY MANAGEMENT UNDER THE FISHERY CONSERVATION AND MANAGEMENT ACT

J. R. Zuboy, A. C. Jones, and T. J. Costello

ABSTRACT TheFishery Conservation andManagement Aciof 1976 PL 94-265! created a new era in fisheryrnanageroent, featuring developmentof comprehensive fishery management plans by regionalfishery management councils and management for "optimumyield." In this paper we discuss the main features ofthe Act, consider the development offishery management plans in general,and brieflydescribe roanagement plans for lobsters.

carrying out a Fishery Management Plan FMP! may result in transfer of managementresponsibility in state waters to federal authority. Aspectsof the FCMA have been discussedat the Symposium on the Fishery Conservation and Management Act of 1976 Washington Law Review, v. 52, no. 3, July 1977!; Optimum Yield Workshop Houston, Texas, June 1977!; and Limited Entry Workshop Denver, Colorado, July 1978!. Brief descrip- tions of the Act have been prepared by Browder 977! and O'Conner 978!. The National Standards for fishery managementspecified in the Act help to ensurethat managementmeasures are objective and equitable. The required contents of FMP's include an esti- James R, Zuboy A. C, Jones mate of the maximum sustainable vield MSY! and the optimum heCongress of the United vield OY! for each fishery. T States established author- MSY is defined as "the largestaverage catch or yield that can ity under the Fishery Conserva- continuouslv be taken from a stock under existing environmen- tion and Management Act tal conditions" Ricker 1975!. Where sufficient scientific data on FCMA! to conserve and rnan- the biological characteristicsof the stock do not exist, or the age the fishery resourcesfound period of exploitation or investigationhas not been long enough off the coasts of the United for adequate understandingof stock dynamics, the MSY will be States. One goal of the Act was estimated from the best scientific information available. The re- to provide a meansof unilateral quirement to estimate MSY on the basis of the best available management to replace the scientific information allows the first generation of FMP's to be then current system of multi- completed and managementto begin. Further scientificinvesti- stateregulations and multilateral gations will improve the data base and the estimatesof MSY, fishery agreements.The United and allow managersto choose more sopnisticatedmanagement States assumed exclusive T. J. Costello measuresif they believe these are warranted. fishery management authority OY is defined in the Act as "the amount of fish which will over all fish, except highly migratory species e.g.. tunas!, in a provide the greatestoverall benefit to the Nation, with particular FisheryConservation Zone FCZ!extending from the territorial reference to food production and recreational opportunities; limits of the United States to 200 nautical miles offshore. Eight and which is prescribedas such on the basis of the maximum RegionalFishery Management Councils. consisting of state. sustainablevield from such fishery, as modified by any relevant federal, and private representatives,were establishedto prepare economic. social, or ecological factor." Optimum yield has not fisheryrnanagernent plans for eachfishery in accordancewith been reduced to a more specific definition nor have proce- national standardsand objectives specified in the Act, dures for its quantification been specified. OY remains an elu- Regulatoryauthority under the FCMAis limitedto the FCZ; sive concept that can be shaped to meet the objectives of the however, fishery resourcesare mandatedto be managed user. Some persons believe that lack of a precise definition throughouttheir range.The Councils,whose membership in- facilitates the decision process in accomplishing the "best use cludes the principal state officials with marine fishery manage- of the resource." Other persons would prefer to see more rigid ment responsibility, are constituted to help ensure that man- procedures specified for estimating the value of optimum agementis consistentin the FCZand in stateterritorial waters. vield. Our opinion is that Congress wisely provided flexibility In addition, state action or inaction that adversely impacts on in interpretabonof this mainpurpose of the Act. Benefitsfrom

50 Fisheries, Vol. 5, No. 4 agementas specified in theFCMA, and its development by the minimum size limitation on the basis of the best scientific infor- Board was endorsedby the New Englandand Mid-Atlantic mationavailab!e; ! developregional programs to controllobs- FMC's. terfishing effort and regulate lobster fishing mortality rates; ! In this paperit is beyondour scopeto discussthe detailsof implementuniform collection, analysis, and disseminate biolog- eachFMP. We just mention the specific objectives of eachplan ical and economic data; ! increasebrood stock abundance to andthe generaltypes of managementmeasures proposed to minimizethe risk of stockdepletion and recruitmentfailure; ! meetthe objectives.For furtherdetails, please write directlyto minimizelobster injury and mortalityassociated with fishing;! the group responsiblefor the plan of interest addressespro- developstandard gear-marking procedures to the extentpracti- cal; and ! maintainexisting social and culturalfeatures of the vided below!. industry wherever possible. A variety of managementmeasures are spedfiedto meet Caribbean theseobjectives: gear restrictions, size and sexrestrictions, per- The Caribbeanspinv lobster FMP identifiesfour objectives: mit requirements,effort limitation, and reportingrequirements. ! providefor biological conditions consistent with the ability to achieve a maximum sustainable yield; ! promote economic CONCLUSION efficiencyof the commerdalfishery; ! providefor thesocial and cultural needs of Puerto Rican and U.S, Virgin islands' The FCMAhas provided the impetusfor effectivemanage- citizens;and ! providebiological, economic, and socialdata ment of our marine fisheriesresources. The Act has also created basesfor future management of the resource. anunprecedented need for cooperation between state and fed- The followingmanagement measures are establishedto at- eralfishery management agencies because fish stocks do not tain the objectives:size and reproductivecondition regulations; recognizepolitical boundaries, The lobsterfisheries, which are closedareas; improved data collectionsystems; a self-destruct prosecutedin bothstate and federal fisheries jurisdictions, pro- panelrequirement for trapsand pots; owner identification and vide an excitingchallenge for contemporaryfishery managers. markingrequired for traps,pots, buoys, and boats;prohibition of theuse of poisons,drugs, or otherchemicals, spears, grabs, ADDRESSES hooks, or similar devices for taking spiny lobsters; encourage- ment and stimulation of investigationin relation to the construc- Caribbean Fishery Management Council tion of underwater habitats to attract lobsters. Suite 1108 Banco de Ponce Building Gulf of Mexico and South Atlantic Hato Rey, Puerto Rico 00918 The followingfour potentialmanagement objectives are pro- Gulf of MexicoFishery Management Council posedin theGulf of Mexicoand South Atlantic plan, subject to The Lincoln Center adoptionby the Councils:! projectlong-run yields and pre- Suite 881 ventdepletion of lobsterstocks by assuringadequate larval pro- 5401 West Kennedy Boulevard duction;! maximizeyield taken in thefishery given recent and Tampa, Florida 33609 anticipatedfuture levels of fishingeffort; ! developmethods SouthAtlantic Fishery Management Council for effectivelyenforcing provisions of the managementpro- Southpark Building gram;and ! developregulations that conform,to the extent Suite 306 practical,with a! existingstate laws by whichthe fisheryis now 1 Southpark Circle regulated, b! practicesand laws of othercountries within the Charleston, South Carolina 29407 Caribbean,and c! currentmethods and practices in the fishery. Recommendedmanagement measures in the plan include WesternPacific Fishery Management Council sizelimits, season and arearestrictions, gear regulations, recre- Room 1506 ationalbag limits,protection of berriedfemales, import restric- 1164 Bishop Street tions, and permit requirements. Honolulu, Hawaii 96813 Northeast Marine Fisheries Board WesternPacific Leverett Saltonstall Building The Western Padfic FMC plan specifies four major policy Government Center objectivesfor managingspiny lobstersin the northwestern 100 Cambridge Street HawaiianIslands. They are to ! lowerthe risk of depletionof Boston, Massachusetts02202 the stock;! acquireas much informationas possibleon the fishery;! maximizeeconomic efficiency; and ! avoidunfa- LITERATURE CITED vorableimpact on monk sealsand the NWHI WildlifeRefuge. Baker,J. s., J.s, Kamlntt,and R. E. tttarrtean,1977. Enatranmentat impact statementr A A finalset of regulationswill be developedfrom the broad guideta preparattanand review. Practicing Law institute, New York, N.Y. 334 pp. categorieswhich follow; size and seasonrestrictions, protection Brawder,J. A. 1977.nte FnhetitAct af 1976.Sea Grant Spec. Rep. Na. i2. Univ.Miami, Miami,Ra. 16 pp. of egg-bearingfemales, quotas, time-area restrictions, limited Costello,T.J., and L M. paine.1979. How ta preparettshety management plans. teOAA entry and/oreffort, permits, logbooks, observers, and gearre- TechnicalMemorandum NMFS Na 4 SoutheastRrhertea Center, Miami, Fla. 62 pp. strictions. O'Canner,C.R. 197S.The Fishery Canaereatlan andManagement Actaf 1976.The need tar stateand iederal cooperation. sea Grant spec. Rep. Na. 13, Univ. Miami, Miami, Fla. 13 pp. Rlcker,W. E. 1975 Computation and interpretaaan af biological statistics ai 8rh papalattans 1tlorthwestMarine FisheriesBoard NMFB! Flrtt ReaBoard Can BaB. 19i. 38Zpp. The NFMB has adoptedthe followingmanagement objec- tivesin its planfor theAmerican lobster fishery: ! adjustthe

Fisheries, Vol. 5, No. 4 52 ECONOMIC CONSIDERATIONS IN THE MANAGEMENT OF THE FLORIDA SPINY LOBSTER FISHERY

Fred J. Prochaska and James C. Cato

researchin numerousdisciplines as thev apply to the spiny lobsterresource. The purposeof this paper is to discusshow conceptionaleconomic ideas pertaining to the managementof the fisherymay be appliedand to relav empiricalresults from economicmodels that have been estimated for the Floridaspiny lobster indushy.

ECONOMIC MODELS Economicanalysis can be appropriatelvdivided into three basicfunctional areas, The first is the industryproduction func- tion or the bioeconomicmodel, which addresses the questionof maximum economic yield MEY! and optimum allocation of Fred J. Prochaska James C. Cata resourcesfrom a totalindustry perspective. The secondtype of uring the past two decades tremendous growth in the analysisis often referredto as the "firm" productionmodel, D spiny lobsterfishery in Horida hasoccurred. Landings which addressesthe questionsof efficiencvand profitability increasedfrom lessthan one million pounds before the 1950's from the view of the individualfishing firm. The third type of to about 11 million pounds between 1972 and 1974. Part of economicanalysis is concernedwith marketing,consumption, this growthin landingsresulted from U.S.fishermen fishing in and demandfor the product.An additionalcategory can be foreign waters.The Bahamashave been the major foreign definedas eitherindustry or systemsanalysis. This fourth cate- water source and in the early 1970's accounted for as much as gory combinesthe separateareas of analysesand addresses 80 percentof foreignwater landings. Between 1969 and 1975, broadindustry and managementprograms. The bioeconomic spiny lobster landings from U.S. waters accounted for 42 73 modelis the principalsubject of this paper.Uses of the other percent of U.S. landings. economicmodels in fisherymanagement will be briefly dis- At the beginningof the 1975 76 season,the Bahamian cussed in the concluding section. government banned foreign lobster fishing. The effect of the ban was that landings from foreign watersin 1976 and 1977 are Bioeconomic Model about one-thirdof the level in previousyears. The trend in Bioeconomicmodels are estimated bv relatingresultant yield landingsfrom domesticwaters gradually increased until a 1974 to fishingeffort over a period of time or productionseasons. peak, after which landings from domestic waters have de- Yield is usuallymeasured as eitherlandings or catch.Effort in creased.This paper uses models that referto landingsfrom the theoreticalliterature and in manvapplied statistical models is domesticwaters. Specifically, landings from Monroe County, normallymeasured as an indexof inputs such as fishermen, Florida Florida Keys are used since almost all domesti- vessels,or gear units.The index allows for adjustmentsin callvcaught and landedlobsters occur in that county.Most technology,scale of operation,etc. For example,the number of Bahamianlobsters were landed in DadeCounty. fishermenmay be the basicunit of effortbut thiseffort unit may Increasedharvesting resources devoted to the fisheryhave changethrough time because of changesin gear units used per causedeffort to increasesubstantially more than landings. This fishermen.The technological efficiency of eachunit of gearmay hasled to decliningaverage productivity or decliningcatch per alsochange over time; that is, newor differenttrap styles may unit of effort. Decliningcatch per unit of effort aloneis no indica- "catch"better than previousstyles. Indexing inputs adjusts for tion of the well beingof the stockor industry.Dedining average these factors so that "effective" units of effort are measured and marginalproductivity occur in any productionprocess be- throughthe time periodof analysis.An alternativemethod for foreeither maximum physical or maximumeconomic produc- adjustingthe effort variable s! is to include the individual items tion occurs.However, the extentof the decliningcatch per unit in a statisticalmultiple-regression model, The regressionproce- of effort and the apparentdedine in totalproduction from dure essentiallyaccomplishes the samething as indexingfor domestic waters have been considered a sufficient reason for standardizingunits of effort.However, the latter approachhas concernand justification for a fisherymanagement program and theadvantage of being more directly applicable to ananalysis of

July August 19SO 53 proposedmanagement programs, since specific variables analyzedare thosesubject to regulationsand are basedon 00 theoryapplicable to the fisheiy. The estimatedparameters in the bioeconomicmodel that show the effect of different levels of effort on catch are usually statisticallyconfounded. This means that the parameters reflect both ! the effecton landingsbecause of stocklevel changes as a result of fishing effort and ! the returns to proportion re- flectedin anyproduction process. Returns to proportionsrefers to increasingor decreasingproduction per unit of variableinput dueto someother fixed inputs used in the productionprocess.! 7!i eeech Ideally,variables measuring stock in eachtime period would be includedso that the separateeffects of effort on landingscould be determined.Unfortunately, these data are not normallyre- corded on a continuous time seriesbasis. Environmental vari- ablessuch as watertemperature are often includedas altern- ce!00 tives for the stock variables.Water teinperature reflectsto some degreeenvironmental conditions that affect the stock as well as seaconditions for somefisheries, particularly the spinylobster. 100 303 300 000 500 500 700 0 Tressere firn A bioeconomic model estimated for the Monroe County Figure1. Averageand marginalcatch of lobstersper trap with spinylobster fishery provides an example of theseeffects as well 399 firms in the industry. as how the model can be used to answer management ques- tions,' Firmsin MonroeCounty, Florida,account for 80 90 The marginaleffects of landingsof increasingor decreasing percentof Floridadomestic landings in recentyears.' Monroe trapsper firm or numberof firmsin theindustry are determined Countywas thuschosen as the areafor study. by takingthe partialderivatives of equation!. Averagecatch At the time the followingmodel was estimated in 1976,time perunit of effortis determined by solvingequation ! for vari- seriesdata were available for the 1963 1973 time period. The ousunits of T andF andthen dividing by the numberof unitsof parametersand standard errors estimated with the modelare effort. presentedin the following equation: The effectof increasing or decreasing!the numberof traps fished when 399 firms are in the indushy is shown in Figure 1. Y = 28.38 1,439,98 465.17 .24 E ! Marginalcatch per unit of effort declinesthroughout the range T F of trapsconsidered, while averagecatch per unit of effort in- 65.9! 16.5! .2! creasesat first and then declines after approximately 350 traps are fishedper firm.' It is importantto note the differencebe- where tweenaverage and marginalcatch per unit of effort traps!.For example,if eachfirm adds the 600th trap, the additional yield Y annual landings' of lobstersin million pounds in for thattrap would be approximately10 pounds; whfie at 600 Monroe County, 1963 73 trapsthe averagecatch for eachtrap is approximately26 T number of traps fished per firm pounds.The management implication is thata regulationthat F = total number of firms in the industry requiresa reductionin effortwill havesubstantially less impact E = seasonalmean surface water temperature measured thanexpected if projectionsare based on averagecatch per unit in degreesFahrenheit. of effortestimates, In the exampleof eachfirm fishing600 traps, a reductionin industrytrap usageof one trap per firm for each VariablesT, F, andE explained80 percent R' .80! of the of the 399 firmswould reduce industry catch by 3,990pounds, annual variation in lobster landings. The three variables were not the reduction of 9,975 pounds suggestedwhen average statisticallysignificant at the99, 90,and 80 percentconfidence catchper unit of effortis considered.The differencein the two levels,respectively. Based on 1973 reportedlevels of firms estimatesresults becausethe average catch for each of the re- 99!, trapsper firm 29!, andmean temperature 7.591' F!, mainingtraps increases when the 399 trapsare removedfrom the equationsolves at the meanfor 5.2 millionpounds. the industry. Similar conclusions are drawn when consideration is given to 'Completediscussion of methodology, assumptions, data base, and increasingor decreasingthe number of firms in the industry conclusionsof the bioeconomicmodel are presented in Williams1976, Fig.2!. Afterapproximately 150 firms have entered the indus- The models are being updated with current data, and alternative try, averagecatch per firm beginsto dedine.The dedineoccurs mathematicalspecifications are beinginvestigated to reflectcurrent in- becauseless stock is divided among a given number of firms in dustry conditions. the short run and the occurrence in the long run of a negative 'A firm is definedin this studyas a lobstervessel and all the inputs necessaryto catchlobster. See Prochaska and Williams 1976 for budget information on lobster firms, 'Thisproduction function operates throughout its range where output 'Catch is normallyused as the yield measurein any analysisthat is increasingat a decreasingrate. This is normallydefined as the rational estimatescatch per unit of effort.Landings are used in thisanalysis since economicstage of productionfor the firm, lt allowslandings to ap- the analysiscovers one entire geographicarea Monroe County, proacha maximumbut does not allow them to decrease.Rorida's Florida!and almostall lobsterscaught in the countyare landed in the regulationsprovide for minimumsize limits and prohibittaking or strip- county.All parametersare used on a countybasis and thus catch and ping egg-beafing females. Both of these regulations assure minimum landingsin this caseare virtuallyidentical. stock levels and justify the form chosen.

Fisheries, Vol. 5, No. 4 effecton stocksresulting from increasedfishing pressure, A re- ductionin the numberof firmsfishing for example,through a limited entry program! would causea much smaller catch reduc- tion than is suggestedby observingaverage catch per unit of effort firms!. A reduction of one firm at the 1973 level of 399 firmswould reduceindustry catch by lessthan 3,000 pounds, not the approximate13,000 pounds suggested by the average catch per unit of effort estimate. See Williams and Prochaska 1977 for a more complete discussion. Beforeconsidering the maximumeconomic yield for the in- dustryit is appropriateto notethat the catchper unit of effort estimates for number of firms and the estimates for the number ro of traps are not independent. At current effort levels, a decrease in traps fishedper firm will shift the catchper firm estimates upward in Figure 2. Likewise, a decreasein the number of firms wfil shift the estimatedcatch per trap functionin Figure1 up- ward. Revenuesand costsmust be consideredto determine the level of effort in the industry consistent with maximum eco- nomic yield. The maximum economic yield MEY! level is de- terminedwhere the marginalrevenue resulting from an addi- 1!! i GOJ 3OO 4OO SGG G !'! 7'! ! .'OG I !r'!s tionalunit of effortis equalto the marginalcost of the additional Figure2. Averageand marginal catch of lobstersper firm when unit of effort.Marginal revenue is simplydetermined by multi- each firm Ashes429 traps. plying the marginal catch per unit of effort functions shown in solution implies that a considerablereduction in the number of Figures1 and2 by the priceof lobsters.To determinemarginal firmsin the industrywould be necessaryto achievemaximum cost per unit of effort equation ! was estimated, economic efficiency, However, each firm would fish consid- erablymore traps. This reorganization would decreaseindustry C $1,876 + $11,55T ! coststo $2,355,407from the 1973 costlevel of $2,725,549. .17! Savings measured in reduced costs are $370,172. These sav- ings of approximately14 percent must be comparedto the where socialcost of displacing186 firms from the industrybefore an C = annual total cost per firm "optimum" yieldcan be determined.In addition,the redistribu- T number of traps fished per firm. tion of incomemust be considered.Predicted profits would be $18,350per firm compared to $6,677before the reorganization Theestimated cost equation used data from a sampleof 25 of the industry. lobsterfishermen Prochaska and Williams1976!. Equation ! explained75 percent R' = .75! of the variationin costsamong Other Production Models firmsand wasstatistically highly significant. Fixed costs per year attributableto lobsterfishing are estimatedto be $1,876,while Economic models estimatedin equations ! and ! can be marginalcosts of fishingan additionaltrap are estimatedto be madecurrent by simplyusing current lobster prices in the reve- $11.55per trap. nue calculationsand multiplyingthe cost estimatesby an ap- Since yield per trap is not independentof the number of propriate cost index. However, if substantial changes in fishermen firms! in theindustry, MEY must be determinedby technology and/or fishing effort have caused population stock simultaneously considering variafions in both the number of levelsto vary,it will be necessaryto makenew physical produc- firmsand the numberof trapsfished per firm. This is done by tion parameter estimateswith current data. If overfishing has maximizingthe industry-profitfunction described bv equabon occurred or regulations on which the model assuinptionswere ! with the temperature effect added to the constant. based have changed,it would be necessaryto formulatea mathematical model that allows for the occurrence of a Profit Total revenue Total cost ! maximum catch. Profit = P! Y! ] = [ F! Cost per firm! ] The bioeconomicmodel estimated in equation! is not ap- g P 9,773,481 1,439,976,169 465,173,997 propriate for use in analysis of some economic problems if F $1,876 + $11.55T! managementregulations insure that stock will be maintained at somedesired level, For example,the 1974 catchappears to where have been the maximum from the fishery. If management measures were instituted that ensured future effort levels would docksideprice of lobstersper pound PY not affect the stock, economic models would have to be esfii- predictedpounds landed for givenlevels of F andT mated from data that reflects the fixed stock level. Since the F total numberof firmsin the industry stocklevel and effectsof fishingpressure on stocksare likelyto number of traps fished per firm have changed over the time period from which the data were industry profits generatedfor the bioeconomic model, the model would not be Docksideproduct prices in 1973were $1.08 per pound.At this appropriate to answer economic questions for one stock level. price the profit maximizingsolution is 795 trapsper firm with 'Thesolution is determined by setting the first partial derivatives of g 213firms in theindustry.' Substituting these values into equa- with respectto T andF equalto zero,and solvingthe setof simultane- tion ! givesan MEYof approximately5.8 millionpounds. This ous equations for T and F.

July August 1980 55

in 1929 resulted in establishmentfor the Rorida Panulirus argus are also caught with adult, legal-sizedanimals in traps without fishery of a one-pound minimum size limit, which was sub- escapevents Bowman 1963!, One consequenceof even minor sequentlv reduced to a three-inch carapacelength size. dainage is significantly reduced growth rates Davis 1978 and Fifty years later, it appears that the 1929 limit would have 1979!, which in turn means a reduced yield per recruit as the protected many animals through their first reproductive season, growth curve is prematurely flattened. whereas the present limit does not Davis 1975!, The reduction Fishery managementactions to improve growth rates include in minimum sizeto below sizeof maturity probably reflected the escapevents, prohibifions on possessionor transport of shorts, growing influence of the market in determining "optimum" size and establishment of nursery sanctuaries, particularly where during the 1950's, and the suggestionthat larvae recruited to recreational diving has been a significant part of the fisheiy the Florida fishery originated in the Caribbean Ingle et al. 1963; pressure,Any of these actions,by reducing the time required to Sims and Ingle 1967!. While the growth of the Rorida fishery reach the minimum harvest size, will reduce mortality and in- after the reduction of the minimum harvest size below the me- crease YPR. dian size of maturity could be taken as evidence that size of maturity is not a necessary consideration in setting minimum Mortality harvest size, it more likelv reflects the geographically marginal Reducing mortality is a corollary of improved growth rates. position of the Florida fishery, ln the Florida fishery, density- The major problem of reduced growth rates resulting from in- independent factors acting on the planktonic larvae appear to juries is the subsequentincrease in mortality incurred before the be more important Sims and Ingle 1967; Little 1967! than lobsters reach minimum harvest size. In the Western Australian density-dependentfactors acting on juveniles Davis 1978 and fishery,juvenile habitat and food are other major limiting factors 1979!. This situation appears similar to that demonstrated for on growth that result in high juvenile mortality Chittleborough the peripheral areas of the Western Australian fishery 1970 and 1975!, and they may be amenableto amelioration by Chittleborough 1970!. In fisheries where definite stock- management action. Artificial reefs have been used successfully recruitment relationshipsare known Chittleborough and Phil- to increaselobster production in some areaswhere natural shel- lips 1975!, minimum size is set above the size of maturity. ter was limited Sheehy 1976!. However, as Chittleborough Where definition of this relationship is stilt lacking, it would seem 970! suggested,adding artificial shelters at the periphery of prudent to consider size of maturiiy in setting a minimum har- the Western Australia fishery would probably not be e8ective vest size. since postlarval recruitment was the most important population Regardlessof the stock-recruitment relationship, two other limit in such areas. Recent observations on the Florida fishery factors play a major role in selecting a minimum harvest size; support the view that in areasof low, highly variable postlarval they are growth and mortality rates. Both of these rates change recruitment, shelter in the juvenile habitat is not a major limiting throughout the stagesof the lobster life cycle, with growth gen- factor on the population Davis 1978 and 1979!. erally increasingin a sigmoid manner, and mortality decreasing logarithmically. Once these curves have been defined for a Critical Habitat population, selectinga minimum harvest size becomessimply a Juvenile spiny lobsters utilize nearshore habitats, in contrast matter of determining the point at which the biomass of the with the open water planktonic larvae and offshore reef- surviving lobsters is maximized. At maturity, growth rates gen- dwelling adults Little 1977!. Man's well-known destructive ac- erally decline as energy is redirected into reproductive activity tivities in the coastal zone frequently reduce the quality and and mortality rates are already relafively stable, so the size of quantity of this juvenile habitat, and by doing so reduce the maturity often approximates an optimal minimum harvest size potential yield of available posflarval recruits. As estuariesand for palinurid lobsters, Although perhaps the best approach, a seagrass beds are dredged,filled, or otherwisereduced by poor minimum harvest size and protected gravid females will not water quality o'r increasedsedimentation, their canying capacity ensure a successfulfishery, as witnessed by the California P. for juvenile shrimp and many other speciesis reduced or elimi- interruptus and New Zealand's Chatham Islands ed- nated Lindall and Saloman 1977!. uiardsii fisheries. It is clear that other approaches are also If the maximum yield is to be achieved for each postlarval needed, recruit reachingtheir shores,spiny lobster fishery managersmust Growth Rate look beyond the biology of their target speciesto their ecological requirements, including protection of sufficient areas of highly One of the major factors considered in setting an optimum productive juvenile nurseries.National parks that are managed size at first capture is growth rate. Management actions to in- on an ecosystembasis provide protection for about one-third of crease growth rates directlv can also be used to increase the the juvenile spiny lobster habitat 200 km'! in south Florida, yield per postlarvalrecruit. Sublegal-sized lobsters handled by With a moratorium on harvest to reduce injuries and the resul- fishermenare frequently damaged. Thesedamaging encounters tant mortality, not only is input into the fishery maximized from occur in several ways. In some areas, fishermen intentionally these areas, but naturally functioning ecosystems are made fish in nursery areas to catch sublegaljuveniles to use as attrac- available for comparisons with other similar but disturbed or tors bait! in traps for legal-sized lobsters elsewhere Florida manipulated systemsto evaluate various managementregimes, Stats. 370.14!. Recreational divers must catch an animal to and natural high concentrations of lobsters are available for determine if it is actually large enough to harvest, and since studying and public viewing. most divers fish in shallow nearshoreareas, they are frequently in nurseriesand must catch many lobstersto obtain a few "leg- SUMMARY als" Davis 1978!. The gregariousnature of juvenile spiny lobsters Chittleborough 1970; Bemll 1975! practically insures In discussingthese potential techniques for increasing or re- that many small lobstersare inadvertenfly damaged in the gen- storing yield per postlarval recruit, not all the possibilitieswere eral melee of pouncing and grabbing that ensueswhen amateur exhausted. In fact, not all of the possibilities have even been divers commenceIobstenng in nurseries.Finally, a few juveniles explored. The step-down diagram in Figure 1 outlinesthe steps

Fisheries, Voj. 5, No. 4 necessaryto pursue these topics further, Which management Bowen,B. K. 1971.Management of the western rock lobster Panulirus !ongipes cygnus GeorgeJ Pages 139 154in Proc.14th Sess Indo-Pacific Fishery Council, Bangkok. actions are selected and how well they achieve the desired ob- Chittleborough,R.G. 1970Studies on recruitment in the western Austrakan rock lobster jective will depend on the ecologicalconditions of the area and Panufirusiongipes cygnus George Density and mortality of juveniles Aust. J. Mar.Freshw the nature of the fishery. A juvenile habitat enhancement pro- Res21! 131 148. 1975Environmental tactors affecting growth and survival of iuvenilewestern rock grafn that would be effective in the central part of the Western lobsters,Panufirus longipes Milne-Edwardsl. Aust J Mar.Freshw. Res 26: 177 196. Australia fishery may not be effective when applied to the Chittleborough,R.G., andB. F. Phillips.1975. F!actuation of year-class strength and re Florida or California fisheries, where juvenile habitat is not a cruitmentin thewestern rock lobster, Panuiirus fongipes Mfine-Fdwards! Aust. J. Mar FreshwRes 26 major limiting factor, Recognition of the diversity of conditions Davis,G. E. 1975Minimum size ot maturespiny lobsters, Panvfirus angus, atDry Tortugas, that determine effective ways to maximize yield per recruit is an FloridaTrans Am. Fish Soc 104!:675 676 1978Prrxeedings. Sea Grant, Key West Florida, important first step in designing a functional fishery manage- 1979Management recommendations t'orluvenile spiny lobsters, Panukrus argus, in ment program. BiscayneNational Monument, Florida LI S Nat.Park Serv.. S. Fla. Res. Cent Manage. Rep There are three basic elements in the fnanagement of spiny M-350 Ingle,R. M., B, Eldred,H. W. Sims, and E. A. Eldred.1972. On the possible Canbbean lobster fisheries:adequate larval production, efficient and equit- originof Florida'sspiny lobster populations Ra BoardConserv. Tech Ser frio. 40. 12 pp able allocation of available stocks among fishermen; and Lewis.J. B. 1951.The phyllosoma larvae of thespiny lobster Panufirus argus. Bull Mar Sci. maximization of yield per postlarval recruit. This paper dis- GulfCaribb 1! 89 103. Lindall,W. N.. andC. H. Saloman.1977. Alteration and destrucrion ot estuaries atfecting cussedfour ways in which yield per recruit has been improved fisheiyresources ot the Gulf of MexicoMar Fish Rev. 39 9!:1 7 or maintained in spiny lobster fisheries: establishment of a Little,E. J., Jr. ! 977. Observations onrecmitment ofposriarval spiny lobsters, Pan uiirus angus. to thesouth Horida coast. Fla Nat. Resour Mar Res Publ. No. 29. 35 pp. minimum harvest size, maximization of growth rates, reduction Menzies.R. A.,and J. M.Rerrigan. 1980 The larval recruitment problem of thespiny lobster of mortality rates, and protection of critical habitat. Most meas- Frshenes5!.42 46 ures to maximize yield per recruit can be implemented with Phenicie,C.I .,and J. R. Lyons. 1973. Tactical planning infish and ualdlife management and research.L!S. Fish Wildl. Sen Resour.Pub!. No 123.19 pp. existing knowledge, but require additional inputs from recruit- Phillips,B. F. 1977.A reviewof thelarval ecology of rocklobsters. Pages 175 186iri B F ment potential and socioeconomic conditions to provide real Philbpsand J S.Cobb, eds Workshop onlobster and rock lobster ecology and physiology. optimum yield for American fisheries. CSIRODiv FishOceanogr Circ. No. 7 Sheehy,D. J. 1976.Lrri!ization of ariit'iaafshelters by theAmerican lobster Homaius americanuszJ Fah ResBoard Can 33!:1615 1622. LITERATURE CITED Sims, H. W.,and R. M. Ingle. 1967. Caribbean recruitment of Florida's spiny lobster popula- tion.Q. J. Fla.Amd. Sci 29! 207 242. Berrill,M. 1975Gregarious behavior of!uvenr!es of the spiny lobster, Panufi rue argus Crus- Zuboy,J.RA. C. Jones, and T. J. Costello.1980 Lobster hshery management under the tacea:Decapoda!. Bull. Mar. Sci. 25!:515 522 FisheryConservahon andManagement Act Fisheries5! 50- 52.

SPINY LOBSTER CULTURE: AN ALTERNATIVE TO NATURAL STOCK ASSESSMENT

George R. Tamm

ABSTRACT Routineculture of larvaehas not yet beenachieved, whereas it hasfor juvenilesand adults,A long and complexlarval life, inadequate knowledge of nutritional needs, and the maintenanceof high water quality standardsare major constraintsin larval culture,Growth in older animals,although naturally slow, can be enhancedby environmentalmanipulation. Several species appear well suited for advanced culture efforts including polycuiture. Coordinated programs of aquaculture and fisheriesman- agement appear feasible.

aising spiny lobsters in szJr,g adequately the feasibility of rearing lobsters either in intensive R captivity is one manage- or extensiveculture systems.This review will cover previous cul- ment strategy to consider to ture attempts, major constraints in raising palinurids, optimal cope with intensifying fishing ef- conditions for cultivation, and growth and mortality in captivity. fort and decreasing lobster Some thoughts on optimal species to raise and the future of stocks. Successful cultivation of spiny lobster culture are presented. palinurids might dampen cur- rent fishing pressures on lobster PREVIOUS CULTURE ATTEMPTS stocks by creating a competing source, thereby dropping prices Egg-bearingspiny lobsterswill releaseviable larvae in captiv- and lesseningthe monetary in- ity. Japanese researchersworking with P, japonicus were the centives for fishing for lobsters first to have any degree of successin raising phyllosoma larvae Those raised in captivity could in the laboratory Nonaka et al. 1958!, Inoue and Nonaka also be used to restock areas 963! are creditedwith rearingphyllosomas from hatching where populations once George R. Tamm throughthe VI instarin 40 48 days.Pore recently,Inoue flourished,but are now depletedor are in the decline.High 978! has succeededin culturingP. japonicusfor 253 days expectations notwithstanding, spiny lobster culture is in its in- from egg up to the last !IXIj phyllosornastage. Attempts to raise fancy and much basic research is still required to assess other palinurids have met with less success.P. argus and P.

July August 1980 guttatus have been raisedthrough 8 or 9 molts with some larvae most successfulin keeping the larvae alive. Inous 978! cul- living for about 70 days Robertson 1968!. Dexter 972! tured P. japonicus into its final phyllosoma stage, offering them reared phyliosoma larvae of P. interruptus for 114 days in a a diet of Artemta, Sagitta spp., and fish fry. Polychaete worms series of 8 molts, through their VI stage, Minor successesin were accepted by larval J. edLvardsti Bantham 1967!, Thomas culturing palinurid phvllosomas have been achieved for P, in- 963! has observed late-stagephyllosomas of attached flatus Johnson and Knight 1966!, P. polyphagus Ong 1967!, to scyphozoan medusae, and although feeding was not wit- Jasus lalandit Sheard in Robertson 1968!, J, edurardsii nessed, the pigmented material inside the transparent larvae Bantham 1967!, J. verreauxi McKoy 1979!, and P, ornatus matched that of the jellyfish in color. According to Williamson Michel 1979!. Thomas 1963!, Jasus will feed on hydromedusae. Provenzano Whereas culturing spiny lobster larvae is fraught with many 968! draws attention to the morphology of phyllosomas and difficulties, rearing postiarval and older animals has proven to suggeststhat the elongated legs and delicate mouth parts of be a relatively straight-forward task, given current aquaculture advanced stages are well stuited for clinging onto large, soft- technologies and a moderate amount of care. Lewis et al. textured prey such as jellyfish and sucking out the nutrients. 952!, Witham et al. 968!, and Sweat 968! have demon- Future successesin larval cultivation will hinge on advanced strated the feasibility of rearing juvenile P. argus under con- nutritional studies, trolled conditions for extended periods of months to years. Ser- Another consideration in rearing phyllosomas is maintaining fling and Ford 975a! reared groups of P, interruptus from exceptionally high water quality standards in the laboratory. pueruli to 14 months of age. Salinities,temperatures, oxygen concentrations,and pH should Chittleborough 974a! hand caught P. cygnus pueruli in the all match those naturally encountered in pelagic, oceanic wa- field and raised them in the laboratory for six years. Captive ters. Overcrowding of larvae should be minimized to prevent animals matured in 4.9 5.7 years, a rate comparable to the physicalentanglement of larvaeand possibleappendage loss developmental rate expected for wild populations. Under opti- Provenzano 1966; Robertson 1968!. mal environmental conditions, growth rates of captive lobsters The most acute problem in raising postlarvaeand older spiny exceeded that of wild stocks Chittleborough 1975, 1976!. lobsters is attaining adequate numbers of individuals to stock a Other palinurids that have been successfullymaintained in ex- culturesystem, The literatureis filled with remarkson the diff- perimenta! culture systemsfor months, or even years, include icult in trapping pueruli in the field see, for example, Serfiing Fielder 1964!, J. edwardsii Kensler 1967!, J. and Ford 1975b!. Phillips 972! has devised a lobster trap that verreauxi McKoy 1979!,P. homarus Berrv 1971!, andP. or- is fairly effectivein catchingP. cygnuspueruli, and the Witham natus Michel 1979!. habitat trap has on occasion proved quite successfulin collect- ing P. argus Sweat1968!. Many nursery reefs of P. cygnus MAJOR CONSTRAINTS IN CULTIVATION are overcrowded and trapping young juvenile lobsters age 2 + years!in largenumbers may be possible Chittleborough 1970! . The inability to culture spiny lobsters through their entire A final negative factor in rearing spiny lobsters is that growth phyllosoma larval phase has inhibited any large-scaleattempts is rehtively slow. It takes 3 to 4 years for P. argus to reach the to rear them experimentally or commercially. Several slipper minimum legal size Davis 1979!, 5 to 6 years for P. cygnus lobsters Scyllaridae!, with phyllosoma larvae morphologically Chittleborough 1974a!, and 7 to 10 years for J. verreauxt' similar to those of palinurids, have been cultured through all McKoy 1979!. Growth rates can be increased notably with their larval stages.However, Scyllarusamericanus required only superior diets and increasedwater temperatures. 40 days or less at 25 C to complete its larval development Robertson 1968!; Ibacus ciliatus and I. novemdentatus OPTIMAL CONDITIONS FOR metamorphosedinto posfiarvaein 76 and 64 days, respectively, CULTIVATION at comparable temperatures Takahashi and Saisho 1979!. Palinurids, on the other hand, may require more than 8 months Culturing phyllosomasis more of an art than a science.Sev- to develop in the laboratory Inoue 1978!. Chittleborough and eral environmental and nutritional requirements for rearing Thomas 969! estimatea 9 11 month phyllosoma phase for them have already been mentioned. For further details, see P. cygnus under ambient oceanic conditions. Estimates for Provenzano 966!, Robertson 968!, and Dexter 972!. other spiny lobsters i.e., 6 9 months for P. argus! are some- Juvenile P. cygnus have been held under an anay of con- what less Lyons and LitHe 1975!. Not only is the spiny lobster trolled environmentalregimes to determine those best suited for phyllosoma phase protracted, but it is comprised of a dozen or growth and survival. Lobsters two years of age or older should so instars, requiring at least that many molts Dexter 1972; be held in groups, provided ample shelter, and maintained at Inoue 1978!. If larval molting is as physiologically complex as 25 C ambient temperaturesof 15 26'C are encountered an- that reported for other life stagesof Crustacea Passano1960!, nually!, and the seawatershould be saturated with oxygen. P. and there is no reason to expect that it is not, then spiny lobster cygnuspreferred live food to stale or preparedrations and larvae are subject to a series of highly vulnerable periods at selected shellfish over fish, Daily feeding was necessary for which time feeding ceases,defenses are nil, and even small maximum growth Chittleborough 1974b, 1975!. Phillips et al. physical disturbancesmay lead to limb loss and possible death, 977! found that growth was enhanced in juvenile P. cygnus Inadequate diet is conceded to be a major obstacle in the when a mixed diet of lvfyttlusedulis!, abalone Haliotis rearing of spiny lobsters. Most researchersfeed larvae initiallv roeij, and pieces of teleost fish were provided. Because the with live brine shrimp Artemia! nauplii. Survival is usually poor speciesfeeds at night,food shouldbe offeredjust prior to dusk beyond Stage III Inoue and Nonaka 1963; Robertson 1968; or the dark phase to minimize the time for food decomposition Dexter 1972!. In feeding experimentsMitchell 971! and Dex- and decay. Uneaten food should be removed during the next ter 972! showed thatP. intenuptus phyllosomaswill also feed light phase. Kensler 967! has suggestedthat a detritivore be on Tubifex, Ivfytilus gonads, Lytechinus eggs, fish larvae, housed with the lobsters to act as a biological vacuum cleaner. ctenophores, and chaetognaths, The latter four items proved In all cases the trauma associated with handling should be

Fisheries, Vol. 5, No. 4 minimized,especially when animalsare in proecdysisor have Although few records are available on survivorship of spiny just molted. At these times risk of autotomy is high, and al- lobsters in the laboratory, the evidence suggests that under though lost appendages will be regenerated, growth is often near-optimal conditions, natural mortality is low. Phillips et al, slowed Chittleborough 1974b!. Similar holding conditions, 977! state that aquarium mortality over four years was about with slight modifications in temperature and food regimes, will one percent in P. cygnus, and Serfling and Ford 975a! report probably be effective in rearing many of the shallow-waterJasus that natural mortality of P. intenuptus was low in their culture andPanulirus species, although these hypotheses need testing. system. When culturing palinurids in redrculated seawater,additional concerns arise regarding water quality. Filtering systemsshould OPTIMAL SPECIES FOR CULTIVATION include substratefilters in which suspendedparticulates are re- moved,toxic metabolicwastes i.e., NH3! are oxidized,pH is Of the fifty or so extantpalinurids, few havebeen extensively buffered, and the water is oxygenated. Charcoal filtration re- studied,only a handfulhave been maintained in the laboratory ducesthe dissolvedorganic carbon load in the system.Algae for any length of time, and no specieshas been successfully addedto the filteringsystem tend to removeexcess NH3 NO3, reared through its entire life cycle. Therefore, nominating and CO,. Serflingand Ford 975a! haveemployed these fea- speciesbest suited for aquacultureis predictedon a greatdeal of tures in raising juvenile P. intenuptus. Details of filtration de- guesswork. signs can be found in Wheaton 977!. Most culturistswould agree that rearing phyllosoma larvae of any speciesis equallydifficult. For now, any seriousattempt to GROWTH AND MORTALITY IN raise spiny lobsters should begin with rearing juveniles. Some CAPTIVITY important criteria in selectinga speciesfor culture are that large numbers of juveniles be readily available to "seed" a culture From an aquaculturist'spoint of view, growth of spiny system;the speciesmust be adaptableto husbandryand en- lobsters held under natural conditions is rather slow. For in- vironmental manipulation, exhibiting fast growth and low mor- stance, Sweat 968! reared P. argus pueruli to juveniles, tality; it must be cultureden masseto minimizeoperational 50-mm carapace length CL! in size, in 28 months. Travis costs,and easilyprocessed, shipped, and marketed.Although 954! showed that P. argus, 50 90-mm CL in size, confine many of the palinurids examined to date meet most of these their growth to seven months May-November!, molt 3 to 5 criteria, few can be obtained in large number as juveniles on a timesper year,and increasean average9 to 12 mm in CL per routinebasis, Newman and Pollock974! haveobserved high annum.Thus, P. argusis expectedto take4 to 5 yearsto grow unspecifiednumbers! densities of J. lalandiion the Capewest from pueruli to a legally harvestablesize animal 6.3-mm CL! coast of South Africa. P. cygnus densities of about 2 x 10' underrearing conditions similar to thoseused by Travis954! juveniles per hectare have been recorded on certain shallow andSweat 968!. Wildstocks of P. argusare expected to reach nurseryreefs off WesternAustralia. Density-dependent mortal- legal size in 3 to 4 years in the Florida Keys Davis 1979!. ity is thoughtto be high and thesedense reef popualtionsmay Captive Jasus lalandii in the 50 90-mm-CL size class also be croppedwithout alteringrecruitment into the fishery growslowly, increasing in sizeaboui 3 mm CL! per molt, and Chittleborough and Phillips 1975!. A final consideration in moltingonly 2 to 3 timesper year Fielder1964!. Slow growth choosinga speciesto rearis the extentof the biologicalinforma- in individualsheld under "naturalconditions" is alsoexpected tion presentiy known on that species. inP. interruptus Lindberg1955!,P. homarus Berry1971!,P. In lightof thesecriteria, I suggestthat current spiny lobster cygnus Chittieborough 1974a!, J. ediJardsii, and J. uerreauxi culturefocus on raisingP. cygnus,possibly P. ornatus,and a McKoy 1979!. Contrastingly, Michel 979! claims to have large-growing,colder-water inhabitant such asJ. uerreauxi or J. rearedfive posflarvajP. ornatusto 300 g in weightwithin 10 edwardsii.P. cygnusin one of the few palinuridsavailable in months,These animals also began to mateand spawnin their large quantities as juvenfies; a fair amount is known about its second year of captivity. ecologicaland physiological requirements; ii appears to adapt Throughmanipulation of environmentalconditions, growth well to captivity,and its growthrate is easilyaccelerated when ratesof P. cygnus Chittleborough1974b, 1975; Phillips et al. temperatures and food availability are increased.P. ornatus is 1977! and P. interruptus Serfling and Ford 1975b! have been apparently a very hardy and fast-growingspecies Michel markedly enhanced. When P. interruptus were cultured at a 1979!.Its widespread distribution in theIndo-Pacific is a plus; constant 22'C, they reacheda mid-juvenile size 5-mm CL! in however, the availability of its juveniles is unknown. There is one-halfthe tiine expectedfor wild populations.Serfling and some publishedliterature on the distribution Chitfleborough Ford 975b! estimated thai three lobsters would reach their 1974c!, ecology Debruin 1969!, and fishery of P. ornatus minimumlegal size 82-mm CL! in 3 vearsas opposedto 7 Moore and MacFarlane 1975!, but other information is scant. yearsat ambienttemperatures in Californianwaters, Raising the TheJasus spp. are suggested because their natural growth is temperatureto 28 C inducedeven faster growth, but also an most likely minimalin winter, when water temperaturesmay appreciableincrease in food consumption.P. cygnushas been dropto 10 15 'C. Byrearing them in water20 to 25'C year raised from two years of age 0 40-mm CL! to minimum round,they probablywill molt throughoutthe yearand, if legal size 6-mm CL! in 57 to 80 weeks when held under amplefood is provided,they shouldrapidly increasein body near-optimalconditions in the laboratory Chittleborough weight. The cultured animals may reach a marketable size 1974b!,as opposedto the 150 to 200 weeksrequired for two- within a year to two. year-oldsto reachmaturity on nurseryreefs Chittleborough 1974a!. These observationsdemonstrate that growth in warmwater-dwellingpalinurids, as in coldwater-inhabiting FUTURE PROSPECTS nephropidlobsters Hughes,Sullivan, and Shleser1972!, can Raisingspiny lobsters commercially in highlymechanized or be improvedgreatly by optimizing the environmental regime of "intensive" culture systems,dependent on a continualsource of the culture system. conventionalenergy for the operationof pumps,filters, feeding

July August 1980 61