Grouper Stocks of the Western Central Atlantic: The Need for Management and Management Needs

YVONNE SADOW Fisheries Research Laboratory, Department of Naturaf Resources, P. 0.Box3665. Mayagiiez. Puetto Rico a0681

Grouper stocks fhrouglrout much of thz western cenlnl Athtic rue being heavily fshed. An overview of the infon3tion on stocks in the region indicates that nianv are erowth and/or recruiunent overfished. and that m&arrement is essential io ins&. their continued commercial and &rational viab&ty. Dam necessary for effective management of prouDer are identified. md the need for stock monitoring and asseshent to bg stihdardized and co-ordinated. on a regionwide basis, is emphasized. KEYWORDS: grouper; fisheries management

INTRODUCTION The purpose of this paper is to focus the nuention of biotogists, ftsheries managers, and public officials on the precarious smte of gouper stocks in much of the western cenlnl Atlantic. In particular, I want to emphasize the need for smdardizing and improving data collection protocols in the region, and also to identify the major daw. gaps precludimg development of more effective management approaches. I have drawn together information widely dispersed in the scientific titeratwe, unpublished repans, manuscripts, newsteners, and management plans to produce an overview of (1) the current status of stocks in the region, (2) the most pressing data needs for managing and monitoring stocks, and (3) the current approaches to mmayemeni Whie I have focused on grouper, many of the problems of resource status and use, monitoring, and managment are relevant to dcmersal (subsmte-associated) fisheries resources, in general, throughout the region. Grouper are of considerable commercial and recreational significance throughout the Caribbean. Bermuda, the Gulf of Mexico, Florida, and southeastern United Smtes. They are also species of major ecological imponance and of Beat aesthetic and scientif~value. A total of 20 species is exploited, mging in length from the giant jewfish (Epinephelus itajara) which may exceed 2 m to the creole fsh, or barber (Paranthiusfurcifer), which barely reaches 0.3 m. Wable 1). Among the most economically valuabIe species are the red grouper (E. morio), followed by the gag (Mycteroperca microlepis) in nonhem continental areas. and the Nassau grouper (E. striatus), at least historically, and red hind (E. guttatus). among the islands of the region (Fischcr, 1978: Mahon. 1987). Proceedings of the 43rd Gulf and Caribbean Fisheries Institute Peer Reviewed Section

Table 1. Grouper species of commercial and/or recreational importance in the STATUS OF GROUPER STOCKS Western Central Atlantic. Information on the condition of grouper stocks (a stock is the part of a fish population which is under consideration for actual orpotential We) in the region Epinephelus adscansionis rock hind is patchy and largely incomplete. Few databases were initiated prior to the wly Epinephelus cruentatus graysby ^' 1970's. and even now much of what exists derives from a handful of the island Epinephelus dmmmondhayi speckled hind nations of the region, and from southeastern United Sta~esand the eastern Gulf Epinephelus flavolimbalus yellowedge grouper of Mexico. Epinephelus fulvus coney" The condition of a fishery may be monitored in a number of ways. Data on Epinephelus guftafus red hind '^ catch per some unit of effort, such as per fisherman day, nip, or gear hour, may Epinephelus itajara jewfish be used to follow trends in catch over time in a standardized manner. Changes in Epinephelus morio red grouper" catch composition (i.e., relative numbers of different species cnmpriring the Epinephelus rnysracinus misty grouper catch), in the mean size of captured individuals, or in catches from spawning Epinephelus ngnrus Warsaw grouper aggregations, also provide indications of the response of stocks to fshing Epinephelus niveafus snowy grouper pressure (Mum, 1983). Fimally, formal stock assessments. although Epinephelus strialus Nassau grouper ' time-consuming and expensive to do, permit more rigorous assessments of Myctemperca bonaci black grouper " actual and potential yield under u range of fshery conditions. Myctemperca inferstitialis yellowmouth gmuper I have divided available information into five categories (Table 2): a. Myctempem rnicmlepis gag " iandigs or catch per unit effon (CPUE); b. catch composition; c. mean size Mvctemoerca rrhenax scam0 (length or weight); d. chancteristics of spawning aggregations; and e. formal Myctemperca tgris tiger grouper " stock assessments (yield-per- remit and spawning-stock-biomass per-recruit). Mycteroperca venenosa yellowfin grouper Alphestes afer mutton hamlet a. CPUE and Landings Paranthias furcifer creole fish Long-term landings and CPUE data are available hom few locations. Moreover, since data on grouper species were originally combined, assessment and "-evidence exists for female to male sex change hv-, individual soecies in the wly years of exploitation of a number of gouper '-Carter eta/., (1994) stocks was no; possible. This si(uation has improved and gouper arc more " - Shapiro. (1987) Erequently monitored on a species by species basis. Dam from Bermuda show an almost fourfold decline between 1975 and 1989 for reported landings of larger grouper species and a steady decline in Grouper spies share a number of life-history chancteristics believed to CPUE (Ibs. per trap haul) beween 1975 and 1985. with a slight rise there&& render them particularly vulnerable to human exploimtion (Manmh, 1987; (Repon of the Commission of Inquiry. Bermuda, 1991). Particularly striking Raiston, 1987). They are cmivores, have a relatively long life-span. large size have been the declines in reported weight landed of certain species, principally at sexual maturation, slow growth, and appear to be relatively easy to catch. the yellowfin (M. venenosa), Nassau and tiger groupers (1M. tigrisf, as much as being susceptible to a wide mge of sizes and types of fshing gear. In particular, 15-fold over the &year period between 1975 and 1981 (Bannerot el al., 1987. two chancteristics may make them especially vulnenbie. Many exhibit a sexual Fig. 13.9). pattern incorporating adult sex change (female to male sex change, known as In Puem Rico, a six-fold decline (in weight) in reported grouper landings pmtogyny) making them more sensitive to fshing pressure than gonochore (non (all species combined) was reported between 1977 and 1989 following a steady sex-changing) species (Banneror 1984: Bannemt ef al., 1987), under certain increase as the fsheries of che island were developed during the mid-1970's conditions. Aixt, some species spawn in large numbers at weU-dehed times (Sadovy and Figuemla, 1992, Fig. 1; unpubl. &, Fisheries Research and locations each yw. Many of these spawning "aggregations" are thus easily Labontory (FRL), CODREMAR. now the Depment of Natural Resources - located in both time and space and have ken heavily %bed. (DNR). Mayaguez, Pueno Rico). Since the number of fshermen in Puem Rico is not believed to have declined markedly during this 12-year period, the picture Proceedings of the 43rd Gulf and Caribbean Fisheries institute Peer Reviewed Section IJ of consistently dimiinishmg landings likely also reflects a decrease in CPUE. Pyticularly alarming have been the declines in landings of Nassau grouper in Puerto Rico and the U. S. Virgin Islands. In Pucrto Rico, this species was once one of those most Frequently landed (Sujrez-Caabm. 1970). "A common and very important food-fish, reaching a weight of 50 pounds or more" (Evmann, 1900). Now it is essentially extinct for fisheries putposes (Bohnsack et al., 1986). In Martinique, as in much of the Lesser Antilles, grouper comprise less than 10% of the demersal arch with three smaller species dominating: the coney, E. fulvus, the red hind, and the muuon hamlet, Alphesres afer. I Species-specific trends in CPUE by depth across the island platform indicate that grouper biomass is severely reduced on the shelf (Goben 1994). In 1 Jamaica, MUNO(1983) reported that the mean catch rixe of red hind in exploited areas was significantly less than that of unexploited areas. In gene&, grouper are found to conuibute relatively less to landings in arm that are heavily exploited than in those which are lightly exploited (Appeldwrn er al.. 1987). For a number of species taken off eastern and southern Florida and the states of Nonh and South Cmii. United Slates. catch by weight and by number, as well as CPUE, declined during the 1970's and 1980's. The most affected species have been the Nassau, black;snowy, and warsaw (E.nigritus) groupers, and the speckled hind. as well as the red grouper in southern Florida. Data indicate that scamp, M. phenar, and gag. speckled hind, E. drwnmondlmyi. and snowy grouper, E. nivearus. can quickly become depleted by heavy unregulated fishing activity (J3unlrman and Diion, 1975: Meare and Labiisky. 1984: Bohnsack, unpubl. ms.; Plan Development Team - National marine^, Fisheries Senice (NMFS) Beaufon North Carolina - Report. South Atlantic Reef Fish Snapper-Grouper Assessment (SARSGA). October. 1990). The most seriously impacted species appears to be the jewfish, which in southeastern United Staw is heavily overfrshed. Evidence of overfishing in both recreational and commercial sectors include comments from divers and dive boat opentors who report declining numbers (Amendment 2 February. 1990, Fisheries Managment Plan for Reef Fish of the Gulf of Mexico Fisheries Management Council (FMP-GMFMO, 1981). Thcse data consistently indicate that rapid, heavy declines have occurred in landings and CPUE for many, if not most. grouper species following unregulated exploitation of suxks, assuming that landings uends reflect populauon levels.

b. Catch Composition Gmupcr exhibit another classic fiheries response to heavy fshing pressure: a change in catch composition from larger to smaller individuals and/or species over time. Catch composition dam indicate that over the last 13 years the proportion of the towl repaned landings (in pounds) from the commercial Proceedings of the 43rd Gulf and Caribbean flsheries lnstltute Peer Reviewed Sectlon ftshery of Pueno Rico (all reef fsh included) comprising gouper species has commercial fisheries of gag. scamp. warsaw, and snowy groupers and in the dropped from 16% to 7% (1977-1989) (unpubl. &fa FRL.-CODREMAR-Dm, speckled hind. Data on gag from the Carolina headboat fishery recorded that, Maws and Sadovy, 1990). A similar decline (48% to 19%) over a 14-year period while in 1973 virtually no fish were less than 800 mm in total length, by 1988, (1975-1989) was reported from Bermuda. 97% of fsh taken were less than this in size (SARSGA. 1990). Grouper landings in Bermuda have exhibited a marked differential loss (by weight) of Same of the larger species, such as the Nassau gouper (16% -

(1971) repaned that the smali size of the red hirendered it of minor 8 have been reported for the Nassau gonper. the red hind, and the yeuowfin commercial imponance in Bermuda prior to the 1970's. but that its abundance grouper (Smith. 1972; Bumeu-Herkes, 1975: Olsen and LaPlace, 1979: Colin er made it useful as bait for larger species; it is many yeam since the red hind was al., 1987; Carter, 1988: Beets and Friedlander, 1992; Colin, in press). One has taken solely for bait in Bermuda Grouper species that were once common in the recently been reponed for the tiger grouper in eastern Puerto Rico, and markets of Jamaica are now rare Wlton Haughton. Jamaica, pa.comm.). In aggregating behavior appears likely for the gag and scamp (Gilmore.1994), as southeastern United States shifts in catch composition to less desirable species well as for the black grouper (Carter el al., 1994). Reports from heavily fished in both commercial and recreational fisheries, especially loss of the largest aggregations indicate declining catches. and, in some cases. declining mean predators, have been noted (SARSGA, 1990). Since gouper species are among sizes and progressively female-biased sex ratios over time (Carter e! al.. 1994; thc largest of Ihe multispecies fishcry components, it is not surprising that they Beets and Friedlander. 1992; Sadovy. Rosaiio and Roman unpubl. data; Grant are among the first to be affected by fishing activity. Gilmore, pers. comm.; Jack Ward. pers. comm.). Whether these changes result Overall, dafa indicate declines in both absolute and relative terms of directly horn fshing pressure on the aggregations, or simply reflect pressure on exploited grouper species, with a shift in the proportion of gouper landed from swks in the yea throughout the year, or perhaps some combination, is nor predominantly larger to predominantly smaller individuals and species over known. However. Carter et at. , (1994) showed that an exploited aggregation in time. Belize yielded smaller individuals than either an unexploited aggregation or an I exploited population during the non-reproductive period. r Trends in Size Particularly hard hit have been the Nassau grouper aggregations. Several, Fishing activity tends to select for largerheavier individuals of a species. with a history of heavy Ctshing in Puerto Rico and St. Thomas (Olseu and For this reason changes in mean fish sue over time can provide indications of LaPlace, 1979: Carlos Cumpiano, CODREMAR, pers. comm.; Beets and species' responses to exploitation. Friedlander. 1992). have now completely disappd, at least from their Analysis of size-frequency &la of the red hind from St. Thomas indicates a original locations. It is not known whether disturbed aggregations develop gradual, but consistent, decline in mean size between 1984 and 1988 with a elsewhere de novo, although this has never been reponed. Catches from complete loss of the largest size classes over this period (Bohnsack el al.. 1986; long-fished spawning aggregations in the Cayman Islands and in Belize hwe Beets and Friedlander, 1992). A similar progressive decline in size was recorded exhibited panicularly marked declines during the 1980's (Carter el a1.,1994; for the coney in St. Croix between 1984 and 1989 (Bees and Friedlander. 1994). Bush and Ebanks-Peuie, 1994). Reduced catches from grouper aggregations in In Jamaica, differential loss of larger red hind and Nassau gmuper were noted the Dominican Republic led, recently. to prowtion of spawning Senanidae on hm more heavily exploited arcas (Munro, 1983). Over the last decade, the nonh coast, and prohibition of fishing at spawning sites (Colin. 1988). commerciill samples of the once abundant Nassau grouper from Puerto Rim In Mexico, an aggregation at Mahahual was fished by hook and line since have been scarce and have consistently fallen almost exclusively within the the 1950's but numbers apparently declined sharply following introduction, in immature size range (i.e., c 450 mm standard length) (unpubl. &ta the late 19M)'s. of spwyns (Aguilar-Percn and Sosa-Cordem. 1994). The use FRL-CODREMAR-DNR). of spearguns on aggregating grouper appears to be a very efficient technique. Analysis of weight dmfrom the 1970's and into the 1980's from Florida For example, many thousands of pounds of tiger gouper were reported &en by and the Camlinas show a progressive decline in mean weight for most spccies in a couple of divers on just one day of an aggregation lying in about 120 it of both headboat (boats that charge on a per person. or per 'head', basis) and Prodeedings of the 43rd Gulf and Caribbean Fisheries Institute Peer Reviewed Section water off Puem Rico (ishmael Rivas, pers. comm.); tiger grouper may be remained high andstocks have rapidly declined. For gag, speckled hind, and particularly susceptible to spar fshing because of their curiosity toward divers snowy grouper, for example, mean size captured in 1985 equalled the minimum (Appeldoom et a!., 1987). size considered necessary to produce a reasonable yield-per-recruit (Huntsman and Wiis, 1989; SARSGA, 1990). e. Stock krsessment Recent assessments of red grouper and jewfish fmm the Gulf of Mexico Assessment of stock status at current levels of fishing mortality, and age, or indicate that spawning stock biomass per remit for both species may be below size, at which individuals of a species enter (recruit into) the fshery, have been the theoretical minimum necessary for maintaining stocks (Goodyear 1988: predominantly made using yield-per-remit WIR) (Bevaton and Holt, 1957). Amendments 1 and 2,1990. FMP-GMFMC. 1981). Similar stock assessmenwof and spawning-stock-biomass-per-wruit (SSBIR) (Goodyear. 1989) models. speckled hind, gag, scamp. snowy, black. red and Warsaw groupen from the ab These models enable the fishery manager to establish what would happen to south AtLantic indicate that same stocks, especially Warsaw gouper, are stock yield (i.e., catch), or spawning stock biomass (i.e. .reproductive potential recruitment overfished (SARSGA, 1990: Huntsman et al., 1994). of a stock), at different levels of fshing mortality (death in fish stock caused by With few exceptions, therefore, stock assessments indicate that both growth fishing), or at different sizes of recruitment into the fishery. and/or wruitment overfishing is occuning for many species at a number of The models allow fsheries managers to establish the recruitment age (and locations. This is, in pan, because age (or size) of recruiunent into the fishery hence size) and level of fishing mortality likely to produce Ihe thearetical has not been kept sufficiently high to manage yield approprintely under the maximum sustainable yield (the largest average catch. or yield, that can conditions of the fishery or insufficient adults remain to replenish stocks. continuously be taken from a stock under existing environmental conditions) of Management mwures need to be effected, improved or enforced. depending on the stock. If fishing mortality is grater than this theoretical level, or fsh enter location, to regulate fishing mortality, or size of recruitment into the fishery. the fishery at too small a size, then yield will decline and overfi~shingwill occur. Overfiihing may take one of two basic forms. Growth overfishing occurs DATA NEEDS when fish are caught before they have had an adequate chance lo grow. More Stock monitoring and effective management are impossible without severe is recruitment overtishing, when fishing reduces adult numbers to a level knowledge of the biology, smlus, and exploitation levels of the species to be which greatly decreases egg production and hence increases the chance for managed, as well as the socio-economics of resource usen. Significant gaps in recruitment failure (i.e.. not enough juveniles will be produced to replace the our knowledge of grouper biology, stock characteristics, and exploitation levels adult population (Plan Development Team (PDT), 1990)). undermine stock monitoring and management More attention must be given to In the Caribbean, a recent yield-per-recruit analysis of red hind from the following five mas: western Puem Rico and SL Thomas showed that. in both cam, slocks are growth overfished (Sadovy and Fiyemla, 1992). By the late 1970's. Nassau a. Landings, CPUE, Catch Composition and Size Frequencies of Exploiled grouper were found to be likely overGshed in the U. S. Virgin Islands, and Stocks The recording of landings, fishing effort (the total fishing gear in use for underexploited off northwestem Cuba (Baisre and Piez, 1981). a specified period of time) catch composition, and individual size and weight Fmm the wtem Gulf of Mexico. Moe (1969) reported declining stocks of data, on a long-term basis. is necessary for any fishery monitoring pmgnm. red grouper between 1965 and 1968 following 15 years of flucovting or While these data are essentially simple to collect, care is needed to ensure that increasing catch, and expressed concern that management would be necessary if they are adequately and consistcnlly recorded, and are appropriate for stock yield continued to decline despite increasing fishing effort (see also analyses by monitoring and management For example, effort and size frequency data should Blanco et of., 1980; Contreras et at., 1994 and Burton, 1994 on red grouper). specify gear type, including mesh size for traps or nets, because of the influence For gag. scamp, snowy, and black groupers, and the speckled hind, in of gear on the data collected (see Munm, 1983). Units of fishing effort should be southeastern United Swles, stock assessments conducted in the early 1980's specified (e.g., catch per trap haul, per hook how, per fshing tlip, etc), as showed that if the age (and hence size) at which individuals fmt enter the should the sector of the fishery being monitored i.e.. commercinl or recreational. fishery (recruitment age) could be kept moderarely elevated. then When catch composition is recorded, any dam biases should be noted, such as yield-per-recruit would stay high regardless of how great fshing monality the possible undersampling of small individuals which may have been discarded should become (Huntsman er al.. 1983: Matheson and Huntsman. 1984; prior to landing. When subsamples are taken from complete catches, it should be Manooch and Mason, 1987). However, age (and size) of recruitment has not Peer Reviewed Section Proceedings of the 43rd Gulf and Caribbean Fisheries Institute determined whether large. small, or rare species tend to be preferentially 240 km between spawning areas in Belize waters (Caner er at, 1994). An measured (Bohnsack el al., 1986), and whether subsamples are t;lken randomly. individual gag was found to have moved 621 km, during a possible spawning Furthermore, data collection should aim to sample all principal gws used $for migration, off sonthwtern United States (van Sant er al.. 1994). We need to grouper these would be hook and line, fish trap, and spearyn), ideally for all know the geographic locations and ddons of significant aggreghons. For seasons, and in all principal fuhing areas: modifications in gear use not only example, despite the presence of exploitable stocks of red hind in the islands of influence the nature of the data collected, they also reflect user response to the Lesser Antilles. no aggregations arc known for this species in much of the changes in the condition of a fishery. eastcm Caribbean. Duntion of spawning activity cach year may be extremely Data collection should incorporate lengths and weights (units of length and brief. In the red hind, for example. all annual spawning activity at two spawning weight - whole or gutted - and precision of measurement should be specified), locations in Puem Rico, in different years, was recorded to ncur over no more and length-weight and length-length (i.e., smdard length to fork lengtNtot;.l chan two weeks (Sadovy eta/., in press: Sadovy. Rosario and Roman. unpubl. length) relationships should be established. This enables comparison between data). Given the apparent impomnce of spawning aggregations, we need to data sets which have been collected in difierent ways. Data on size distribution know how aggregation fishing is llkely to affect counship or spawning behavior by sex should also be collected whenever possible, although for grouper this (see also discussion in symposium summary). Data recorded from aggegation may be difficult using gross inspection of gonads outside the spawning season. sites need to be specific regarding date, location. and fishing gear, and should be However, if examined carefully, ripe individuals taken at spawning aggregations collected on a long-term basis. may be sexed. Relatively little is known for many species concerning the minimum size of Collection of data on lengths, weights, and sex of ripe individuals does not sexual maturation relative to size of enuy into thc fishery. This information is require sophisticated equipment or extensive mining of personnel. Nonetheless. necessary to determine the proportion of the catch which is composed of if smdanlized and well documented, such information provides an essential juveniles. If this proportion is too high, then management measures must aim to database which may be used for comparative purposes bath over time and reduce it. Information is also needed on the relationship between fecundity and between geographic locations. Because many stocks are likely shared by many body size, the male to female sex ratio, and on the size of sex change, if this of the nations of the western cenml Atlantic (Mahon, 1987). standardized and occurs. Sex change has been confiied for less than half of the groupers of compatible data collection programs over as wide a geographic area as possible TabIe 1, and certainly not all gouper are exclusively hemphroditic (Sadovy would greatly assist our understanding of slacks on a regionwide basis. and Colin, 1990). Mode of reproduction (i.e.. hermaphroditism versus gonochorism) (Sadovy and Shapim. 1987) must be established to determine the as b. Reproduction possible impact on stocks of fishing activity (Bannemt ef a[., 1987). well as A better undersonding of grouper reproductive biology would greatly their response to management measures. For example. fishing activity can result faciliate stock management In particular, among those species that assemble to in a reduction of mean individual size, thus reducing the number of males, since males are generally larger than females in protogynous species. If male reproduce, we need to know whether most annual reproduction occurs exclusively at the spawning aggregations. If so, and there is no evidence to the availability is a limiting factor for successful reproduction, and if sex change is conhary, then the limited number of known aggregation sizcs that have been convclled by age (or size) rather than by some socially-meditcd factor, then a identified (Colin et at., 1987; Beets and Friedlander, 1992; Sadovy et a!.. in heavily fished stock may not be able to compensate for the differential removal press) suggest that these aggregations are of critical importance to stocks in of the larger males. If, on the other hand, sex change is behaviorally induced. much of the region. Moreover, since individuals of a number of species are then a decline in males may be compensated for by increased numbers of known, from mark-recapture studies, to be able to move over considerable females changing sex. However, this could result in a decline in mcan female distances, it is not unlikely that each aggregation site services individuals fmm size leading to a reduction in stock egg production. Until more is known of extensive areas. For example, red hind may move at least 18 km. through water reproductive dynamics, the response of grouper populations to exploitation and of at last 194 m to reach an offshore bank in westcm Pueno Rico (Sadovy, management cannot be fully evaluated. Figueroln and Roman, submitted ms.). individual Nassau grouper have been reponed to mvel as far as 110 km (Colin, in press) to a spawning site in the c. Recruitment Bahamas, and an individual Nassau grouper was recorded to move a distance of It is not known to what extent grouper recruit (in this context "recruitment" refers to the seulement of planktonic larvae onto the substrate) locally or from Proceedings of the 43rd Gulf and Caribbean Fisheries Institute Peer Reviewed Sectlon

lanrae originating from upstream andlor off-island locations. This information longevity, mortality, and other panmetee used in formal stock assessment. is essential for establishing the necessary geographic extent of monitoring and Unfonunately. these data are difficult and/or extremely expensive to collect, management initiatives. The fact that some grouper spawn in large especially on a regular basis, and, for many small. understaffed, and concentrdtions and are known to be able to migrate over substantial distances to underfunded fisheries divisions. may he impossible to obrain. reach aggregation sites, indicates that adults move acms international For these reasons, small fisheries divisions may have to rely, at least partly, boundaries where platforms are shared by more than one nation. This latter has on population parameters established elsewhere and corrected for local obvious management implications. conditions, rely on relatively simple methods for stock assessment (Pauly. Assuming a larval life of between 3 weeks and 2 months (Colin er 01.. 1987) 1980). or use appropriate length data in length-frequency analyses (the use of for grouper species, spawning aggregations in one location may well supply data on fish lengths to determine growth panmeters) such as WAN(Pauly, recruits well downstrwn, on different geological platforms, depending on 1987) to produce the necessary information (Caddy. 1986; Beets et 01.. 1994). prevailing currenu. Colin el d. (1987) For example, proposed that, given the However, for the larger, slow-growing, long-lived species of grouper, heuse of current patterns off southwestem Pueno Rico and in the westem Caribbean, length-frequency techniques may be limited because of considerable size larvae from spawning areas in this location could potentially reach at least to variation at different ages. Finally, it needs to he established whether standard Hispaniola, Jamaica, Cuba, Turks and Caicos Islands. and the southeastern stock assessment models, originally developed for temperate, gonochore, Bahamas. However, on the Pueno Rico and Viin Island geological platform species, can be effectively applied to tropical. hermaphroditic species. For area anecdotal information suggests the likelihood that local recruitment is of example, adult sex ratios are assumed to be 1:l for spawning st& biomass per some importance: there appears to be an association between local declines in recruit analysis. Xis ratio may not apply to species of grouper that rnnsform catches of the Nassau grouper over this platform and loss of local spawning from female to male during their lifetime because of the frequent female bias to aggregations; the losses of both being most marked for Pueno Rico to the west, the sex ratio (Goodyear, 1989). Yield-per-recmit models deal with sexes-lumped and least for the British Virgin Islands to the east (pec;. obs.). It has been data; the impact of sex-specific differences in age and size dishibutions, suggested for gag off southeasrem United States, that a single stock may be associated with sex change, on the estimation of the population parametee used involved, indicating the need for a uniform monitoring and management strategy in these models, has yet to be evaluated (Sadovy and Figuemla, 1992). in that area (van Sant er al., 1994). Powles (1977) hypothesized that grouper might he recruited to southeastern United States from the Gulf of Mexico. It is e. Sociu-economic Factors thus imponant for stock management to establish the extent to which local A knowledge of local social and economic factors associated with the stocks are self-recruiting, and to what extent they depend on larvae generated at fishery would assist assessment of the possible impact of management options aggregation sires upstream. Collection of landings and other data in a on the different resource user groups. It would also facilitate the improvement of standardized and co-ordinated manner throughout the region, combined with dam collection prognm design. genetic studies to distinguish different populations, would be needed to address some of these imponant issues (Appeldoorn el al., 1987). MANAGEMWT AND CURRENT MANAGEh4ENT MEASURES Little is known of the chmteristics of critical juvenile habitat for the By November 1990, four classes of management measures were in effect majority of grouper species, nor of their principal settlement periods. Juvenile specifically to manage gouper stocks in the region: minimum size (weight or Nassau grouper are reponcd from gnss beds, jewfish from mangrove areas, red length), prowtion of spawning aggregations, quow/bag limits, total harvest ban hind from shallow mky, rubble and sandy areas @en. obs.), and gag from Ojble 3). These measures are summarized below. Not included are broader estuarine areas (MuIlaney.1994.). Hence, juveniles of several species depend on measures, such as marine resenres or prohibition of spetyuns or mps, which shallow. neatshore habirats which are those most susceptible to coastal human would also impact these smks, nor included are a number of harsher measures activity. Nonetheless. for many species. little or nothing is known of juvenile presently under consideration in Amendment 4 of the South Atlantic Fisheries habitats. If critical juvenile habitat is being impacted then management of adults Managemcnt Council (SAFMC News Release. Nov. 7,1990). may be of little consequence For the condition of local stocks. a. Minimum Size of Capture d. Stock Assessment The most widespread management measure currently utilized is minimum Biological data on exploited stocks are necessary to determine growth, capture size (for grouper this ranges from 12" and 20" TL.. depending on species Proceedings of the 43rd Gulf and Caribbean Fisheries Institute I Peer Reviewed Section and jurisdiction; minimum weight of 3 lbs. for the yellowfin and Nassau groupers in the Bahamas) (Amendments 1 and 2, 1990, FMP-GMFMC, 1981; Fishery Management Plan for the Snapper-Grouper Fcshery of the South Ahtic Region. 1983 and its Amendment 2, 1990: Florida Smc Law; Luckhurst, 1990). The effectiveness of minimum size restrictions depends on two critical factors; adherence to the law and survivorship of released, undersized, individuals (Waters and Huntsman, 1986). the lauer a variable yet to be fully evaluated; survivorship is generally believed to be low for released gmuper species, in panicular for those taken from deep water which exhibit pronounced stomach eversion on surfacing. Imperfect survival of released fish reduces thc expecwt increase in yield-per-recruit resulting from size limit regulations. Moreover, the effectiveness of minimum sue limits for protogynous (female to male sex changing) species under conditions of high ftshing mortality are not known (Huntsman and Waters, 1987). Recent evaluation of the impact of minimum size regulations implemented in the early 1980's. in Florida, indicated that there was little obvious incrrase in sizes recorded from subsequent hdings of Nassau and black grouper (Bohnsack, unpubl. ms.). Whether this is due to lack of compliance to, or enforcement of, the law, or the inappropriateness of minimum size limits for grouper management is not known. h. Protection of Spawning Aggregations Swsonal andtor zeal closurcs are designed to protect stocks which may be panicularly vulnerable to fishing pressure at specific times andlor locations, such as during spawning periods. Such closures also reduce fishing mortality on the spdwning stock Aggregations in Bermuda and in the Dominican Republic are protected from all fishing activity. In the Cayman islands, aggregzuion fishing is permiucd but access is restrictod to local residents and hook and line fishing. In St. Thomas, U. S. Virgin Islands, no fishing at all is permitted over the three month period identified as the red hind spawning season (Dec. 1 - Feb. 28) at a location lying in fedenl waters south of the island. A similar measure has been proposed for a red hind spawning area in federal waters off western Puem Rico (Sadovy, Rosario and Roman, unpubl. dam). There is much support, from both theoretical and pmtical standpoints, for the protection of aggegations (Bohnsack, 1989: Beets and Friedlander, 1992; Sadovy and Fiyerola, 1992). However, the extent to which protection of aggregations, in the absence of additional management measures. is likely to protect the spawning stock cannot yet be fully evaluated.

c. Catch Quotas Esrablishment of catch quoras requires sound knowledgc of fishing effort and catch and of morlality rates of released individuals. This information is Proceedings of the 43rd Gulf and Caribbean Fisheries Institute Peer Reviewed Section I often difficult to obLlin. and limits hence difficult to establish, monitor, and Nonetheless, many island nations of the Caribbean ye extremely limited in enforce. Bag limits have been implemented in Bermuda and in some U. S. their capacity to evaluatc and resea~~htheir own resources, having to deal with mainland waters for a number of recreational and commercial fihery species. considerable shortages of funding and personnel, combimed with the intrinsic The effectiveness of bag limits for the management of gouper is not known. . difficulties of monitoring nrtisanal fisheries. Moreover. the low natural productivity of the idand platform amand the relatively limited economic d. Total Harvest Ban value of individual components of multispecies fiheries. result in low priority This measurc aims to protect individual species believed to be severely for tisherics funding, little or no law enforcement, and little interest from local overfished. A total ban on harvest has been implemented for only one species, legislators (Kimmel and Appeldwrn, in press). Many nations do not even have a the jewfish, in U. S. federal waters of the'~u1fand South Atlantic, and in Rorida national policy regarding their fishery stocks, which frustrates aaempts to assign state waters. This measure has also been proposed for a number of other species priorities to resource use and management (speckled hind. warsaw, Nassau, snowy, misty, and yellowedge gmupers) in One option for the protection of grouper, in psrticular, and reef stocks, in southeastern United Swtes. as a result of recent stock analyses (SAFMC News gene&. may be marine fshery reserves (areas set aside for no consumptive Release - Nov. 7,1990). usage) (PDT, 1990). Carefully located, such reserves would. among other things. serve to protect spawning stock biomass and recruitment, maintain SUMMARY population age smcrure and the balance of natural communities, insure against Information on the status of grouper stocks in the tropical western Atlantic growth overfihmg and management failurc, and provide undisturbed provides undeniable testimony of the vulnerability of grouper species to anything underwater park areas for public enjoyment (PDT, 1990). of pdcular but low levels of fishing effort, in the absence of regulation. For almost dl smks imponance for tourism development Indeed, marine fishery reserves may be the assessed, there have ken sharp ddecnes in mean size and weight reduced only realistic solution to managing multispecies fisheries, given the lack of landings and CPUE, and loss of, or ahing reductions in, spawning enforcement and the limitations of stock monitoring, assessment and aggregations, all classic indications of overfishing. Hence. many stocks in both management capabilities regionwide. Another option to be explored is continental and islicnd mas am growth andfor recruitment overfihed. More mariculture mcker, 1994). While only in the very earliest stages of s@ingent management measures, and their enforcement, am badly needed. At the development, concerning the technical details of gouper culhlre, mariculture very least, spawning aggregations should be protected, monitoring programs holds some promise for the future. It will not, however, substitute for stock implemented or refined, and management options evaluated. Some species have management if we wish to continue exploiting natural populations. been more heavily impacted than others. Nassau grouper, for example, have Finally, while the focus of this paper hm been gouper, many of the shown paniculariy dramatic decliies in both mainland United Sues and island comments concerning use and management of ffieries resources apply to other arras, and many spawning aggregations have been so badly depleted that the components of the fishery. There clearly and urgently needs to be a change in species may be considered extinct for fisheriespurposes in a number of lmuons, perception regarding the exploitation of marine resources. Management must to The largest of the grouper species, the warsaw grouper and jewtish, have also be seen to be an integai component of sustainable resource use. Communication been heavily impacted, and have declined markedly off mainland United States. between fisheries offtcers and those in government respn~iblefor careraking As larger gouper have disappeared from landings, smaller individuals and public resources must improve. There is now no longer any excuse for species have come to comprise an increasingly grcater proportion of thecatch. proposing or promoting the development of marine fisheries resources without a Considenble data are required to better understand and manage gouper scienac database an which such development must be founded. The lessons stocks. Data collection prognms necessary to monitor these resources must be learned from past failures following poorly-planned attempts to "develop" must long-term and well-planned, and should be s-zed over as wide a not be forgotten. geographic area as possible because of the possibility of shared stocks over extensive regions. Socio-economic data also necd to be collected to determine ACKNOWLEDGEMENIS the possible impact of management measures on various resome user groups. A I wish to thank Jim Bohnsack for access to unpublished manuscripts. I am number of biological questions, particularly those regarding recruitment and grateful to Jim Bohnsack and Miguel Figuerota for reviewing various versions reproduction, need to be addressed Much of thin information is relatively simple to collect, with the appropriate planning and mining of personnel. Proceedings of the 43rd Gulf and Caribbean Fisheries Institute Peer Revlewed Section

of this paper. I also thank Cwlos Cumpiano and Ishmael Rivas for sharing with Bohnsack, J. k 1989. Protection of grouper spawning aggregations. Coastal me their extensive knowledge of the groupcr fshery of Puerto Rico. I am Resources Division Contribution 88-89-06. graeful to the Gulf and Caribbean Fisheries Institute for providing the forum for Bobnsack. J. A. Black and Nassau grouper fishery irends. Unpublished the 1990 grouper symposium. for which this paper was elaborated. A modir~ed manuscript version of this manuscript was presented at the 1990 nmual meeting of the Bohnsack. J. A,, D. L. Sutherland. A. Brown. D. E. Harper, and D.B. McClellan. American Fisheries Society. I wish to than Herb Kumpf for organizing the 1986. An analysis of the Caribbean biostatistical database for 1985. Pan-Caribbean Session at the AFS meeting. Coastal Resource Division Report for the Caribbean Fishery Management Council. Contribution Number CRD-86/87-10. LITERATmE ClTED Burnett-Herkes, J. 1975. Contribution to the biology of the red hind. Aguilar-Peren. k 1994. Preliminary obscwations of the spawning aggegarion Epinephelus gutratus, a commercially imponant settanid fsh Emm the of Nassau grouper, Epinephelus strintus, at Mahahual. Quintam Roo, uopical westem Atlantic. Ph.D. Disscttation, Univ. Miami, Coral Mexico. Proc. Gulfand Caribb. Fish. Inrr. 43: 111-121. Gables. Florida. 154 p. Appeldwm. R. S., G. D. Dennis. and 0. Monlerrosa Lopez. 1987. Review of Bush. P. G. and G. Ebanks-Petrie. 1994. The Cayman Island Nassau gmuper shared demersal resources of Puerto Rico and the Lesser Antilles study - A progress repoh. Proc. Gulf and Caribb. Fish. Insf. region. In: Reporr andproceedings of rhe expert conrultafion on shnred 43389-390. fishery resources of rhe Lesser Aniilles region. Mayagriez, Puerro Rico, Caddy, J. F. 1986. Size frequency analysis in stock assessment - some 8-12 September 1986. FA0 Fisheries Repon, 383:36-106. petspeclives, approaches and problems. Proc. Gulf and Caribb. Fish. Bannerot, S. P. 1984. The dynamics of exploited groupers (Serranidnc): An Insi 39:212-238. investigation of the protogynous hetmaphnxlitic reproductive strategy. Caner, 1. 1988. Grouper mating ritual on a Caribbean reef. Undenvarer Ph.D. Dissertation, Univ. Miami, Coral Gables, Florida. 393 p. Naturalist17:8-11. Bannerot. S. P., W. W. Fox Jr., and J. E. Powers. 1987. Reproductive strategies Caner, J.. G. 1. Mmow, and V. Pryor. 1994. Aspecs of the ecology and and the management of snappers and goupers in the Gulf of Mexico reproduction of Nassau grouper. Epinephelus striarus. off the coast of and Caribbean. In: J. J.Polovina and S. Ralston (eds.). Tropical Belie, Cenual America.. Proc. Gulfand Caribb. Fish. In&. 43:64-109. snappers and groupers: biology and fisheries management Westview Colin, P. L., D. Y. Shapim, and D. Weiler. 1987. Aspects of the reproduction of Press. Boulder, Colondo. pp. 561-603. two species of groupers. Epinephelus gutfatus and E. srriarus in the Baisre, J. A. and 3. Piiez. 1981. Los recursos pesquems delarchipielago cuhano. West Indies. Bull. Mari. Sci.. 40(2):220-230. Estudios WECAF No. 8. PFDgtama interregional de ordenacion y Colin, P. L. 1988. Management and Issues. Grouper Wafcher 1(2):4. desvrollo pesqueros. FAO-Naciones Unidas. 79 p. Colin, P. L. in press. Reproduction in the Nassau grouper. Epinephelus srriarus Beets, J. and A. Friedlander. 1992. Stwk analysis and management strategies (Piies: Sernnidae) with relationship to eovironmenral conditions. for red hind, Epinephelus gurmrus, in the U. S. Virgin Islands. Proc. Environmental Biology of Fishes. Gurand Caribb. Fish Conuens, M.. F. Aneguin-Sjnchcz, J. A. Sbnchez, V. Moreno, and Im-~~ 4266-RO ~ Bee& J. and k Friedlander. and W. Tobias.1994. Stock analysis of coney. M. A. Cabrera. 1994. Mortality and population size of the red grouper Epinephelusfulvus, in St. Croix, U. S. Virgin IslandF. . Proc. Gulfand Epinephelus morio) fishery from the Campeche Bank. Proc. Gulfand Caribb. Fish. Imt. 434051118, Canbb. Fish. IN[. 43:394-403. Evemann, B. W. 1900. Fishes and Fisheries of Porto Rico. United States Bevenon. R. J. H. and S. J. Holt. 1957. On rhe dynamics ofexploited fish Commission of Fish and Fisheries, Washington, D. C. 350 p. populations. United Kingdom Minimy ofAgriculm and Fisheries., Fischer, W. (ed). 1978. FA0 species identifcation sheets forj'ishery purposes. Fisheries Investigation (Series 2)19.533 p. Western Central Atlanric (Jishing area31). Vols. 1-7. Rome, FAO. Blanco, W.. R. Vald& y A. PCrez. 1980. Evalwi6n de lapesqueria de la chema Gilmore, R. G. and R. S. Joncs. 1994. Color variation and associated behavior in Mycreroperca microlepis americana (Epinephelus morio,Perciformes. Semidae), en-.. el-- --.""hanrn ""rie the Epinepbeline groupers, (Goode and Bean) Campeche. Revista Cubana de Investigacione$ Pesquerns 5: and sJonian and Swain.. Proc. Gulfand Caribb. Fish. 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Gobcrt. B. 1994. Preliminary analysis of the exploitation of groupers in Manwh. C. S., III and D. L. Masan. 1987. Age and gmwth of the warsaw Maninique.. Proc. Gulfand Catibb. Fish. Im. 43:447-456.s grouper and black grouper hmthe southeast region of the United Goodyear, C. P. 1988. Yield per renuit analysis for sevenl reef fish species of States. Northeasr Gulf Science 9(2):65-75. the Gulf of Mexico. A report to the Gulf of Mexico Fishery Matheson, R. H., El and G. R. Huntsman. 1984. Growth, monaiity, and Management Council by the Miami Laboratov, Southeast Fisheries yield-per-recruit models for speckled hind and snowy grouper from the' Center, NOAA. U. S. Department of Commerce. CRD 87/88-14. United States South Atlantic Bight. Trans. Am. Fish. Sac. lU:M)7-616. Goodyear, C. P. 1989. Spawning stock biomass per recmic The biolo&cal basis Matos, D. and Y. Sadovy. 1990. Overview of Puem Rim's small- scale for a fisheries management, tool. -ICCAT Worlring Document fisheries statistics 1988-1989. Fisheries Reswch Laboratory, SCRS/89/82. I0 n. CODREMAR, Department of Natural Resources-Puem Rico, Huntsman. G. R. and R. L. Dixon. 1975. Recreational wrches of four species of Technical Repon 1(4):1-17 Volume 1. Number 4. groupers in the Carolina headboat fishery. Pmceedings of the 29th Mae, M. A. 1969. Biology of the red grouper, Epinephelus morio Annual Conference of the Southeastern Association of Game and Fish (Valenciennes) from the eastern Gulf of Mexico. Florida DeparUnent of Commissioners, 1975:185-194. Natural Resources. Marine Research Laboratory Professional Papers Huntsman, G. R.. C. S. Manwh 111, and C. B. Grimes. 1983. Yield per recruit Series 10:l-95. models of some reef fishes of the U. S. South Atlantic Bight. Fishery Moore, C. M. and R. F. Labisky. 1984. 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Polovina and S. Ralston (eds.). Tropical snoppers and groupers: Oisen, D. A. and J. A. LaPlace. 1979. A study of a Virgin islands grouper biology and frrheries managemenl. Wes~iewPress, Boulder, Colorado. fishery based on a breeding aggegation. Proc. Gulf and Caribb. Fish. pp. 533-560. inrr 31:130-144. Huntsman. G. R. and P. W. Willis. 1989. Status of reef fish stocks off Nonh Pauly, D. 1980. A selection of simple methods for the assessment of tropical fish Camlina and South Carolina as revealed by headboat catch statistics. stocks. FA0 Fisheries Circular No.729.54~. National Oceanographic and Atmospheric Administration-National Pauly. D. 1987. A review of the ELEFAN system for analysis of Underwater Research Program Report, 89-2. length-frequency data in fish and aquatic invertebrates. pp. 7-34 In: D. Kimmel, 1. J. and R. S. Appeldwrn. In press. A critical review of fisheries and Pauly and G. R. Morgan (eds.) Length based methods in fisheries %heries management policy in Puem Rico. 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Criteria for the diagnosis of Wildlife Canservarion International hermaphroditism in f~hes.Copeia 1987 (1):136-156. Division of New York ZoologicnI Society Sadovy, Y. and P. L. Colin. 1990. Hermaphroditism in the Nassau grouper, Bronx. New York 40005 Epinephelus sniarus. American Society of Ichthyologists and ZDepartment of Life Sciences Herpetologists Annual Meeting, Charleston. 1990 (Abstract). University of New England Sadovy. Y., and M. Figueroia, 1992. The status of the red hind fishery in Puem Biddeford, Maine 04005 Rico and St Thomas. as determined by yield-per-recruit analysis. Proc. 30ceanography Program Guyand Caribb. Fish. Inst 42:23-38. University of Delaware Sadovy, Y., Shapiro. D. Y.. and M. A. McGehee. In press. Size, composition College of Marine Studies and spatial smcture of the annual spawning aggregation of the red Lewis, DE 19958-1298 hid. Epinephelus guttatus (Pisces: Serranidae). Copeia. Shapiro, D. Y. 1987. Repmduction in gtoupers. In: J. J. Polovina and S. Raiston ABSTRACT (eds.) Tropical snappers and groupers: biology md fisheries Aspects of the fie histoiy of Nassau grooqer, Ep!nephe!m suiatur, f@m management. Westview Press, Boulder, Colorado. pp. 295-327. Befi in the western Caribbean arc described, iclu&ng food hablLS. Smith, C. L. 1971. A revision of the american groupers: Epinephelus and allied movements, pmtogyny, sexual matuntion, ~nality,penodicity of spawnins. fecundity, and papuladon sex ratios. Adduona! information is provided on $c genera. BuIIerin of the American Museum. of Natural History 146 potential effect overfishing spawning nggepuons has on future rcpmduc~ve (2):67-242 potential of the population. Smith, C. L. 1972. A spawning aggregation of Nassau grouper. Epinephelus Gonad structure and ontogenetic development,p?ttems are described and sfriarus(Blcch). Trans. Am. Fish. Soc. 101:257-261. ev~dencefor sex-revers& is given. Seasonal vanmon in lhe percentnge of Stevenson. D. K 1978. Management of a tropical fish pot fishery for maximum sexually active fishes indicate the majority of fishes aggregated and spawned along the shelf edge at the reef promontories around the full moon m December sustainable yield. Proc. Gulf md Caribb. Fish. Imt 30:95-115. and January. A comparison of sex ntios and length frequency disuibution Suhz-Caabm. 1. A. 1970. Pueno Rico's fmhery sratisucs 1968-1969. among three sites indicates that there is a mked increase in the number of Departamento de Agricuitura de Pueno Rico, Conaibuciones females relative to males at heavily fished sites, as well as a sipinificant decrease Agropecuarias y Pesqueras 2(1):l-38. in the size of both sexes relative to the genenl papuletion oulside of the Waters, J. R., and G. R. Huntsman. 1986. Incorporating monality from catch spawning banks and at pristine spawning aggegauon sites. Spawning of Nassau grouper is discussed in relationship w environmental and release into yield-per-recruit analyses of minimum-size limits. conditions (e.g., reef geomorphology, tempenm, photopenod. cunenl e~.). North American Journal of Fisheries Management 6:463471. Finallv.~.~.... , food habits of Nassau grouper are described and related to fonglng suategy and daily patterns of mo;ement

KEYWORDS: Belize, ecology, pmwgyny. reefs, reproducuon, sen;lnidae. Nassau grouper. Proceedings of the FORTY-THIRD ANNUAL Gulf and Caribbean Fisheries Institute

MIAMI. USA NOVEMBER, 1990

Library of Congress Catalog Card Number 52-33786 Edited by Melvin H. Goodwin and Gregg T. Waugh Senior Editor - Peer Review Mark J. Butler Associate Editors - Peer Review Richard S. Appeldoorn John Hunt Brian E. Luckhunt Wiliam Lyons David Miller Yvonne Sadovy

CHARLESTON, SOUTH CAROLINA 1994 CONTENTS SECTION A: PEER REVIEWED PAPERS

GULF AM) CARIBBEAN IXSERlES - GENERAL Harper, DE.. J.A. Bohnsack, Investigation of Bycatch fmm and D. McClellan the Wm F&-Trap Fshery in Federal Waters off Southem FIorida 3 STOCK ASSESSMENTS AND TROPEIC STUDIES Godez. ME., J.A. Sanchez. Population Dyoamics of the and F. Arreguin-Sanchez Red Snapper (Lutjnnus cmpechnnus) fishery fmm Campeche Bank, Mexim 29

SYnIPOSIUM ON THE BIOLOGY AND MAVAGEhIENT OF GULF AND CARIBBEAN GROUPERS Sadovy, Y. Gmuper Stocks of the Western Central Atlantic: The Need for Management and Mmagement Needs 43 Caner, J. Aspects of the Ecology and Reproduction of Nassau Grouper (Epinephelus strialus) off the Coast of Belize, Central America 65

Aguiiar-Pw%~,A. Preliminary Obse~ationsof the Spawning Aggregation of Nassau Gmuper, Epinephplus striatus. at Majahual, Quintana Rw,Mexico 112 Stiles, T.C. and MA. Bunon Age. Growth, and Mortality of Red Gmuper, Epinepltelus morio. from the Southeastern U.S. 123 Colin, PL. Preliminary Investigations of Reproductive Activity of thc Jewfish, Epinephelus itajara (Pisces: Scnanidne) 138 ,