ent OF Coy, Marine Fisheries %,6 *. S * < € WY 1991 w i National Oceanic and Atmospheric Administration + National Marine Fisheries Service 7 Ly a ~; = Dd ~ ~ a i | Ph | hs | ~~ ms FALE + MtTh t » BEILPHP IT eu. com |4j Menhaden On the cover: A menhaden fishing scene. Special Issue 53(4), 1991 Menhaden: The Resource, the Industry, and a Management History Preface Douglas S. Vaughan Population Biology and Life History of the North American Menhadens, Brevoortia spp. Dean W. Ahrenholz Overview of Mark-recovery Studies on Adult and Juvenile Dean W. Ahrenholz, Atlantic Menhaden, Brevoortia tyrannus, and Gulf Menhaden, B. patronus Donnie L. Dudley, and Eldon J. Levi The Atlantic and Gulf Menhaden Purse Seine Fisheries: Origins, Harvesting Technologies, Biostatistical Monitoring, Recent Trends in Fisheries Statistics, and Forecasting Joseph W. Smith 28 New Products and Markets for Menhaden, Brevoortia spp. Malcolm B. Hale, Paul E. Bauersfeld, Sylvia B. Galloway, and Jeanne D. Joseph 42 Assessment and Management of Atlantic Menhaden, Brevoortia tyrannus, and Gulf Menhaden, B. patronus, Stocks D.S. Vaughan and J. V. Merriner 49 Biological Analysis of Two Management Options for the Atlantic Menhaden, Brevoortia tyrannus, Fishery Douglas S. Vaughan and Joseph W. Smith 58 U.S. DEPARTMENT OF The Marine Fisheries Review (ISSN 0090-1830) the Director of the Office of Management and is published quarterly by the Scientific Publica- Budget. COMMERCE tions Office, National Marine Fisheries Service, The NMFS does not approve, recommend or en- NOAA, 7600 Sand Point Way N.E., Bin C15700, dorse any proprietary product or proprietary Barbara Hackman Franklin, Seattle, WA 98115. Single copies and annual material mentioned in this publication. No refer- Secretary subscriptions are sold by the Superintendent of ence shall be made to NMFS, or to this publica- Documents, U.S. Government Printing Office, tion furnished by NMFS, in any advertising or sales NATIONAL OCEANIC AND Washington, DC 20402: Single copy, $5.50 promotion which would indicate or imply that ATMOSPHERIC ADMINISTRATION domestic, $6.88 foreign; annual subscription, NMFS approves, recommends, or endorses any $9.00 domestic; $11.25 foreign. proprietary product or proprietary material men- John A. Knauss, Under Secretary Publication of material from sources outside tioned herein, or which has as its purpose an in- the NMFS is not an endorsement and the tent to cause directly or indirectly the advertised for Oceans and Atmosphere NMFS is not responsible for the accuracy of product to be used or purchased because of this facts, views, or opinions of the sources. The NMFS publication. Second class postage is paid William W. Fox, Jr., Assistant Secretary of Commerce has determined that the in Seattle, Wash., and additional offices. POST- Administrator for Fisheries publication of this periodical is necessary for MASTER: Send address changes for subscriptions the transaction of public business required by for this journal to—Marine Fisheries Review, c/o National Marine Fisheries Service law of this Department. Use of the funds for Superintendent of Documents, U.S. Government Editor: W. L. Hobart printing this periodical has been approved by Printing Office, Washington, DC 20402. MENHADEN: THE RESOURCE, THE INDUSTRY, AND A MANAGEMENT HISTORY A Special Issue of the Marine Fisheries Review Smaller purse-seine menhaden fisheries for bait (e.g., crab and lobster) are found on both coasts. Preface The National Marine Fisheries Service (formerly Bureau of Commercial Fish- DOUGLAS S. VAUGHAN eries) has maintained records of daily vessel landings and fishing activity since 1940 onthe Atlantic coast and since 1946 on the Gulf coast. Sampling for age and size of menhaden landed has been con- Four species of menhaden, Brevoortia currents where they metamorphose and ducted by staff in the NMFS menhaden spp., are found along the Atlantic and develop as juveniles. research program since 1955 on the Gulf of Mexico coasts of the United Menhaden form large surface schools Atlantic coast and since 1964 on the States. The Atlantic menhaden, B. tyran- susceptible to purse seines which are Gulf coast. Captain’s daily fishing nus, is found from Nova Scotia, Can. , to now the principal fishing gear. Major reports containing data on individual West Palm Beach, Fla.; the yellowfin fisheries for Atlantic and Gulf menha- menhaden purse-seine sets have been menhaden, B. smithi, is found from Cape den exist on each coast (Fig. 1). Although collected on both coasts since 1978. In Lookout, N.C., tothe Mississippi River neither fishery is directed toward the addition, extensive mark-recapture Delta, La. ; the Gulf menhaden, B. patro- yellowfin or finescale menhaden, both studies (using internal ferromagnetic nus, is found from Cape Sable, Fla., to may be part of these other catches to a tags) on adult and juvenile Atlantic and Veracruz, Mex.; and the finescale men- small degree. During the 1980s, the com- Gulf menhaden have been conducted haden, B. gunteri, is found from the Mis- bined landings for the menhaden reduc- since the late 1960’s. Also, studies on sissippi River Delta, La. , to Campeche, tion fisheries comprised about 40% ofall the biology and estuarine distribution Mex. U.S. commercial landings, ranging be- of juvenile menhaden have been con- Menhaden are euryhaline species that tween 934,000 t and 1,342,000 t, with ducted along the U.S. Atlantic and Gulf inhabit coastal and inland tidal waters. Gulf menhaden landings representing coasts since the early 1960’s. These Spawning occurs principally at sea (in between two-thirds and three-fourths studies culminated in extensive juvenile northern areas some spawning occurs in of the total. Menhaden are processed abundance surveys along both coasts bays and sounds). Eggs hatch at sea and primarily at reduction plants for fish during the 1970’s. the larvae are moved to estuaries by ocean meal, oil, and solubles. The meal and Management of Atlantic and Gulf men- solubles are used mostly in poultry and haden fisheries is by the respective states Douglas S. Vaughan is with the Beaufort Labora- livestock feeds, and increasingly in and coordinated through the Atlantic and tory, Southeast Fisheries Science Center, National aquaculture, whtihe olil ies u sed in paints Gulf States Marine Fisheries Commis- Marine Fisheries Service, NOAA, Beaufort, N.C. 28516. and as an edible oil in Europe and Canada. sions. State, industry, and Federal inter- 53(4), 1991 LANDINGS OF ALL BILLIONS OTHER OF SPECIES IN U.S. POUNDS b4y]wa s SANAARAMRANAANRAJNAR ARARRSANASRS SISS MSS 77% SSS T NNANANNANANMMAAAANSSSSS)A DASOOAOXOASORYODSO AOOSNOAYO N AO NASsN ARAASAORSMMNASRSAAQ ASN[ AMA N AAAAAAS[AQASAAA NSSA A NNJQS R A MAetnlhaantdiecn NANSASASA ANRSARSANRSAAN NAMAAANAA)A A NANNAAANANAAAAANAANA NARSPARQAEAAAAN RAMNASS Fishery SMMSASSS SQ SM — GULF AND ATLANTIC MENHADEN ~ 23% 1970 71 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 8889 CALENDAR YEAR - .. ; Ca: Gulf Menhaden Fishery WACTIVE PLANT LOCATIONS IN 1989, AU.S.-SOVIET IWP VENTURE Figure 1.—Gulf and Atlantic menhaden contribution to total U.S. commercial fisheries landings during the calendar year, 1970-87. ests are represented on the Atlantic and This special issue of the Marine Fish- interactions including a comparison of Gulf Menhaden Advisory Committees as eries Review provides an historical management options for the Atlantic part of the interstate commission fishery perspective on the menhaden resource menhaden. These papers draw on management process. Information from and fisheries. These papers summarize previously published material and on NMFS is provided to these groups, in- information on menhaden biology, current research. They are intended to cluding stock assessments on these spe- environmental influences on recruit- help define ‘‘the state of our knowledge’”’ cies. Coastwide management plans have ment, results of tagging studies, the and provide guidance in developing been prepared for both Atlantic and Gulf fisheries, current and potential products future studies to improve our understand- menhaden with recent updates for both from menhaden, recent assessments of ing of menhaden biology and population species. stock status, and describe management dynamics. Marine Fisheries Review Population Biology and Life History of the North American Menhadens, Brevoortia spp. DEAN W. AHRENHOLZ Introduction sumers, feeding on phytoplankton, or Geographic Ranges secondary consumers, feeding on zoo- Menhaden are members of the world- plankton, or both. Many clupeids are Reintjes (1969) summarized the geo- wide family Clupeidae, one of the most in turn prey for various piscivorous graphic ranges for the four menhaden important families of fishes both eco- predators through virtually their entire species. Atlantic menhaden are seasonal- nomically (Hildebrand, 1963), and eco- lives. Life history patterns for this fam- ly found from Nova Scotia, Can., to logically. Clupeids are characteristically ily of fishes include species which can southeastern Florida, near West Palm very numerous and form large, dense complete their entire life cycle in either Beach. Gulf menhaden range from schools which enhance our ability to fresh or marine waters, or are anadro- southwestern Florida, near Cape Sable, harvest them. Many of the species are mous species, or marine migratory (estu- to Veracruz, Mex. Yellowfin menhaden filter feeders, being either primary con- arine dependent) species. overlap the ranges of all three other The large-scaled menhadens, the At- menhaden species and are found from lantic menhaden, Brevoortia tyrannus, Cape Lookout, N.C., to the Mississippi and the Gulf menhaden, B. patronus, River Delta. Finescale menhaden over- have received considerable attention in lap the ranges of both the Gulf and fishery science research due to their large yellowfin menhaden, and are found from ABSTRACT—Four recognized species of menhaden, Brevoortia spp. , occur in North population sizes and resulting economic just east of the Mississippi River Delta American marine waters: Atlanticmenhaden, and ecological importance. The small- (Turner, 1971) to Campeche, Mex. B. tyrannus; Gulf menhaden, B. patronus; scaled menhadens, the yellowfin men- The numbers of Gulf menhaden rela- yellowfin menhaden, B. smithi; and finescale haden, B. smithi, and the finescale men- tive to numbers of yellowfin menhaden menhaden, B. gunteri. Three ofthe menhaden haden, B. gunteri, are less numerous become reduced proceeding southward species are known to form two hybrid types. Members of the genus range from coastal and have received far less consideration on the Gulf of Mexico coast of Florida. waters of Veracruz, Mex., to Nova Scotia, in the scientific literature. The contrast in There appears to be a similar distribu- Can. Atlantic and Gulf menhaden are ex- relative importance is quite marked. On tion pattern for relative numbers of At- tremely abundant within their respective one extreme, the purse-seine reduction lantic and yellowfin menhaden pro- ranges and support extensive purse-seine reduction (to fish meal and oil) fisheries. All fishery (to fish meal and oil) for Gulf ceeding southward along the Atlantic menhaden species are estuarine dependent menhaden was the largest U.S. fishery by coast of Florida. The coastal area be- through late larval and juvenile stages. De- weight from 1963 through 1988, and tween West Palm Beach and Miami, Fla. , pending on species and location within the Atlantic menhaden purse-seine reduction where menhaden are relatively rare range, spawning may occur within bays and landings, currently one-third to two- (Dahlberg, 1970), geographically sep- sounds to a substantial distance offshore. Menhadenare considered to be filter-feeding, thirds those for Gulf menhaden, were the arates the Atlantic menhaden from the planktivorous omnivores as juveniles and largest for the U.S. from 1947 to 1962. Gulf menhaden, as well as apparent east- adults. Menhaden eggs, immature develop- On the other extreme, finescale menha- ern and western populations of yellowfin mental stages, and adults are potential prey den are apparently not directly sought by menhaden. for alarge and diverse number of predators. North American menhadens, including two any recognized fishery, and yellowfin A large amount of hybrid introgression hybrids, are hosts for the parasitic isopod, menhaden (and their hybrids) are only occurs between Atlantic and yellowfin Olencira praegustator, and the parasitic cope- harvested by specialized bait fisheries on menhaden on the Atlantic coast of pod, Lernaeenicus radiatus. Although the data both coasts of Florida. The following is Florida, and Gulf and yellowfin men- are quite variable, a dome-shaped Ricker a general description of the population haden onthe Gulfcoast of Florida. Areas function is frequently used to describe the spawner-recruitment relationship for Atlan- biology and life history of these four with pure strains of yellowfin menhaden ticand Gulfmenhaden. Each of these species North American menhaden species. are yet tobe defined. As the relative den- is treated as a single stock with respect to ex- sity of Gulf menhaden decreases pro- ploitation by the purse-seine reduction fishery. Dean W. Ahrenholz is with the Beaufort ceeding southward, the number of Gulf Estimates of instantaneous natural (other) Laboratory, Southeast Fisheries Science Center, x yellowfin menhaden (B. patronus x mortality rates are 0.45 for Atlanticmenhaden National Marine Fisheries Service, NOAA, and 1.1 for Gulf menhaden. Beaufort, NC 28516-9722. B. smithi) hybrids increases along with 53(4), 1991 pure strains of yellowfin. For example, Table 1.—Distinguishing and comparative characteristics of North American coastal menhadens (modified from Dahiberg, 1970). Turner (1969) reported that collections of menhaden from Panama City to Cedar Large-scaled menhaden Small-scaled menhaden Keys, Fla., consisted of 94% Gulf men- Character B. tyrannus B. patronus B. smithi B. gunteri haden and 6% yellowfin menhaden, Frontal groove Complete Complete Absent Absent while samples from farther south, Tam- Lateral spots Usually present Usually present Absent Absent pa Bay to Cape Sable, Fla., were 7% Gulf above and below above and below the level of the level of menhaden, 56% yellowfin menhaden, shoulder spot shoulder spot and 37 % Gulf X yellowfin hybrids. Het- Ventral fin Middle rays and Inner rays equal Inner rays about Inner rays about sometimes inner to or longer than one-half to two- one-half to two- tler (1968) reported on two collections rays equal in length outer rays thirds length of thirds length of made along the southern Gulf coast of to outer rays fin fin Scale Pointed, length Pointed Rounded tip, Rounded tip, Florida; one near Naples consisted of pectinations ' medium or long shorter shorter 17% Gulf menhaden, 9% yellowfin Body mucus” Copius Copius Sparse Sparse menhaden, and 74% Gulf x yellowfin Flesh? Soft Soft Firm Firm Ovarian color Yellow Yellow White ? hybrids, and the other from near Sanibel Lateral scale rows 43-53 (40-58) 42-48 57-73 (54-80) 65-72 (60-76) Island consisted of 5% Gulf menhaden, Opercular Prominent Prominent Faint or absent Faint or absent 54% yellowfin menhaden, and41% Gulf striations (12-31) (13-25) (0-15) (0-18) x yellowfin hybrids. A similar situation Predorsal scales 35-44 (33-46) 29-37 (28-39) 39-51 (37-56) 40-49 (39-52) Vertebrae 46-48 (44-49) 44-46 (43-47) 44-45 (43-46) 42-43 (41-43) apparently exists on the east coast of Ventral scutes 31-34 (29-34) 29-31 (28-32) 30-32 (29-34) 28-30 (27-31) Florida with the distributions of Atlantic ' Older adults. and yellowfin menhaden and the Atlan- ? Fresh specimens. tic X yellowfin hybrids; for example, the menhaden gill-net fishery in Indian River, Fla., is dominated by yellowfin two large-scaled menhaden species can shoulder spot, and have larger and fewer menhaden and the Atlantic X yellowfin be separated from the two small-scaled scales. The small-scaled menhadens hybrids (Dahlberg, 1970). species by a variety of characteristics lack the former characteristics and have (Table 1). Fresh specimens can be sep- smaller and more numerous scales. Gulf Species Characteristics arated simply by feel, as the large-scaled menhaden have a deeper (more convex) Menhaden are generically distin- menhadens have large amounts of body body shape and fewer predorsal scales, guished from other clupeids by their mucus and relatively soft flesh, while the vertebrae, and ventral scutes than their relatively large heads, pectinated scales, small-scaled species have relatively small Atlantic congener (Table 1, Fig. 1, 2). absence of teeth (beyond juvenile stages), quantities of body mucus and their flesh The yellowfin menhaden can be separ- and by their dorsal fin being over the in- is firm. Additionally, the large-scaled ated from the finescale menhaden by the terval between the pelvic and anal fins species possess a frontal groove, acces- yellowfin’s greater number of vertebrae (Reintjes, 1969; Hildebrand, 1963). The sory lateral spots beyond the large and ventral scutes, and relatively smaller \ \ i 12} j bs |E Y Re s' f 7 € a Figure 1.—Adult Atlantic menhaden, 250 mm FL (J. W. Reintjes photo). Marine Fisheries Review Figure 2.—Adult Gulf menhaden, 167 mm FL (R. B. Chapoton photo). Figure 3.—Adult yellowfin menhaden, 300 mm FL (J. W. Reintjes photo). head (Table 1, Fig. 3, 4). More detailed crossing with either parental population Hybrids of B. gunteri x B. patronus descriptions are available from Dahlberg will be predominantly by male hybrids, (finescale < Gulf menhaden) and B. (1970) and Hildebrand (1963). Dahlberg as they dominate the hybrid population. gunteri X B. smithi(finescale x yellow- (1970) also provides divergent char- A self-sustaining population of hybrids is fin menhaden) have not been reported. acteristics between the Atlantic and Gulf unlikely due to the preponderance of Although the ranges of the three species populations of yellowfin menhaden. males. Hettler (1968) found no female overlap, yellowfin and finescale men- The morphological and morpho- hybrids (B. patronus X B. smithi), while haden are not abundant in the area of metrical appearances of the large- Turner (1969) reported finding4 f emales overlap (the Mississippi Delta region). scaled menhaden and yellowfin men- out of 390 hybrids examined. Dahlberg Except for the southeastern Texas coast, haden hybrids are intermediate to those (1970) discovered one female hybrid (B. finescale menhaden are apparently not for the parents (Dahlberg, 1970) (Fig. 5). tyrannus X B. smithi) fromanunknown abundant in U.S. Gulf coastal waters The presence of a gradient of character- number of hybrids examined on the where Gulf menhaden predominate. istics between the parental types sug- Atlantic coast, and found no females Definitive studies on finescale menhaden gests back-crossing also occurs. Back- among Gulf hybrids. are lacking. 53 (4), 1991 vais: aes soe aspshi - Figure 4.—Adult finescale menhaden, 320 mm FL (R. B. Chapoton photo). General Life Cycle existing knowledge of migration and ward. By June, the population is redis- distribution was further strengthened tributed from Florida to Maine. Even Menhaden are estuarine dependent, by an analysis of the age and length though some Atlantic menhaden migrate marine migratory species. Spawning distributions of Atlantic menhaden in the north and south along the U.S. Atlantic generally occurs during the cooler landings (Nicholson, 1971), and finally coast, because the fish distribute them- months in the marine environment, from results of an internal, ferromagnetic selves on the basis of size and age, the and larvae undergo early growth and tagging program (Dryfoos et al., 1973; movement actually represents a seasonal development at sea. About 1-2 months Kroger and Guthrie, 1973; Nicholson, expansion and contraction of the Atlan- later, those larvae that have been trans- 1978). tic menhaden’s range. ported shoreward enter estuarine bays, During summer, Atlantic menhaden Geotemporal aspects of spawning for sounds, and streams, and metamorphose are generally distributed from northern this species are closely associated with the into juveniles. Menhaden juveniles Florida to Maine. The adult population migratory behavior of the adults, and (young-of-the-year) normally reside in stratifies by age and size, with the older some degree of spawning activity is estuarine areas until the following fall or and larger individuals farther northward believed to occur during virtually every early winter when many migrate into and the younger and smaller fish in the month of the year. Some fish ripen and marine waters. Adults generally occur in southern half of the species’ range. Al- some spawning occurs in the more north- nearshore oceanic waters and frequent- though localized movements occur dur- erly portions of the fishes range as the ly reside in large estuarine systems. ing summer, no major systematic move- fish begin moving southward in Septem- ment occurs until September, when the ber. Spawning continues with increasing Migratory Behavior more northerly portion of the population intensity as the fish move progressively and Spawning Season begins to migrate southward. By Decem- farther southward in October and No- ber, a significant portion of the adult vember. Spawning intensity is believed Atlantic Menhaden population that was north of Chesapeake to peak in waters off the North Carolina Early hypotheses of the migratory be- Bay during summer has moved south- coast during winter. Spawning con- havior of Atlantic menhaden were based ward to waters off the North Carolina tinues, but with decreasing levels of in- upon observations of schools appearing coast. These fish are followed by large tensity as the fish move northward the and disappearing along the U.S. Atlan- numbers of juvenile (young-of-the-year) following spring and early summer. Sup- tic coast, and from the examination of the menhaden, which have recently emi- porting evidence for these conclusions age and size composition of catches grated from nursery areas farther north. was obtained earlier by Higham and among fishing ports along the U.S. Atlan- Usually by late January, menhaden Nicholson (1964), subsequently by Ken- tic coast (June and Reintjes, 1959). An schools disappear and schools disperse dall and Reintjes (1975), and later by analysis of the frequency and distribution from nearshore surface waters of North Judy and Lewis (1983). Atlantic men- of purse-seine sets contributed additional Carolina. During March or early April, haden are believed to spawn in oceanic information with respect to the timing schools of adult menhaden reassemble in waters over much of the continental of migrations (Roithmayr, 1963). The coastal waters and move rapidly north- shelf, and in bays and sounds in Long Marine Fisheries Review Figure 5.—Adult yellowfin menhaden (upper), 280 mm FL, and Atlantic x yellowfin menhaden hybrid (lower), 285 mm FL (J. W. Reintjes photo). Island waters and northward (Nelson et of the age/size structure of the spawning U.S. Gulfcoast in nearshore waters. Be- al., 1977; Ferraro, 1980b). Evidence for population. For example, with the bulk ginning in October, they move offshore recent spawning activity was based on of the spawning stock in recent years into deeper waters for winter. Roithmayr the presence of menhaden larvae andeggs consisting of late age-2 fish, relatively and Waller (1963) reported that during in plankton samples. Evidence of immi- less spawning activity would have been summer Gulf menhaden occurred in nent spawning was also provided by expected in the New England and Mid- depths of 1-8 fathoms, while during the presence of near-ripe specimens in dle Atlantic areas as compared to the winter months they were found in 4-18 fish samples obtained from commercial 1950’s when a broader and stronger age fathoms east and west of the Mississippi purse-seine landings. However, spawn- structure was more extant in the popula- Delta, and at 20-48 fathoms in a smaller ing has not been directly observed in tion (Ahrenholz et al., 1987b). area east and northeast of the Delta. the marine environment, and running- Results of tagging studies failed to ripe females are rarely captured. Gulf Menhaden identify any east-west component of The relative magnitude of temporal Gulf menhaden do not exhibit an exten- annual migration for Gulf menhaden spawning activity within and between sive migratory pattern. During late spring (Pristas etal. , 1976; Kroger and Pristas, geographic regions is in part a function and summer they distribute along the 1975); however, multiple-year juvenile 53(4), 1991 tag-recovery data indicated a tendency by February 8 about 25 % of females were sexually mature during their second year for Gulf menhaden from the eastern and ready to spawn. Hettler’s (1970) speci- of life, while from two-thirds to nearly western extremes of their range to move mens from the Atlantic coast were taken all are sexually mature by the end of their toward the center of their range with age from the Indian River, Fla., in Febru- third year (age 2) (Higham and Nichol- (Ahrenholz, 1981). ary. Dahlberg (1970) concluded that son, 1964; Lewis et al., 1987). Female The spawning season for Gulf men- the spawning season for yellowfin men- Gulf menhaden about 150 mm FL and haden was determined by observations of haden was February and March on both larger are generally sexually mature by larvae, gonadal development, and pres- the Atlantic and Gulf coasts of Florida. the spawning season (Lewis and Roith- ence of eggs in plankton samples. Spawn- His conclusion for the Gulf coast was at mayr, 1981), while the smallest sex- ing has not been directly observed. From least in part based on Hettler’s (1968) ually mature female Atlantic menhaden observations of the occurrence of lar- collection just north of Naples during are at least 180 mm FL (Lewis et al., vae in Lake Ponchartrain, La., Suttkus mid-March of tworipe female yellowfin 1987). (1956) concluded that spawning prob- menhaden and Turner’s (1969) collection Age and size at maturation data is ably began in October and ceased in of ripe females during February and limited for the small-scaled menhadens. February; he presumed that this period March off the southern Gulf coast of Gunter (1945) observed a ripe female could fluctuate among years. Combs Florida. Spawning may occur as early as finescale menhaden 150 mm TL, anda (1969) concluded from a histological November, as Houde and Swanson ripe male 125 mm TL. The smaller of the examination of ovaries that spawning (1975) collected yellowfin menhaden two ripe female yellowfin menhaden that ranged from October through February eggs during this month from Atlantic Hettler (1968) found was 186mm FL. No or early March. Christmas and Waller waters off the Florida coast. standing stock ova counts for either (1975), after a literature review and an species of small-scaled menhaden are examination of plankton sampies col- Finescale Menhaden available. lected from much of the Gulf of Mexico, There is no evidence from which to Atlantic and Gulf menhaden are con- concluded that spawning ‘‘.. .for the deduce any systematic seasonal migra- sidered to be multiple (fractional or inter- most part. ..’’ occurred from October tion by the finescale menhaden other mittent) spawners (Higham and Nichol- through March. Shawetal. (1985a) pre- than the notation of an apparent seasonal son, 1964; Combs, 1969). As noted by sented arguments and evidence for an shift of larger finescale menhaden be- Combs (1969), the fishes’ ovaries could even more protracted season. tween Texas bays (Gunter, 1945). Like not contain all the developing ova if they Spawning areas have been determined the yellowfin menhaden, the finescale matured at the same time. Thus, ova by noting the geographic collection sites menhaden appears to occur more in es- mature and are spawned in batches over where Gulf menhaden eggs were taken. tuarine or nearshore areas. Gunter (1945) a protracted spawning season. Based on their own collections and the referred to it as a brackish-water form, as The potential number of ova produced work of Fore (1970) and Turner (1969), opposed to the more saline Gulf men- by an individual female during a spawn- Christmas and Waller (1975) concluded haden, although this species was not ing season has been determined (esti- that Gulf menhaden spawn from near formally described until 3 years later. mated) by counting the standing stock of shore to 60 miles offshore along the en- Gunter (1945) discovered a ripe male advanced oocytes in Atlantic menhaden tire U.S. Gulf coast. during February and a ripe female dur- (Higham and Nicholson, 1964; Dietrich, ing the latter part of March, and noted 1979; Lewis et al., 1987) and Gulf men- Yellowfin Menhaden that the spawning season was probably haden (Suttkus and Sundararaj, 1961; Adult yellowfin do not appear to dis- from midwinter to early spring. He also Lewis and Roithmayr, 1981). For this play any systematic, annual migratory observed post-larval finescale menhaden technique to provide a reasonable esti- behavior. Dahlberg (1970) referred to from January to May. Simmons (1957) mate of true annual fecundity, the num- them as “‘. ..common near shore along reported that this species spawned in the ber of ova produced during a season must both Florida coasts throughout the year.’ upper Laguna Madre of Texas during be annually determinate, like that of the He considered them an inshore or bay February. Given these observations, a multiple spawning Atlantic silverside, form (in contrast to the large-scaled spawning period of November to March Menidia menidia (Conover, 1985), as menhadens). Some larger individuals are appears realistic. Both Simmons (1957) opposed to being annually indeterminate, occasionally found as far north as Cape and Gunter (1945) reported that spawn- similar to the multiple spawning northern Lookout, N.C., during summer. ing occurs in inside (estuarine) Texas anchovy, Engraulis mordax (Hunter and Spawning seasons and some spawning waters. Macewicz, 1985). While some workers areas have been identified by collecting (e.g., Lewis et al., 1987) felt that deter- Maturation and Fecundity specimens for artificial spawning and minate fecundity is likely for the Atlan- rearing. For the Atlantic coast popula- Gulf menhaden become sexually ma- tic menhaden (and thus likely for the tion, Reintjes (1962) began sampling ture near the end of their second year other menhadens), this condition has near Sebastian, Fla., in November. He of life (age 1) (Lewis and Roithmayr, not been demonstrated, nor has batch noted ripening males in December, 1981). By comparison, only a small per- fecundity been estimated for any species several ripening females in January, and centage of Atlantic menhaden become of menhaden. Marine Fisheries Review