806 
Fishery Bulletin 96(4), 1998 
shore areas of South Carolina and Georgia to the shelf 
edge off Florida, perhaps to spawn. However, the 
presence of hydrated oocytes and new postovulatory 
follicles in female gag taken from North Carolina 
through Florida, including specific locations off South 
Carolina and Georgia, shows that spawning is not 
restricted to Florida. Hunter et al. (1985) reported 
that the presence of hydrated oocytes in teleosts is 
an indication that individuals will spawn within 12 
hours. Furthermore, Hunter et al. (1986) found that 
postovulatory follicles are present only in fishes that 
have recently spawned. 
Gag off Florida are subject to intense fishing pres- 
sure by commercial fishermen, sport divers, and an- 
glers, particularly during the late fall through early 
spring, because the continental shelf is very narrow 
and spawning aggregations are easily accessed. 
Therefore, even if most gag are not migrating to 
Florida for spawning, the vulnerability of at least 
part of the stock to winter fishing is of concern. This 
“funneling” of migratory fishes off southeastern 
Florida may increase fishing mortality for many spe- 
cies and requires further investigation. 
In 1993, the SAFMC decided to take no action on 
a proposal to close all fishing for gag during their 
spawning season because insufficient data were 
available on sex ratios and the spatial extent of 
spawning along the southeast coast of the United 
States. The information on sex ratio and size at ma- 
turity presented here indicates that the gag popula- 
tion is overfished. Because there has been a reduc- 
tion in the percentage of males in commercial land- 
ings and a modest decrease in the size at maturity 
for females, the total fishing effort should be lim- 
ited, especially in the removal of males from spawn- 
ing aggregations. Since gag spawn in offshore areas 
from North Carolina to southeast Florida, any regu- 
lation should apply to all southeastern states. How- 
ever, seasonal closure to fishing may not be suffi- 
cient to protect gag from overexploitation because 
males will continue to be lost during the open sea- 
son. Protected areas with no fishing, such as marine 
fishery reserves, may be a possible solution to over- 
fishing of reef fishes along the southeast coast of the 
United States (PDT, 1990). Marine fishery reserves 
can protect population age structure, species diver- 
sity, genetic diversity, and recruitment supply to ex- 
ploited areas (Bohnsack, 1993). Sedberry et al. (1996) 
found that marine reserves in Belize, Central 
America, had a higher diversity of fishes than in ar- 
eas that were not protected. Top predators, such as 
various grouper and snapper species, were more 
abundant and larger in reserves. In addition, popu- 
lations of herbivorous forage species were reduced 
to presumed natural levels in the presence of pro- 
tected predators. Marine reserves in Belize appear 
to have a natural balance of predators and forage 
species in relation to fished areas. There are other 
species in the snapper-grouper complex that are 
showing signs of overfishing, in addition to gag, in- 
cluding red porgy (Harris and McGovern, 1997), ver- 
milion snapper (Zhao and McGovern, 1997; Zhao et 
al., 1997), and black sea bass. 2 In addition, less im- 
portant species in fisheries (e.g. white grunt, gray 
triggerfish) are increasing in abundance. 2 This trend 
is likely to continue unless different management 
regulations are imposed that will protect ecosystems 
and restore a natural equilibrium community. 
Acknowledgments 
We thank personnel associated with cooperative state 
and federal agencies, for their assistance with the 
collection of data, and commercial fishermen for pro- 
viding trip information. In particular, we recognize 
T. Brandt, L. Bishop, M. Burton, D. Codella, C. Den- 
nis, R. Roman, and D. Thiesen of the NMFS, G. 
Rogers and J. Ross of the Georgia Department of 
Natural Resources, and F. Rohde of the North Caro- 
lina Department of Environment, Health, and Natu- 
ral Resources for their efforts. We thank Dr. John 
Grego (University of South Carolina) for helping us 
with the probit procedure. The assistance at sea by 
SCDNR MARMAP personnel and the crew of the RV 
Palmetto is appreciated. We also thank K. Grimball, 
T. Kellison, and other MARMAP personnel for pro- 
cessing gonads that were sent from state and fed- 
eral port samplers. This study was sponsored by the 
National Marine Fisheries Service (MARMAP Contract 
No. 52WCNF6006013PW and Saltonstall-Kennedy 
Grant NA57FD0030), and the South Carolina Depart- 
ment of Natural Resources. 
Literature cited 
Bohnsack, J. A. 
1993 . Marine reserves. Oceanus 36:63-71. 
Carter, J., G. J. Marrow, and V. Pryor. 
1994 . Aspects of the ecology and reproduction of Nassau 
grouper, Epinephelus striatus, off the coast of Belize, Cen- 
tral America. Proc. Gulf Caribb. Fish. Inst. 43:65-111. 
Coleman, F. C., C. C. Koenig, L. A. Collins. 
1996 . Reproductive styles of shallow-water groupers (Pi- 
sces: Serranidae) in the eastern Gulf of Mexico and the 
consequences of fishing spawning aggregations. Environ. 
Biol. Fish. 47:129-141. 
2 1998. MARMAP unpublished data. Marine Resources Re- 
search Institute, South Carolina Department of Natural Re- 
sources, Charleston, SC 29422-2559. 
