American Oyster (left and center) and Pacific Oyster (right)
American Oyster (left and center) and Pacific Oyster (right)

Eastern Oyster (farmed)

Crassostrea virginica

Sometimes known as American Cupped Oyster, American Oyster, Atlantic Oyster, Blue Point Oyster or Cove Oyster.

This species is farm-raised.

Summary

The Chesapeake Bay was once the center of Eastern Oyster production, but overfishing and disease have decimated its oyster population. States bordering the Gulf of Mexico now supply the majority of Eastern Oysters. Eastern Oyster cultivation improves water quality by filtering algae from the water column. The most common method for harvesting Eastern Oysters is dredging, which is harmful to bottom habitat.

Criterion Points
Inherent Operational Risks 2.00
Feed 3.75
Pollution 3.75
Risk to Other Species 3.00
Ecological Effects 2.00
Final Score (average of criteria) 2.90
Color
Final Score Color
2.60 - 4.00
2.20 - 2.59
1.80 - 2.19
1.40 - 1.79
0.00 - 1.39

Last updated December 17, 2007.

Inherent Operational Risks

Core Points (only one selection allowed)

General System Design:

An aquaculture system's design is a good overall proxy measure for the likely effect of the operation on the environment. For example, open systems (e.g., net pens and net cages) are more likely to have pollution, disease, and escape issues than closed systems (e.g., recirculating tanks). With shellfish, which don’t require supplemental food input, the more important question is whether they are harvested on or off of the bottom.

1.00
This species is raised in a high risk system (e.g., net pens; net cages).
2.00
This species is raised in a moderate risk system (e.g., most ponds; raceways; bottom culture of mollusks).

In recent years, oyster beds in the Gulf of Mexico have supplied over 90% of the annual U.S. Eastern Oyster production (NMFS 2006). Before overharvesting and disease ravaged the Chesapeake Bay’s oysters, Maryland and Virginia oyster harvesters supplied the vast majority of Eastern Oysters to the U.S. market (CBPB 2004). In 2004, oyster beds in the Chesapeake Bay only contributed 0.3% of the total harvest of Eastern Oysters in 2004 (NMFS 2006).

In August 2005, Hurricane Katrina caused extensive damage to the Gulf’s oyster beds (CRS 2005). The Louisiana Oyster Task Force, however, predicted a quick recovery. Though they have not released their findings yet, the task force forecast that harvesters would recover 50% of their annual production by the end of 2005 (CRS 2005).

3.00
This species is raised in a low risk system (e.g., re-circulating closed system; suspended culture of mollusks; zero-discharge ponds).

Points of Adjustment (multiple selections allowed)

-0.25
Species is raised at a high stocking density; OR there is a high density of sites in the geographic region, with evidence of environmental impact.

In Louisiana, oyster harvesters spread oyster seed (juvenile oysters that are 5mm – 20mm in length) over submerged mud bottoms at densities that range from about 200,000 to 500,000 oysters per acre (SRAC 2002). Since Eastern Oysters naturally grow in high-density oyster reefs, we chose not to subtract for this factor.

-0.25
Operations do not incorporate best-available, cost-effective technology to reduce environmental impact.
-0.25
There are no effective zoning or permitting practices for siting of facilities.
-0.25
Government programs encourage expansion of high-impact systems.
+0.25
Species is raised at a low stocking density OR there is a low density of sites in the geographic region, which results in minimal impact to the natural ecosystem.
+0.25
Operations incorporate innovative culture methods that limit environmental impacts (e.g., polyculture).
+0.25
There are effective zoning or permitting practices for siting and operation of facilities (e.g., mandatory consideration of hydrographic characteristics; requirements for site rotation).

In the Gulf States, the U.S. Army Corps of Engineers and state public health departments are involved in site and operations permitting for Eastern Oyster farming (SRAC 2001). Other state agencies regulate leasing of underwater land for Eastern Oyster farming. The lease agreements generally stipulate shellfish production quotas (TNC 2002). Land-leasing regulations are especially important for natural Eastern Oyster reefs to prevent over-harvesting (Beck et al. 2004). However enforcement can be expensive and is often poor (Beck et al. 2004). Therefore, we chose to not award points for this factor.

+0.25
Government programs preferentially encourage the expansion of low-impact systems over high impact systems.
2.00
Points for Inherent Operational Risks

Feed

Core Points (only one selection allowed)

Ecological Footprint of Feed:

"Trash" fish, frequently used in developing countries, is an industry term used to refer to whole fish or fish parts fed to farmed fish without being processed into fish meal and fish oil.

Twenty percent was selected as a cut-off because carnivorous species (e.g., salmon; eel; tuna; cobia; etc.) generally consume greater than twenty percent fish products (fishmeal, fish oil, or trash fish), while omnivorous or herbivorous species (e.g., catfish; tilapia; carps; etc.) consume less than twenty percent fish products.

1.00
Typical aquaculture feed includes high levels of fishmeal, fish oil, or "trash" fish (i.e., >20% of the feed; e.g., salmonid feeds).
2.00
Typical aquaculture feed includes moderate levels of fishmeal, fish oil, or "trash" fish (i.e., <20% of the feed; e.g., tilapia and catfish feeds).
3.00
No feed is used (e.g., mollusks and seaweeds) or typical aquaculture feed includes no fishmeal, fish oil, or "trash" fish (e.g., paddlefish; filter-feeding carps).

Raising Eastern Oysters does not require feed (Matthiessen 2001). Oysters filter-feed plankton from the water.

Points of Adjustment (multiple selections allowed)

-0.25
When fish products are used, the major sources score low on the Wild-Caught Fisheries Ranking System.
-0.25
Feed contains greater than 10% of fish products and public or private sectors are not working to reduce fish content in feed.
-0.25
Feed conversion ratio (FCR) is high (i.e., >2.0; e.g., eel).
-0.25
Government policy promotes research, development and commercialization of carnivorous or other highly fishmeal-dependent species.
+0.25
When fish products are used, the major sources score high on the Wild-Caught Fisheries Ranking System; OR the source is innovative and ecologically sound (e.g., fisheries byproducts); OR no feed is used.

Raising Eastern Oysters does not require feed (Matthiessen 2001).

+0.25
Feed contains less than 10% of fish products OR public and private sectors are working to reduce the fish content in feed; OR no feed is used.

Raising Eastern Oysters does not require feed (Matthiessen 2001).

+0.25
Feed conversion ratio (FCR) is low (i.e., <1.3; e.g., salmon); OR no feed is used.

Raising Eastern Oysters does not require feed (Matthiessen 2001).

+0.25
Government policy promotes research, development and commercialization of herbivorous species or other species not highly dependent on fishmeal.
3.75
Points for Feed

Pollution

Core Points (only one selection allowed)

Typical effluent treatment procedures:

1.00
Effluent is not treated before discharge (e.g., salmon net pens).
2.00
Effluent is partially treated before discharge (e.g., infrequently discharged effluent from catfish ponds).
3.00
Effluent is substantially treated before discharge (e.g., recirculating shrimp systems; settling ponds; reconstructed wetlands); OR treatment is not necessary because supplemental feed is not used (e.g., molluscs or seaweeds).

Treatment of effluent is not necessary because raising Eastern Oysters does not require supplemental feed (Matthiessen 2001).

Points of Adjustment (multiple selections allowed)

-0.25
Operations have demonstrated negative impacts on water quality or sediment/benthic characteristics (e.g., elevated nutrient levels; algal blooms; altered benthic communities).
-0.25
Pollutants (e.g., pesticides; parasiticides; antibiotics; plastic; nets; dead fish) are frequently discharged into the environment or otherwise not appropriately discarded.
-0.25
Effluent regulations do not exist, are lax, or are poorly enforced, which allows for degradation of the aquatic environment.
-0.25
Available technologies and practices to reduce or recycle waste (e.g., feed sensors; low-pollution feeds) are not used.
+0.25
Operations generally improve water quality or sediment/benthic characteristics (e.g., oyster farms).

Oysters reefs provide important habitat to other marine organisms, including juvenile fish (Lenihan and Peterson 2004). Also, the filter-feeding activity of oysters improves water quality by removing suspended particles, excess nutrients, and phytoplankton. Oyster filter-feeding activity can enhance light penetration in the water column, stimulating growth of aquatic vegetation on the seafloor, which in turn increases oxygen levels in bottom waters (ECSGA 2002). The removal of nutrients like nitrogen helps prevent excess build up of nutrients in the water, which can lead to harmful phytoplankton blooms (U.S. EPA 2005). At the height of the Chesapeake Bay’s Eastern Oyster population, the oysters filtered the bay’s volume every 3 to 4 days (CBP 2005).

+0.25
Chemicals (e.g., pesticides; parasiticides; antibiotics) are rarely or never used.
+0.25
Robust water quality regulations exist (e.g., permits required; discharge caps; strong enforcement), and regular monitoring occurs.

Eastern Oyster farms do not produce effluent. However, the National Shellfish Sanitation Program (NSSP) must monitor water quality in areas of Eastern Oyster cultivation, because oysters can accumulate toxins harmful to humans from polluted water or water containing high phytoplankton concentrations (e.g., red tides). State and local agencies are responsible for issuing permits, overseeing leasing of beds, exercising control over uncertified areas, monitoring water quality and classifying areas in accordance with NSSP guidelines (Kraeuter and Castagna 2001).

+0.25
Innovative methods and practices to reduce or recycle wastes are used (e.g., integrated systems; effluent and solid wastes used as terrestrial fertilizer); OR innovative methods and practices are not needed because raising this species does not create waste.

Eastern Oysters are filter feeders and raising them does not create effluent (Matthiessen 2001). However, in high concentrations, psuedofeces and feces produced by Eastern Oysters can have harmful environmental effects by creating areas of no or low oxygen levels. Bacteria quickly consume psuedofeces and feces, and in doing so, they take up oxygen and respire carbon dioxide. High levels of bacterial respiration causes anoxic (no oxygen) conditions, which can reduce the diversity of seafloor organisms in the affected area (DFO 2006). Generally, however, this problem is associated with growing oysters in suspended rafts not with cultivated oyster reefs. Since seeded oyster-reef operations account for the majority of U.S. Eastern Oyster production (NMFS 2006), we chose to award points here.

3.75
Points for Pollution

Risk to Other Species

Core Points (only one selection allowed)

Frequency and Impact of Escapes:

1.00
Farmed species regularly or intermittently escape into the wild AND escapes are non-native to the area or otherwise pose a risk to native populations or ecosystems (e.g., most non-native fish raised in outdoor facilities).
2.00
Escape frequency is not known OR farmed species is native to the area where it is raised and poses minimal risk to native populations or ecosystems (e.g., channel catfish in the US; most native mollusks).

Eastern Oysters are native in the Western Atlantic Ocean from Labrador, Canada to Mexico, including the Gulf of Mexico. Historically, Chesapeake Bay harvesters supplied the majority of the U.S. Eastern Oysters, but production is now heaviest in the Gulf of Mexico. "Escape" from Eastern Oyster farming operations via broadcast spawning (release of eggs and sperm into the water) likely helps repopulate natural oyster beds that have been impacted by overfishing, hurricanes, and/or disease (Matthiessen 2001).

Though some Eastern Oyster production occurs on the West Coast of the U.S., West Coast operations contribute very little to the U.S. market compared to the Gulf of Mexico oyster beds. Therefore, we chose to award 2.00 points for this factor.

3.00
Farmed species never (or virtually never) escape to the wild (e.g., species is raised in bio-secure facilities).

Points of Adjustment (multiple selections allowed)

-0.25
This farmed species has been known to survive in the surrounding ecosystem if it escapes; OR would likely survive given its physiological requirements.
-0.25
This farmed species is known or is likely to compete with wild species for food or habitat if it escapes; OR this species is known or is likely to compromise the genetic integrity of the wild species (e.g., through spawning disruption, genetic introgression or establishment of feral stocks) if it escapes.
-0.25
This farmed species is known or is likely to amplify and transmit disease or parasites to wild populations (e.g., infectious salmon anemia or sea lice infestations) if it escapes.

Diseases such as Dermo (Perkinsus marinus) and MSX (Haplosporidium nelsoni) have greatly impacted oyster populations on the U.S. East Coast, particularly in the Chesapeake Bay. In the Chesapeake Bay the diseases continue to prevent the recovery of the depleted oyster population (Paynter 2005). The diseases spread between farmed and wild Eastern Oysters, but temperature and salinity play a more important role determining the spread of the diseases than farming operations (Brown et al. 2005a). Therefore, we chose not to deduct points here.

-0.25
Regulatory authorities are not adequately addressing the risks of escape or spread of disease associated with farming this species.
+0.25
This farmed species has not been known to survive in the surrounding ecosystem if it escapes; OR would not likely survive given its physiological requirements; OR farmed species is a native mollusc.

The majority of Eastern Oysters in the U.S. market are from the Gulf of Mexico, which is within the species’ native range (Matthiessen 2001).

+0.25
Operations employ management protocols and techniques to limit the ecological impacts of escaped farmed fish (e.g., triploidy; sterilization); OR it’s unlikely that escaped individuals will either compete with wild species for resources, or compromise the genetic integrity of wild species.

It is unlikely that oysters that have “escaped” would compromise the genetic integrity of wild populations. The major American producers along the Gulf Coast seed their beds with spat (larval oysters) from wild populations (Coleman 2003).

+0.25
Operations employ effective disease and parasite management protocols (e.g., fallowing of pens; retaining water when disease outbreak occurs); OR incidence of disease or risk of retransmitting disease is low.

The transfer of Eastern Oysters from high to low salinity water can help reduce the incidence and prevalence of MSX (H. nelsoni) and DERMO (P. marinus) infections (Paynter 2005).

Programs such as Dermowatch, an Internet-based monitoring and modeling program, aid growers and resource managers in dealing with Dermo outbreaks. The program forecasts the intensity of infections based on environmental parameters. Dermo is most virulent in high-salinity waters and can cause high levels of Eastern Oyster mortalities. Growers use Dermowatch’s forecasts to determine when they should move their oysters to areas of lower salinity, which helps them prevent high mortality. Growers also use the forecasts to predict when they should harvest market-sized oysters before they die from the disease (Gulf Oyster Watch 2005).

MSX does not infect Eastern Oysters in the Gulf of Mexico, only Eastern Oysters along the East Coast. Chesapeake Bay harvesters deal with the disease by moving oysters to lower-salinity water and using selectively bred disease-resistant Eastern Oysters (Bower 2003). Oyster managers and harvesters also use mathematical models to simulate the possible spread and incidence of MSX.

+0.25
Regulatory authorities are addressing the risks of escape and spread of disease associated with farming this species.

The National Shellfish Sanitation Program (NSSP) oversees the transfer of live shellfish (spat and full-grown individuals) between states to help prevent disease problems (Kraeuter and Castagna 2001).

3.00
Points for Risk to Other Species

Ecological Effects

Core Points (only one selection allowed)

Ecological sensitivity of site used for operations:

1.00
Operations are generally located in areas of high ecological sensitivity (e.g., coastal wetlands; mangroves).
2.00
Operations are generally located in areas of moderate ecological sensitivity (e.g., coastal and nearshore waters; rocky intertidal or subtidal zones; river or stream shorelines).

Eastern Oysters live in intertidal and subtidal zones of bays, inlets, and sounds and behind barrier islands (Matthiessen 2001). Growers often cultivate oyster beds in waters that are uncovered at low tide, so that they are easily accessible for harvesting (SRAC 2001).

3.00
Operations are generally located in areas of low ecological sensitivity (e.g., land that is less susceptible to degradation such as land formerly used for agriculture or land previously developed).

Points of Adjustment (multiple selections allowed)

-0.25
Farming this species causes substantial damage to surrounding habitat, ecosystem or other resources (e.g., groundwater depletion; stream diversion; saltwater intrusion; soil salinization; loss of habitat for juvenile fish; loss of flood control; dredging hard bottoms; etc.).

Eastern Oyster harvesters from the Gulf States to the Chesapeake Bay States mainly use dredges and tongs to harvest oysters. Dredging is most common in the Gulf States operations, and, in Florida, tonging methods dominate (NMFS 2006).

Dredging destroys bottom habitat, decreases biodiversity in the area, and captures and kills significant numbers of non-targeted animals (Morgan and Chuenpagdee 2003). There are two ways to dredge for oysters: power dredging and sail dredging. The difference refers to how the dredge is being pulled across the seafloor: power dredging is by motorboat and sail dredging, by sailboat. In the Gulf of Mexico, harvesters use power dredging exclusively. In the Chesapeake Bay, however, where Eastern Oyster beds remain severely depleted, there is considerable debate over the use of "power" dredging in oyster beds in Chesapeake Bay. Power dredging enables harvesters to dredge more oyster beds, which can cause overharvesting and depletion; while sail-powered boats move slower and dredge less area (Morgan and Chuenpagdee 2003).

Tonging methods cause significantly less damage to the bottom habitat than dredges (Lenihan and Peterson 2004). Tongs have very long handles of either wood or metal with a rake-like end for each handle. The rake-like ends scrape the bottom to capture oysters, which harvesters then bring back onboard to release the oysters on their boat decks.

Farming Eastern Oysters enhances structural habitat diversity and improves water quality (ECSGA 2002). However, oyster dredging damages habitat (Lenihan and Peterson 2004). Since the majority of Eastern Oysters in the U.S. market are harvested with dredges (SRAC 2002; NMFS 2006), we chose to deduct here.

-0.25
Harmful or lethal predator deterrents are used (e.g., bird/seal shootings; acoustic deterrent devices); OR operation otherwise harms wildlife (e.g., dolphin/seal entanglement; disrupting migration routes; bird/animal shooting).

Harmful or lethal predator deterrents are rarely used in Gulf of Mexico oyster operations. In the Chesapeake Bay, however, chemical predator deterrents, anti-predator culturing methods, and the destruction of predators are common. Some harvesters use suction dredges to remove Oyster Drill snails and mops to remove sea stars from oyster beds (Matthiessen 2001). The mops entangle the sea stars, which harvesters then remove from their oyster beds.

Suspended raft culturing is another way of growing Eastern Oysters. This method elevates oysters in the water above their benthic predators (Matthiessen 2001; LSGC 2003) by growing them on floating wooden rafts. Therefore harvesters using suspended rafts to grow oysters do not need to use predator deterrents (Matthiessen 2001; LSGC 2003).

We did not deduct points for this factor because the majority of Eastern Oysters in the U.S. market are from seeded oyster beds in the Gulf of Mexico, where predator deterrents are rarely used (SRAC 2002; NMFS 2006).

-0.25
If seed is collected from wild sources, the intensity of collection is high enough to result in depletion of brood stock, wild juveniles, or associated non-target organisms (e.g., collection of postlarvae shrimp).
-0.25
Government policy encourages aquaculture operations to locate or expand in areas of high ecological sensitivity.
+0.25
Operations enhance habitat structure or function (e.g., constructed wetlands).
+0.25
Predator deterrents are not used OR predator deterrents are used but are not harmful or lethal (e.g., predator exclusion nets), AND operation does not otherwise harm wildlife.
+0.25
Seed comes predominantly from hatcheries or on-farm sites (e.g., seed for trout); OR if seed is collected from the wild, it does not deplete brood stock, wild juveniles, or associated non-target organisms (e.g., collection of oyster or mussel spat).

Eastern Oyster harvesters collect seed primarily from wild oyster stocks. Using seed from hatcheries is less common due to expense. Collection of seed does not likely deplete the wild brood stock. (Matthiessen 2001).

+0.25
Government policy encourages the growth of aquaculture operations in areas of low ecological sensitivity; OR protects sensitive habitats from aquaculture operations (e.g., prohibitions on cutting mangroves).
2.00
Points for Ecological Effects

References

Beck, M.W., T.D. Marsh, S.E. Reisewitz, and M.L. Bortman. 2004. New Tools for Marine Conservation: the Leasing and Ownership of Submerged Lands. Conservation Biology. Vol 18, No. 5: 1214-1223.

Bower, S.M. 2003. Synopsis of Infectious Diseases and Parasites of Commercially Exploited Shellfish: Haplosporidium nelsoni (MSX) of Oysters. Accessed 2 June 2006. Available at: http://www-sci.pac.dfo-mpo.gc.ca/shelldis/pages/hapneloy_e.htm

Brown, B.L., A.J. Butt, D. Meritt, and K.T. Paynter. 2005a. Evaluation of resistance to Dermo in eastern oyster strains tested in Chesapeake Bay. Aquaculture Research Vol. 36: 1544-1554. Available at: http://www.life.umd.edu/biology/paynterlab/labpub/ARE_1377.pdf

Brown, J.J., R. Hildreth, and S.E. Ford. 2005b. Letters: When the world is not your oyster. Science Vol. 309: 244. Available at: http://www.sciencemag.org/cgi/reprint/309/5732/244b.pdf

Chesapeake Bay Program (CBPa). Eastern Oyster. Accessed 10 May 2006. Available at: http://www.chesapeakebay.net/american_oyster.htm

Chesapeake Bay Program (CBPb).Virginia Oyster Aquaculture Program FY 2004 Year End Summary. Accessed 14 July 2006. Available at http://www.cbf.org/site/DocServer/oyster_garden_ar04.pdf?docID=2866

Coleman, E. 2003. Louisiana Sea Grant College Program. The Gulf Oyster Industry: Seizing a Better Future. Available at: http://www.nsgo.seagrant.org/research/oysterdisease/index.html

Congressional Research Service Report for Congress (CRS). 2005. Hurricanes Katrina and Rita: Fishing and Aquaculture Industries- Damage and Recovery. Order Code RS22241. Available online at: http://fpc.state.gov/documents/organization/57873.pdf

Department of Fisheries and Oceans (DFO). 2006. Assessing Habitat Risks Associated with Bivalve Aquaculture in the Marine Environment. Department of Fisheries and Oceans Can. Sci. Advis. Sec. Sci. Advis. Rep. 2006/005. Available online at:

http://www.dfo-mpo.gc.ca/csas/Csas/status/2006/SAR-AS2006_005_E.pdf

East Coast Shellfish Growers Association (ECSGA). 2002. East Coast Shellfish Growers Association. Accessed on 18 January 2006. Available at: http://www.ecsga.org/pages/BenefitsBrochure.pdf

Gulf Oyster Watch. 2005. Texas Sea Grant. Sea Grant Gulf Oyster Industry Program. Dermowatch. Accessed on 25 May 2006. Available online at: http://www.gulfoysterwatch.org/

King, J.L. and K. McGraw. 2004. Status of U.S. Oyster Resource. Habitat Connections. Oyster Restoration Series. Vol. 5: No. 1: 1-4. National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Office of Habitat Conservation. Silver Spring, MD. Available at: http://noaa.chesapeakebay.net/docs/HabitatConnectVol5no1NOV2004.pdf

Kraeuter, J.N. and M. Castagna, Eds. 2001. Biology of the Hard Clam. Developments in Aquaculture and Fisheries Science – 31. Elsevier. Amsterdam, The Netherlands.

Lenihan, H.S. and C.H. Peterson. 2004. Conserving oyster reef habitat by switching from dredging and tonging to diver-harvesting. Fishery Bulletin. Vol. 102: 298-305.

Louisiana Sea Grant College (LSGC). 2003. Coleman, Elizabeth. The Gulf Oyster Industry: Seizing A Better Future. Available at: http://aquanic.org/beginer/shellfish/Final%20Oyster%20Pub.pdf

Matthiessen, G. C. 2001. Oyster Culture. Fishing News Books, Blackwell Science Ltd. Great Britain.

Morgan, L.E. and R. Chuenpagdee. 2003. Shifting gears: addressing the collateral impacts of fishing methods in U.S. waters. Pew science series on conservation and the environment.

National Marine Fisheries Service (NMFS). 2006. Fisheries Statistics and Economics: Commercial Fisheries Information. Available at: http://www.st.nmfs.gov/st1/ and http://www.st.nmfs.gov/st1/commercial/landings/annual_landings.html

Paynter, K. T. 2005. Harvest Reserve Monitoring Report 2005. Accessed on 10 May 2006. Available at: http://www.life.umd.edu/biology/paynterlab/labpub/index.html

Southern Regional Aquaculture Center (SRAC). 2002. Supan, John. Extensive Culture of Crassostrea virginica in the Gulf of Mexico Region. Publication No. 4300. Available online at: http://aquanic.org/publicat/usda_rac/efs/srac/4300fs.pdf

SRAC. 2001. Wallace, Richard K. Cultivating the Eastern Oyster, Crassostrea virginica. Publication No. 432. Available online at: http://aquanic.org/publicat/usda_rac/efs/srac/432fs.pdf

The Nature Conservancy (TNC). 2002. Marsh, T.D., Beck, M.W. and S.E. Reisewitz. 2002. Leasing and Restoration of Submerged Lands: Strategies for Community-based, Watershed-scale Conservation. The Nature Conservancy, Arlington, VA. Available online at : http://www.nature.org/files/lease_sub_lands.pdf

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Wallace, R.K. 2006. Personal Communication. Auburn University Marine Extension and Research Center (AUMERC) / Alabama Sea Grant.

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