Positive Relationship between Freshwater Inflow and Oyster Abundance in Galveston Bay, Texas

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Estuaries and Coasts (2009) 32:206–212
DOI 10.1007/s12237-008-9078-z

 TECHNICAL COMMUNICATION

Positive Relationship between Freshwater Inflow
and Oyster Abundance in Galveston Bay, Texas
David Buzan & Wen Lee & Jan Culbertson &
Nathan Kuhn & Lance Robinson

Received: 29 June 2008 / Accepted: 21 July 2008 / Published online: 19 August 2008
# Coastal and Estuarine Research Federation 2008

Abstract Analysis of fisheries-independent data for                    estuary in order to increase oyster harvest. We have the
Galveston Bay, Texas, USA, since 1985 shows eastern                    opposite concern that oysters will be harmed by projects
oysters (Crassostrea virginica) frequently demonstrate                 that reduce flow, increase salinity, and increase the
increased abundance of market-sized oysters 1 to 2 years               duration of higher salinity periods in a basin with
after years with increased freshwater inflow and decreased             increasing demand for limited freshwater. Turner’s study
salinity. These analyses are compared to Turner’s (Estuaries           and our analysis reflect different aspects of the complex,
and Coasts 29:345–352, 2006) study using 3-year running                important relationships between freshwater inflow, salin-
averages of oyster commercial harvest since 1950 in                    ity, and oysters.
Galveston Bay. Turner’s results indicated an inverse
relationship between freshwater inflow and commercial                  Keywords Freshwater inflow . Oyster productivity .
harvest with low harvest during years of high inflow and               Oyster landings . Galveston Bay
increased harvest during low flow years. Oyster popula-
tions may experience mass mortalities during extended
periods of high inflow when low salinities are sustained.              Introduction
Conversely, oyster populations may be decimated during
prolonged episodes of low flow when conditions favor                   The eastern oyster, Crassostrea virginica, contributes
oyster predators, parasites, and diseases with higher                  ecologically and economically to coastal ecosystems along
salinity optima. Turner’s (Estuaries and Coasts 29:345–                United States’ Atlantic and Gulf of Mexico coasts.
352, 2006) analysis was motivated by a proposed project                Commercial oyster production in Texas, second to Louisiana,
in a basin with abundant freshwater where the goal of the              comprised 20% of the nation’s harvest from 2000 to 2005
project was to substantially increase freshwater flow to the           (NOAA 2007). Oyster reefs filter solids from the water
                                                                       column, influence hydrological patterns, and provide habitat
                                                                       and refugia for a variety of species (Coen et al. 1999; Meyer
D. Buzan (*)                                                           and Townsend 2000; Rodney and Paynter 2006; Zimmerman
PBS&J,
                                                                       et al. 1989).
6504 Bridge Point Parkway, Suite 200,
Austin, TX 78730, USA                                                     Estuaries with substantial freshwater inflows like Ches-
e-mail: dlbuzan@pbsj.com                                               apeake Bay on the Atlantic coast, Apalachicola Bay in
                                                                       Florida, Mobile Bay in Alabama, the Louisiana coastal
W. Lee : N. Kuhn
                                                                       zone, and Galveston Bay in Texas support relatively large
Texas Parks and Wildlife Department,
4200 Smith School Road,                                                populations of oysters. In Texas, bays with productive
Austin, TX 78744, USA                                                  shellfish industries tend to have relatively high rates of
                                                                       freshwater inflow (Montagna and Kalke 1995). Galveston
J. Culbertson : L. Robinson
                                                                       Bay receives the second-highest freshwater inflow of any
Texas Parks and Wildlife Department, Dickinson Marine Lab,
1502 FM 517 East,                                                      Texas estuary with an annual average 11.4 million acre-ft/
Dickinson, TX 77539, USA                                               year (TWDB 2007a) and produces most oysters commer-
Estuaries and Coasts (2009) 32:206–212                                                                                    207

cially harvested in Texas. Galveston Bay reefs grow              patterns in the relationships between freshwater inflow,
primarily in the middle of the bay and cover more than           salinity, and oyster productivity in Galveston Bay. These
10,000 ha of bay bottom (Powell et al. 2003). Monthly            broad annual patterns indicate possible value in future,
average salinities in Galveston Bay range from 12 to 19 psu      more intensive analysis of seasonal salinities or inflows on
(Ray 1987; Quast et al. 1988; TPWD 2006a).                       one or a few reefs in Galveston Bay. Our analysis also
   Turner (2006) compares oyster harvest to river inflow (as     utilizes a shorter period of record, 1985 to 2004, than
a proxy for salinity) for five Gulf of Mexico estuaries          Turner (2006) and oyster abundance data obtained from a
including Galveston Bay. He suggests Gulf of Mexico              fisheries-independent monitoring program rather than
estuaries with salinity above optimum levels will experi-        oyster commercial harvest.
ence increased oyster harvest as salinity decreases towards
an optimum. He further suggests a continued decline in
salinity past some optimal point will lower oyster produc-       Materials and Methods
tivity. Turner (2006) evaluated a 54-year record of oyster
harvest in Galveston Bay. Comparing peaks and troughs in         Oysters are monitored by Texas Parks and Wildlife
oyster landings to freshwater inflows, Turner (2006)             Department’s (TPWD) Coastal Fisheries Resource Moni-
concludes “…the annual variability in oyster landings of         toring Program according to the “Marine Resource
the Gulf of Mexico estuaries are, in general, inversely          Monitoring Operations Manual” (TPWD 2002). Samples
related to freshwater inflow.”                                   are collected with dredges (Louisiana style nine-tooth,
   Freshwater inflow is a critical factor influencing oyster     46 cm wide, 25 cm tall with a 36-cm-deep bag) on reefs
abundance. Lengthy periods of low freshwater inflow allow        where oysters form the bottom and the reefs extend at least
salinities to rise and oyster mortality from predation and       0.2 m above the bottom for a continuous distance of 91.4 m
parasitism to increase (Ray 1987). Floods may suppress           long by 0.5 m wide in any direction. The dredge is towed in
salinities long enough to cause mass mortalities of oysters      a straight line for 30 s. All live oysters are counted and 19
(Wilber 1992). Powell et al. (2003) describe how changes         live individuals are chosen at random and measured.
in location of freshwater inflow, inflow amounts, among          Salinity is measured with a refractometer or digital
other factors interact to determine future oyster abundance.     conductivity/salinity meter 0.3 m above the bottom sub-
   Turner (2006) and Wilber (1992) both studied the              strate at the dredge starting point.
relationship between harvest and freshwater inflow. Wilber          Catch-per-unit-effort (CPUE) of “market-sized” oysters
(1992) calculated oyster abundance by dividing the pounds        is calculated by multiplying the number of live oysters
of oysters harvested by the number of registered oystermen       ≥76 mm dredged in 30 s by 120 to convert dredge results to
for each year and eliminating harvest years which were           CPUE measured per hours. CPUE (market-sized oysters per
confounded by hurricanes, drought, and oyster-reef resto-        hour) is used to indicate market-sized oyster relative
ration. Turner (2006) utilized a 3-year moving average of        abundance. The annual value discussed in this report is
oyster harvest to deal with the highly variable harvest data     the average CPUE for all TPWD samples collected within
and influence from occasional rare events.                       the bay in the same calendar year.
   Although Turner’s conclusions clearly focus on the               Freshwater inflow for Galveston Bay includes all U.S.
relationship between oyster harvest and freshwater inflow,       Geological Survey (USGS) gaged flows to the estuary plus
we are concerned his conclusions will be interpreted by          modeled flows from ungaged coastal watersheds dis-
some to indicate freshwater inflow events are generally          charging to the estuary over the period from 1941 to 2005
detrimental to oyster health. This paper describes a positive,   (TWDB 2007b). The Texas Rainfall Runoff model uses
1- to 2-year lagged relationship between freshwater inflow       rainfall, flow diversion reports, and reported wastewater
and oyster abundance as distinguished from oyster harvest        discharges to model flow from ungaged watersheds. Daily
in Galveston Bay. Wilber (1992) showed 2- to 3-year time         rainfall data are obtained from the National Weather
lags between low flow events and subsequent poor oyster          Service. Return flow and diversion data are from the Texas
harvest and a positive relationship between fall inflows and     Commission on Environmental Quality. Turner’s (2006)
oyster harvest 2 years later. Hofstetter (1988) described a      analysis used Trinity River inflow measured at a USGS
lagged relationship between inflow and oyster harvest in         gage upstream of the bay. Annual trends in Trinity River
Galveston Bay for the 1980 year class following severe           flow into Galveston Bay are generally comparable to
flooding in 1979 which supported record harvests in the          annual trends in freshwater inflow. Approximately 69% of
1982–1983 season. Most recently, Bergquist et al. (2006)         the freshwater inflow to the bay has been gaged flow over
demonstrated positive relationships between four indicators      the period from 1941 to 2005.
of oyster community health and changes in salinity 1 to             This report focuses on changes in oyster abundance
2 years earlier. Our analysis simply compares broad annual       when compared to freshwater inflow and salinity 1 to 2 years
208                                                                                       Estuaries and Coasts (2009) 32:206–212

earlier. Increases and decreases in oyster abundance are         2 years after years of increased freshwater inflow and
utilized, not for their magnitude but for the direction of the   experiences lows 1 to 3 years after declines in freshwater
response they indicate to changing freshwater inflow and         inflow. Three years, 1988, 2000, and 2001, illustrated a
salinity conditions.                                             pattern similar to that recognized by Turner (2006) where a
                                                                 peak in inflow or oyster productivity was matched by a low
Study Site                                                       in the other variable in the same year. As mentioned earlier,
                                                                 each of these years was also preceded by an increase or
Galveston Bay is located along the upper Texas coast             decrease in inflow 1 to 2 years earlier.
(Fig. 1) in the northwestern Gulf of Mexico. The Trinity             When Turner’s analytical approach is applied to oyster
River contributes 54% of the annual average flow to the          CPUE and total annual freshwater inflow to the bay, it
bay while the San Jacinto River/Buffalo Bayou watersheds         illustrates increasing oyster relative abundance from 1989
contribute 38% of the total annual average inflow to the bay     until 1993 which is preceded by increasing freshwater
(TWDB 2007b). Peak flows to the estuary typically occur          inflow from 1988 until 1992 (Fig. 3). The high oyster
in May and June and low inflows occur August through             CPUE in 1999 follows an increasing trend from 1996
October.                                                         which was paralleled by increasing to stable inflow from
                                                                 1995 to 1998. Similarly, as inflow decreased over the
                                                                 period from 1992 to 1995, it was followed by decreasing
Results                                                          oyster CPUE from 1993 to 1996.

Oyster Abundance and Freshwater Inflow                           Oyster Abundance and Salinity

Freshwater inflow to Galveston Bay increased in 1986,            Increases in oyster CPUE in 1988, 1993, 2000, and 2002
1992, 1998, and 2001 and upward trends in oyster CPUE            were preceded by lows in salinity 2 or 3 years earlier, in
occurred in 1988, 1993, 2000, and 2002, 1 to 2 years             1986, 1991, 1997, and 2001 (Fig. 4).
afterwards (Fig. 2).                                                Declines in market-sized oyster abundance occurred
   Conversely, lows in oyster CPUE occurred in 1987,             from 1988 to 1990, a period when salinity was also
1990, 1996, 2001, and 2004 following lows in freshwater          declining from year to year. CPUE rose from 1990 to
inflow that occurred in 1988, 1996, 2000, and 2003.              1993 and 1994 when salinities remained relatively low and
Annual freshwater inflow and oyster CPUE declined for            stable. Slightly increasing salinity from 1993 to a sharp
three consecutive years prior to the 1996 low in oyster          peak in 1996 occurred at the same time oyster CPUE was
abundance. Oyster abundance appears to increase 1 to             declining. From 1997 to 2004, salinity and oyster CPUE

Fig. 1 Galveston Bay, Texas
with major oyster reefs (shaded
areas in bay) illustrated (Texas
General Land Office 2006)
Estuaries and Coasts (2009) 32:206–212                                                                                  209

Fig. 2 Galveston Bay annual
average oyster CPUE compared
to annual total freshwater
inflow to the estuary

exhibited similar patterns, both increasing from 1997 to        genic factors like oyster harvesting techniques and regu-
2000 and dropping in 2001.                                      lations have varied relatively little since the early 1980s.
    Oyster relative abundance is not significantly related      Texas Department of State Health Services (TDSHS)
(rs =−0.418, p=0.0823, n=18) to annual average salinity         regulations affect landings when suspending oyster harvest
2 years earlier (Fig. 5a). The analysis suggests the            to protect human health from pathogens and red tide toxins.
relationship between salinity and oyster CPUE may be            The current system for closing oyster harvest areas has been
negative. Market-sized oyster abundance and annual fresh-       in place since the early 1980s (Kirk Wiles, personal com-
water inflow to the bay 2 years earlier exhibit a significant   munication). Harvest areas are frequently closed following
(rs =0.541, p=0.0137, n=20) positive relationship (Fig. 5b).    rainfall events because of elevated bacterial levels in the
                                                                water. Consequently, oyster harvest tends to decline as the
                                                                number of harvest area closure days increase.
Discussion                                                          TPWD implemented a probability-based, fishery-
                                                                independent monitoring program for oysters in October
We analyzed the period from 1985 through 2004 because           1984 in Galveston Bay. This program combined with
fishery-independent monitoring of oysters and anthropo-         improvements in data management creates a relatively

Fig. 3 Comparison of a 3-year
moving average of annual aver-
age oyster CPUE to a 3-year
moving average of annual total
freshwater inflow. The values
have been normalized by divid-
ing the 3-year moving average
of annual average total freshwa-
ter inflow and CPUE by the
annual average freshwater
inflow for the period from 1985
to 2005 and the annual average
CPUE for the period from 1985
to 2002, respectively
210                                                                                            Estuaries and Coasts (2009) 32:206–212

Fig. 4 Galveston Bay annual
average oyster CPUE compared
to annual average salinity
in the estuary

comparable, objective data set from 1985 to 2004 for                 corresponded with increases in surface water inflow 2 years
analysis of relationships between oyster abundance,                  earlier while two other highs were preceded by increases
salinity, and freshwater inflow (TPWD 2006a).                        in surface flow a year earlier. Hofstetter (1977) noted
   Figure 2 illustrates four periods of increase in Galveston        Galveston Bay spat were usually abundant in years of
Bay oyster relative abundance. Two of the periods of                 above-average freshwater inflow. He stated that oyster
increase, including the year of greatest CPUE in 2000,               mortality from flooding was more than offset by increased
                                                                     spat set. Low salinities enhanced survival in central
                                                                     Galveston Bay, the area historically with the greatest amount
                                                                     of oyster substrate (Hofstetter 1977; TPWD 2006b).
                                                                        Floods of sufficient magnitude may reduce oyster
                                                                     harvest during the years in which they occur by both
                                                                     killing oysters in parts of the bay and increasing the amount
                                                                     of time the bay is closed to harvest. Conversely, the same
                                                                     floods ensure long-term survival of oyster populations by
                                                                     reducing oyster predators and parasites such as the oyster
                                                                     drill, Stramonita haemastoma, and Perkinsus marinus.
                                                                     Floods that kill oysters may also aid in the long-term
                                                                     control of Perkinsus epizootics by periodically, temporarily
                                                                     eliminating the oyster host (Ray 1987; Hofstetter 1977;
                                                                     Powell et al. 2003). Analysis of P. marinus in oysters
                                                                     showed infection increased with increasing salinity (TPWD
                                                                     2006b). Oysters colonize formerly flooded areas, particu-
                                                                     larly dead oyster shell, beginning a new cycle of growth
                                                                     with reduced numbers of predators and parasites. Fresh
                                                                     dead oysters release a pheromone from tissues that attracts
                                                                     larval oysters resulting in increased spat set on the inside of
                                                                     these shells and increased spat survival (Keck et al. 1971).
                                                                        During periods of low rainfall, the number of days oyster
                                                                     harvest areas are closed is reduced and harvest may
                                                                     increase as a result of increased oystering days. On the
                                                                     other hand, as salinity increases during periods of drought,
                                                                     oyster reefs may become heavily infested with predators
                                                                     and parasites resulting in increased oyster mortality.
Fig. 5 Correlation a between annual average oyster CPUE and lagged
                                                                        Pierce and Conover (1954) suggested oysters in systems
(2 years) annual mean salinity (psu) and b between annual average
oyster CPUE and lagged (2 years) freshwater inflow (million acre-    with fluctuating salinities grew better than those under
feet) to Galveston Bay 1985 to 2004                                  relatively constant conditions. La Peyre et al. (2003) found
Estuaries and Coasts (2009) 32:206–212                                                                                           211

numerous freshwater releases to the Caloosahatchee River       are a key factor to the health and survival of oyster
kept P. marinus infection intensities low, resulting in low    populations.
prevalence of infection, low oyster mortality, and good
growth. They speculated that repetitive and well-timed         Acknowledgements We thank Dr. Sammy Ray with Texas A & M
                                                               University-Galveston for his critical review of this report. This
freshet events can maintain P. marinus infections at non-      analysis was supported by many of the coastal ecologists at Texas
lethal levels in oyster populations. Bergquist et al. (2006)   Parks and Wildlife Department including Brenda Bowling, Fernando
noted disappearance of reefs in the lower intertidal reaches   Martinez-Andrade, Page Campbell, and Lynne Hamlin. Thanks to
of the Suwannee estuary during a prolonged dry spell while     Kirk Wiles and Gary Heideman with the Texas Department of State
                                                               Health Services for the support. Finally, we recognize the inspired
new reefs appeared higher in the estuary closer to             efforts of Robert Hofstetter who passed away in the spring of 2007
freshwater sources. Modeling of total oyster abundance in      and whose 30 years of study of Galveston Bay oysters will continue to
Galveston Bay and possible changes in hydrology predicted      contribute to our understanding of oyster and estuarine ecology far
lower adult oyster abundance resulting from P. marinus         into the future.
infections associated with lowered freshwater inflows and
increased salinities (Powell et al. 2003). Hofstetter (1977)
advises that for Galveston Bay, we should not conclude that
                                                               References
reduced freshwater inflow “…will necessarily be beneficial
to oysters or other estuarine species.”
                                                               Bergquist, D.C., J.A. Hale, P. Baker, and S.M. Baker. 2006.
   The State Water Plan for Texas (TWDB 2006) concludes             Development of ecosystem indicators for the Suwannee River
Texas will not have enough water to meet future demand              estuary: Oyster reef habitat quality along a salinity gradient.
during drought unless new water supply projects or                  Estuaries and Coasts 29: 353–360.
management strategies are implemented. Turner’s (2006)         Coen, L.D., M.W. Luckenbach, and D.L. Breitburg. 1999. The role of
                                                                    oyster reefs as essential fish habitat: a review of current
conclusion that higher oyster harvests are associated with          knowledge and some new perspectives. American Fisheries
higher average salinities in Gulf of Mexico estuaries may           Society Symposium 22: 438–454.
be misinterpreted as valid rationale to reduce freshwater      Hofstetter, R.P. 1977. Trends in population levels of the American
inflows to other estuaries. Wilber (1992), after review of a        oyster Crassostrea virginica Gmelin on public reefs in Galveston
                                                                    Bay, Texas. Texas Parks and Wildlife Department Coastal
number of studies, concluded that relationships between             Fisheries Division Technical Series No. 24, 90 pages.
flow and oyster production vary among different estuarine      Hofstetter, R.P. 1988. Trends in the Galveston Bay oyster fishery,
ecosystems. Water managers may focus on the inverse                 1979–1984. Texas Parks and Wildlife Department Coastal
relationship between oyster harvest and freshwater inflow           Fisheries Division Management Data Series Number 125. 36 pp.
                                                               Keck, R.D., D. Mauer, J.C. Kaver, and W.A. Sheppard. 1971.
without considering the complex, important role high fresh-         Chemical stimulants affecting larval settlement in the American
water inflow events play in ensuring long-term oyster               oyster. Proceedings of the National Shellfish Association 61:
health.                                                             24–28.
   We concur with Turner’s (2006) caution that sustainable     La Peyre, M.K., A.D. Nickens, A.K. Volety, G.S. Tolley, and J.F. La
                                                                    Peyre. 2003. Environmental significance of freshets in reducing
and healthy ecosystems are based on a wide variety of               Perkinsus marinus infections in eastern oysters Crassostrea
factors, natural and anthropogenic. Estuarine management            virginica: Potential management applications. Marine Ecology
should be done in “…the context of the whole estuary and            Progress Series 248: 165–176. doi:10.3354/meps248165.
not just part of the estuary.” In order to maintain healthy    Meyer, D.L., and E.C. Townsend. 2000. Faunal utilization of created
                                                                    intertidal eastern oyster (Crassostrea virginica) reefs in the
estuaries, oyster communities, and the coastal communities          southeastern United States. Estuaries 23: 34–45. doi:10.2307/
that rely on them, we must understand the unique relation-          1353223.
ship between freshwater inflow and estuary health for          Montagna, P., and Kalke. 1995. Ecology of infaunal mollusca in South
each estuary (Wilber 1992; Bergquist et al. 2006). Society’s        Texas estuaries. American Malacological Bulletin 11: 163–175.
                                                               National Oceanic and Atmospheric Administration. 2007. Fisheries
growing ability to capture and manage water in watersheds           Statistics. Commercial Fisheries. Annual Commercial Landing
with limited water supplies may potentially reduce the              Statistics for Eastern Oyster for the Chesapeake Region.
frequency and magnitude of freshwater inflow events,                http://www.st.nmfs.noaa.gov/st1/commercial/landings/annual_
causing salinities to gradually increase for longer times to        landings.html.
                                                               Pierce, M.E., and J.T. Conover. 1954. A study of the growth of oysters
unhealthy levels for oyster populations.                            under different ecological conditions in Great Pond. Abstracts of
   Paradoxically, floods may have short-term negative               Papers Presented at the Marine Biological Laboratory. The
consequences for oyster harvest but play a vital role in            Biological Bulletin, Woods Hole 1072: 318.
ensuring the long-term health of oyster populations and        Powell, E.N., J.M. Klinck, E.E. Hofmann, and M.A. McManus. 2003.
                                                                    Influence of water allocation and freshwater inflow on oyster
consequently the long-term sustainability of oyster harvest.        production: A hydrodynamic-oyster population model for
We believe variations in freshwater inflow to an estuary            Galveston Bay, Texas, USA. Environmental Management 31:
(particularly associated with moderate floods and freshets)         100–121. doi:10.1007/s00267-002-2695-6.
212                                                                                                   Estuaries and Coasts (2009) 32:206–212

Quast, W.D., M.A. Johns, D.E. Pitts, Jr., G.C. Matlock, and J.E. Clark.   Texas Parks and Wildlife Department. 2006a. Marine resource
    1988. Texas Oyster Fishery Management Plan. Fishery Manage-               monitoring program: Coastal fisheries database. Austin, Texas
    ment Plan Series Number 1. Source Document. Texas Parks and           Texas Parks and Wildlife Department. 2006b. Unpublished data on
    Wildlife Department, Austin, TX. 178 pp.                                  commercial oyster landings.
Ray, S. M. 1987. Salinity requirements of the American oyster,            Texas Water Development Board. 2007a. In Trinity-San Jacinto
    Crassostrea virginica. In: Freshwater Inflow Needs of the                 Estuary Freshwater Inflows (Galveston Bay). http://hyper20.
    Matagorda Bay System with Focus on Penaeid Shrimp. NOAA                   twdb.state.tx.us/data/bays_estuaries/TxEmp/galv_index.htm
    Technical Memorandum, NMFS-SEFC-189, Galveston, Texas.                Texas Water Development Board. 2007b. In Coastal Hydrology, Basin
Rodney, W.S., and K.T. Paynter. 2006. Comparisons of macrofaunal              Wide Inflow Summary for Galveston Bay. http://hyper20.twdb.
    assemblages on restored and non-restored oyster reefs in                  state.tx.us/data/bays_estuaries/hydrology/galvestonsum.txt
    mesohaline regions of Chesapeake Bay in Maryland. Journal of          Turner, R.E. 2006. Will lowering estuarine salinity increase Gulf of
    Experimental Marine Biology and Ecology 335: 39–51.                       Mexico oyster landings. Estuaries and Coasts 29: 345–352.
    doi:10.1016/j.jembe.2006.02.017.                                      Wilber, D.H. 1992. Associations between freshwater inflows and
Texas General Land Office. 2006. Internet. Available from www.glo.            oyster productivity in Apalachicola Bay, Florida. Estuarine,
    state.tx.us http://www.glo.state.tx.us. Accessed September 1, 2006.       Coastal and Shelf Science 35: 179–190. doi:10.1016/S0272-
Texas Parks and Wildlife Department. 2002. Marine resource                    7714(05)80112-X.
    monitoring operations manual. 104. Austin: Texas Parks and            Zimmerman, R., T. Minello, T. Baumer, and M. Castiglione. 1989.
    Wildlife Department.                                                      Oyster reef as habitat for estuarine macrofauna. National Oceanic
Texas Water Development Board. 2006. Water for Texas 2007.                    and Atmospheric Association Technical Memorandum NMFS-
    Volume I. Austin, TX.                                                     SEFC-249.
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