Melissa Southworth

Population Studies of the Native Eastern Oyster, Crassostrea virginica, (Gmelin, 1791) in the James River, Virginia, USA

ABSTRACT We describe oyster population trends in the James River, VA from 1993 through 2006 using quantitative fishery independent survey data collected using a stratified random design. The 23 reefs contained in the study area cover a total of 2.41 × 107 m2 and vary in individual size from 1.26 × 104 m2 to 4.98 × 106 m2. There is a marked pattern in density of oysters among the reefs: during the study period a small group of reefs comprising 5.4% of the total area consistently contained between 25.7 and 55.5% by number and 35.8 and 54.8% by biomass of the total oyster population. The highest density reefs exhibit, with very few exceptions, mean densities well in excess of 200 oysters m-2, typically between 300 and 500 m-2, with a single maximum value of 773 oysters m-2 in 2002 coincident with the highest annual recruitment observed during the study period. Recruitment events were usually followed by very high mortality with very small percentages of the population reaching ages ≥3 y of age. A strong stockrecruit relationship is absent; rather population demographics appear to be dominated by periodic high recruitment events. Biomass maxima tended to lag one to two years after recruitment maxima. Standing stock for the total system varied between 1.07 × 108 g and 3.31 × 108 g (107 and 331 metric tonnes) in 2003 and 2005, respectively as the 2002 recruits grew and suffered mortality. Age-at-length relationships were estimated from demographics: using a July 1 birth date and a November 1 survey date giving lengths of 37.3 mm at 0.33 y, 58.9 mm at 1.33 y, 80.5 mm at 2.33 y, 102.1 mm at 3.33 y and 123.7 mm at 4.33 y Length demographics were recast as age demographics to estimate annual proportional mortality. Mean proportional mortality values for age 1 oysters range from a low of 0.2–0.4 to a high in excess of 0.7. Age 2 mean proportional mortality values range from a low of 0.41 to a high exceeding 0.75. The proportional mortality for age 3 and 4 y olds generally exceeded mean values of 0.6 with highest values approaching 0.95. In all cases, these values exceeded mortality estimates calculated using traditional box count methods by a considerable margin. The ability to accurately estimate age specific mortality allows the construction of shell (habitat) budgets for the individual reef systems. Shell half-life loss rate estimates in the most productive reefs is between 2 and 3 y, and the population is maintained by the continual and extraordinary recruitment in the face of high mortality - the latter driven by disease (predominantly Perkinsus marinus) epizootics. The shell resource, even on the most productive reefs, is modest, equivalent to little more than a monolayer several centimeters thick. Individual reefs demonstrate remarkable stability as either high shell density + high population density associations (high:high) or low shell density - low population density associations (low:low), even in the face of temporal population and demographic fluctuations associated with disease related mortality. The probability of manipulating either shell and/or live oyster density to effect the transition of a low:low reef to a high:high reef is considered bleak in the face of extant recruitment and mortality patterns. The primary impediment to population expansion or rebuilding is high and uncontrolled mortality rather than a lack of recruitment. Given the large numbers of oysters in low salinity refugia that have the ability to continually contribute to the larval pool, active selection against disease susceptible oysters on a system wide basis is unlikely.

OYSTER (CRASSOSTREA VIRGINICA, GMELIN 1791) POPULATION DYNAMICS ON PUBLIC REEFS IN THE GREAT WICOMICO RIVER, VIRGINIA, USA

ABSTRACT We describe oyster population trends in the Great Wicomico River, VA, from 2009 through 2009 using quantitative fishery independent survey data collected using a stratified random design. The seven public reefs examined cover a total of 2.8 × 105 m2 and vary in individual size from 1.36 × 104 to 7.16 × 104 nr. The river is functionally divided by a sand spit into upriver and downriver regions. Oyster densities on the upriver reefs were typically an order of magnitude higher than densities on the downriver reefs within the same time period. Throughout the system, the highest observed densities were coincident with high annual recruitment events (2002, 2006). Recruitment events were usually followed by high mortality, with small percentages of the population reaching ≥3 y of age. A predictive stock—recruit relationship is absent; rather, population demographics appear to be dominated by periodic high recruitment events. In the absence of seed removal, biomass maxima follow 1–2 y after recruitment maxima. Standing stock for the system varied between 1.56 × 106 g and 3.63 × 107 g in 2005 and 2006. Year-specific age-at-length relationships were estimated from demographics data. Length demographics were recast as age demographics to estimate mortality. Observed proportional mortality between young of the year and age 2 oysters was approximately 0.88 for the 2006-y class, which is slightly higher than the 0.62–0.71 observed for the 2007-y class. The ability to estimate age specific mortality accurately allows the construction of shell (habitat) budgets for the individual reef systems. The Great Wicomico oyster population appears to be maintained by episodic and extraordinary recruitment in the face of high mortality—the latter driven by disease (predominantly Perkinsus marinus) epizootics. The shell resource is modest, equivalent to little more than a monolayer several centimeters thick. Over short timescales (years), the available shell resource oscillates in concert with mortality. The shell accretion rate on upriver reefs is consistently 4–5 times greater than that observed on downriver reefs. Periodic modest shell planting has maintained the habitat base (the shell resource) throughout the system over decadal scales.

Shell Length-at-age Relationships in James River, Virginia, Oysters (Crassostrea virginica) Collected Four Centuries Apart

Abstract Eastern oysters were ecologically and structurally dominant features of the Chesapeake Bay prior to European colonization. Four centuries of harvest pressure, habitat degradation, and, more recently, disease activity have affected extant oyster population demographics. We compared population demographics and age-at-shell length relationships for modern mesohaline James River oyster populations with James River oysters collected in the years 1611 to 1612 by Jamestown settlers. Historic oyster collections made by hand included a more complete demographic than modern samples collected with patent tongs. Historic oysters had significantly faster growth rates than modern oyster populations. Modern oysters larger than 30–40 mm SL or age 1 grow more slowly than historic oysters of comparable ages. Unlike historic oyster populations, modern James River oyster populations are affected by Dermo and MSX. The downward trend observed in the modern age at length relationship (Fig. 4B see later) between 1 to 1.6 y is probably related to the seasonal onset of disease with increasing temperatures. Observed changes in oyster demographics and growth rates across four centuries reflect changes in the environment as well as changes in oyster biology because of chronic pressure from two oyster diseases.

MANAGEMENT OF THE PIANKATANK RIVER, VIRGINIA, IN SUPPORT OF OYSTER (CRASSOSTREA VIRGINICA, GMELIN 1791) FISHERY REPLETION

ABSTRACT The Piankatank River is a trap-type estuary on the western shore of Chesapeake Bay that has been managed for seed oyster production since 1963. Market oyster production in the river is minimal. Repletion efforts include shell planting and seed removal. We describe sequential changes in population demographics and habitat in relation to repletion activities on eight Piankatank River public oyster reefs from 1998 through 2009. Two reef groups (northern and southern) may be distinguished by density (oysters/m2), biomass (g dry tissue weight), and shell volume (L/m2) data. Age-at-length relationships were estimated from demographic data using a quadratic model. Observed mortality rates were high, and age 3+ oysters were essentially absent. A strong recruitment signal was observed in 1999 and 2002. Between 1998 and 2009, about 30% of the live oysters in the river were harvested as seed, corresponding to ∼7.5% of the total shell base in an average year. Typically, for every 5 bushels of shell planted, 1 bushel of seed was harvested (20% return). Even with shell planting (∼10 L/m2/y), the river shell budget showed a deficit with respect to the accretion rate required to balance sea level rise and natural degradation processes. During the study period, the mean river recruit-to-stock ratio was ∼4. The unusual and consistently high recruit-to-stock ratios suggest that management for modest continuous seed removal may be accomplished without shell planting. Annual stock assessment to identify low recruitment years is recommended as a method to adjust annual seed harvest quotas.

NORTHERN QUAHOG (HARD CLAM) MERCENARIA MERCENARIA ABUNDANCE AND HABITAT USE IN CHESAPEAKE BAY

Abstract Recent (2001–2002) surveys of hard clam Mercenaria mercenaria density and distribution, using patent tongs in a stratified random design (n = 7,358 stations) in lower Chesapeake Bay are not consistent with historic descriptions of clam habitats and densities. The highest average densities observed, up to 3.1 clams m−2, were in the lower James River. The highest modern average density observed is half that of clam densities commonly observed in these same habitats during the early 1970s. Current distribution is significantly affected by water depth and substrate composition. Hard clam density in Chesapeake Bay is positively associated with increasing sediment grain size; 78% of all clams collected were found in shell or sand habitats. However, 44% of sand habitats and 54% of shell habitats were unoccupied suggesting that even habitat types that typically support higher clam densities may currently be underused.

Comparison of Crassostrea virginica Gmelin (Eastern Oyster) Recruitment on Constructed Reefs and Adjacent Natural Oyster Bars over Decadal Time Scales

Abstract Since 1993, oyster reef replenishment efforts in the Virginia portion of the Chesapeake Bay have relied heavily on construction of oyster shell reefs with enhanced vertical relief. We evaluated the performance of six reefs constructed in proximity to natural subtidal oyster bars by comparing recruit densities (spat m-2, where spat are young-of-the-year oysters with shell heights less than 50 mm) between habitats. Recruitment was higher on the reefs than bars during the first 1–3 yr post-construction, usually by at least an order of magnitude. Within 7 yr, recruitment was similar between reef-bar pairs although both reefs and bars received additions of shell, live oysters, or both during the study period. At decadal time scales, constructed oyster reefs did not show enhanced recruitment relative to adjacent natural oyster bars. The rapid decline in reef recruitment post-construction is likely related to three processes: (i) shell degradation by taphonomic processes, (ii) biofouling that occludes the shell surface to recruitment, and (iii) inability of extant oysters on the reef to produce new shell at a rate commensurate with losses to (i) and (ii). There appears to be a requirement for continued replenishment activity to maintain the shell base on these reefs, contrary to the dynamics of a healthy natural oyster population. The similarity in recruitment between constructed reefs and natural bars at decadal time scales suggests that subtidal shell plants or shell additions to natural bars may be a more cost-effective repletion strategy because they provide equal population enhancement per unit area.

Observations of blue crabs (Callinectes sapidus, Rathbun 1896) on Shell Bar oyster reef, Great Wicomico River, Virginia.

ABSTRACT Blue crabs (Callinectes sapidus, Rathbun 1896) were sampled with commercial crab pots on Shell Bar oyster reef in the Great Wicomico River, VA, from May through October during 2006 and 2007. Weekly catch per unit effort (CPUE), sex ratio, and size (carapace width, measured in millimeters) were evaluated in the context of water temperature (measured in degrees Celsius), salinity, and daylength (measured in hours) conditions on the reef. The total number of crabs collected in 2006 and 2007 was 5,221 and 3,303, respectively. Blue crab CPUE was highest from mid-June through mid-September at water temperatures at or more than 26°C, with maximum CPUEs observed in late July. The overall annual ratio of females to males was 0.47 in 2006 and 0.60 in 2007. Males made up more than 50% of the catch at Shell Bar reef from May through August. The observed sex ratio shifted from male dominated to female dominated as salinity increased, whereas water temperature and daylength decreased seasonally. Female crabs (median carapace width (CW), 142–144 mm) were approximately 20 mm in CW larger than males (median CW, 125 mm) in both years. Annual average CPUE (13.42 crabs per pot; SE = 1.57) and maximum CPUE (32.06 crabs per pot) was twice as high in 2006 as it was in 2007. The observed interannual differences in crab CPUEs may be the result of an enhanced forage base on the reef in 2006 relative to 2007. Approximately 9 million cultchless oysters (40–80 mm in shell height) were planted on Shell Bar reef between May and October 2006 as part of a concurrent oyster rehabilitation program.

Oyster (Crassostrea virginica [Gmelin, 1791]) Mortality at Prolonged Exposures to High Temperature and Low Salinity

ABSTRACT Mortality of two size classes (<35 and >35 mm) of eastern oysters Crassostrea virginica when exposed to combinations of low salinity (1, 2, 3, and 4) for extended periods (up to 30 days) at summer water temperatures typical of the Virginia Chesapeake Bay subestuaries was examined. A critical salinity-temperature combination of less than two at greater than 28°C for more than 1 wk exposure for oyster mortality is suggested. A review of limited historical salinity-temperature tolerance data suggest selection of local populations of oysters having differing salinity tolerances. Such selection may prove critical to persistence of low-salinity populations in the Chesapeake Bay subestuaries with projected climate change.

Oyster Planting Protocols to Deter Losses to Cownose Ray Predation

ABSTRACT The utility of shell overlays to oyster (Crassostrea virginica) plantings as a cownose ray (Rhinoptera bonasus) predator deterrence mechanism was examined. Typical industry practice of oyster seed planting was followed in an experimental design employing treatment areas of 0.5–1.0 acre (0.2–0.4 hectare). Areas were prepared in the Lower Machodoc Creek, Virginia, by the initial application of shell to insure a stable substrate under planted seed oysters. Seed oysters were planted using standard industry methods. Experimental areas were located, two upstream and two downstream, of a constriction in the Lower Machodoc that dictated differing physical environments in the respective locations with downstream locations being more exposed to northeast wind-driven stresses and, historically, a greater incidence of ray predation. Once oysters were planted, two of the areas, one upstream and one downstream of the aforementioned constriction, were additionally treated with a shell overlay as a predation deterrent. Oyster seed were planted in the experimental plots in February 2012. Market oysters were harvested from the experimental plots in December 2013 and January 2014. Final harvest data demonstrated that shell overlays do not offer additional protection to planted oyster seed with respect to possible cownose ray predation. Evidence of predation in the form of characteristically broken oyster valves were recorded in all treatment areas. Concurrent stomach content analysis of rays captured at the study location and observations of fouling community associated with the cultured oysters taken during the harvest operation indicate broad dietary preferences for rays when such a variety exists in the foraging region. For rays, oysters are not the singular preferred diet item, although localized and intensive feeding on oysters remains an option with a wide foraging range. Areas without overlay demonstrated higher production than those with shell overlay. Shell overlays are not recommended as predator deterrents for cownose rays in large deployments of unprotected oyster seed.

Temporal Variation in Fecundity and Spawning in the Eastern Oyster, Crassostrea virginica, in the Piankatank River, Virginia

ABSTRACT Oysters of the genus Crassostrea are considered good examples of an r-selected marine invertebrate with small egg size, high fecundity, and multiple spawning events per year, each characterized by significant individual weight loss. Historical (decadal) data for the Virginia portion of the Chesapeake Bay support these generalities. We present recent (subdecadal) data, collected for natural Crassostrea virginica broodstock of populations in the Piankatank River, Virginia. The relationship is described between oyster size, fecundity, spawning periodicity, and egg viability for natural broodstock. Oysters collected throughout the summers of 2010 through 2012 and induced to spawn by thermal cycling released viable eggs on 7 dates (n = 119 oysters, 35 male, 84 female; shell length (SL) range, 58–113 mm). Oysters were opened to examine sex ratio on four additional dates (total n — 242 oysters, 82 male, 160 female). Fecundity varied in the range 105-1.2 × 108 eggs. When all data are considered in unison, no strong relationship with SL is evident; however, when eliminating the artifact of data corresponding to minimal egg release, a much stronger relationship, comparable with that reported in older literature, emerges. Female fraction (Female/(Female + Male)) was consistently more than 1 in oysters larger than 60 mm in SL (estimated age, ≥2 y), generally in accordance with recently published literature on the species in the mid-Atlantic. The size-versus-fecundity relationship does not appear to be greatly influenced by disease prevalence/ intensity. The temporal sequence of spawning activity was not observed to continue after midsummer and is not commensurate with a cumulative degree-day estimator during the latter half of the well-documented historical spawning season. A size-fecundity estimator for the Piankatank River oysters provides a basis to estimate the disproportionate value of larger/older (≥3 y) oysters in the system, and provides additional input to the fine-tuning of a previously developed rotational harvest schedule for the river stock. The possible impact of recent changes in water quality, seasonal occurrence of dinoflagellate blooms, and/or long-term impacts of changing regimes were not examined in detail in this study but are suggested as worthy lines of future investigation.