The nearly forgotten oyster: Ostrea lurida Carpenter 1864 (Olympia oyster) history and management in Washington State.ABSTRACT Overexploited fisheries are a worldwide problem. Restoration efforts aimed at these fisheries often involve a combination of reduced catch, hatcheries, and habitat improvement. The native oyster of western North America, Ostrea lurida, ([dagger]) was commercially extinct in most locations more than a century ago. In this paper, we track the history of its management for insight into its demise and failed recovery in Washington State. We document six phases of management: open access, aquaculture, control of water pollution sources, substitution by nonindigenous oysters, harvest regulations and marine reserves, and restoration. Three general lessons emerge from this historical analysis, which may apply generally to exploited fisheries that fail to recover. First, the introduction of substitute species led to neglect of the native species for many decades. Second, reserves were not fully protected and instead were designed for commercial removal of newly settled oysters, thus they largely failed. Finally, current restoration efforts are hampered by several biological problems, including water pollution, invasive competitors and predators, and habitat loss.
KEY WORDS: oyster, Ostrea lurida, Crassostrea gigas, Olympia oyster, management, overexploitation, restoration
Oyster fisheries have poor records of sustainability worldwide. Native species have dwindled to commercial extinction in Europe, New Zealand, and eastern North America (Brooks 1891, Dinamani 1991, Goulletquer & Heral 1991, Kirby 2004). The native oyster of the west coast of North America also followed this trend: commercial yields of Ostrea lurida in Washington State dropped by more than 98% between 1881 and 1961 (Fig. 1), and today the oyster represents a tiny fraction (<1%) of the United States west coast oyster harvest (Ruesink et al. 2005). O. lurida is exceptional only in how little scientific attention has been paid to understanding causes of decline and methods of restoration. Scientific articles about O. lurida total just 13 in recent peer-reviewed journals (Aquatic Science and Fishery Abstracts: search terms Ostrea or Ostreola and conchaphila or lurida from 1970 to 2005), compared with 1499 for Crassostrea virginica, the native oyster of the east coast of North America. In this paper, we compile historical data from personal accounts of oyster growers, government agency reports, academic reviews, newspapers, and newsletters. The emerging story of the nearly forgotten oyster reveals several lessons about why commercial fisheries decline and fail to recover.
More than a century ago, O. lurida had clear cultural and economic importance. The presence of shells in middens demonstrates that Native American tribes harvested O. lurida for at least 4000 y (Blake 2003). The oyster may have played a role in the selection of the city of Olympia as Washington's capitol, as a feast of O. lurida was provided to delegates at a dinner in Olympia (Steele 1957). The demand for oysters in western United States urban centers largely drove commercial exploitation. San Francisco initially represented the largest market, and oysters were shipped from successively more distant locations as nearby stocks dwindled (Kirby 2004). The period of commercial exploitation was brief: Willapa Bay shows a common pattern in which the fishery began in the 1850s and essentially ended by 1915. Washington's Puget Sound was unusual because measurable numbers of O. lurida were still harvested in the 1940s (Fig. 1). Current production of O. lurida derives primarily from aquaculture, providing a "boutique" oyster of small size and unique metallic flavor, considered a distinct native food source with strong cultural ties to family harvest operations on the west coast (Warren & Downey 2003).
The propensity for overexploitation of O. lurida stems in part from its life history characteristics. Growth is slow, requiring several years to reach a marketable size of 5 cm (Couch & Hassler 1989). This small adult size requires ~2,400 individuals for a gallon of shucked meat (4,000 individuals/sack [Brennan 1939]; 1.66 gallons/sack). The species is relatively sensitive to heat, cold, and desiccation stress and was naturally found primarily in accumulations of shell below mean lower low water (Steele 1957, Lloyd 1996). Completion of the life cycle requires clean shell in suitable habitats for larval attachment (White et al. 2009). Oysters also settle on other hard substrates such as glass plates, rock, and concrete (Hopkins 1935), but these are not typically available in soft sediment environments. The removal of oysters resulted in the degradation of their shell habitat, because oysters were often shipped live and not shucked in place.
Despite depressed population sizes, larvae of O. lurida can be abundant in the plankton, as observed in Puget Sound in the 1970s (Terry Nosho, pers. comm.) and more recently in Willapa Bay (Trimble et al. 2009). This provides circumstantial evidence that recovery is not limited by broodstock but by habitat. O. lurida matures rapidly (<1 y) and produces abundant larvae (250,000 per female). It is an alternating hermaphrodite and, when female, broods larvae for 10-12 days in the mantle cavity prior to release into the plankton (Hopkins 1937). This reproductive mode requires large ostia (gill pores), that prevent capture of small particles by the gills. Overall, four characteristics distinguish the species from most other commercial oysters: subtidal habitat, slow growth and small size, ovoviviparity, and large ostia influencing particle size selection (Couch & Hassler 1989).
[FIGURE 1 OMITTED]
Management of O. lurida went through six overlapping phases in Washington State: open access, aquaculture, control of water pollution sources, substitution by nonindigenous oysters, harvest regulations and marine reserves, and restoration. Our review of this history reveals three general lessons for fisheries restoration: substitution of species may divert attention away from threatened species, marine reserves should be fully protected to function effectively, and control of biological problems such as point and nonpoint pollution sources, invasive species, and habitat loss may be necessary for recovery to occur. With restoration projects underway, O. lurida may be able to rebound on the West Coast, but the oyster's history holds lessons for marine species management that may help explain the difficulty of restoring overexploited fisheries.
PHASES OF MANAGEMENT
Open Access Fishery
For Native Americans of western Washington, O. lurida provided food and a trade item (Hutchinson 193-, Steele 1957, Lloyd 1996, Gordon et al. 2001, Blake 2003, Cook 2004). Nevertheless, large natural beds of this native species still existed in the 1850s (Mogan 1931, Lloyd 1996, Rogers 2004). Around the year 1850, during the Gold Rush in San Francisco, oysters sold for as much as S20 a plate (Rogers 2004). Oysters were a party food, symbol of wealth, and generally a popular dish. The last meal of convicted criminals often included oysters--Hangtown Fry (Gordon et al. 2001). Only canned oysters were available through import from the East Coast, so the high demand and premium price for fresh oysters led to the development of a new fishery for the native oyster of the North American west coast. For instance, J. W. Russell first took O. lurida from Willapa Bay to San Francisco in 1851 (Lloyd 1996), and two years later regular imports were arranged by John Mogan (Lloyd 1996, Rogers 2004).
European settlers removed more than 5 billion individual O. lurida from Willapa Bay between 1851 and 1915 (Fig. 1), a figure that is surely an underestimate because of missing data. These oysters were harvested from thick beds built up over time, so we can infer an initial population on the order of billions of individuals. The subsequent shipment of live oysters to distant ports prevented the replacement of shell (cultch) necessary for recruitment of new oysters. This reliance on built-up stocks and removal of habitat led to rapid overexploitation and decline of O. lurida.
Overexploitation occurred despite low accessibility of the oysters, which were largely low intertidal and subtidal, and harvested by nonmechanized methods. In the 1850s in Willapa Bay oystermen poled or sailed boats to beds and collected oysters with long-handled tongs or dredges (Swan 1857). Initially, schooners sailed directly from Willapa Bay, although rail and steamboat travel reduced transport time by 1889 (Lloyd 1996).
Hatcheries and aquaculture are common responses to revive failed fisheries. In Puget Sound, O. lurida was harvested for decades after failures in other locations because of the development of cultivation techniques. Oyster growers built 6- to 12-inch thick concrete or creosote wood dikes to maintain water over their intertidal oyster beds (Hopkins 1937, Steele 1957). This innovation extended the growing area for oysters intertidally and reduced early mortality because oysters were protected from temperature fluctuations. However, this technique still required natural recruitment of seed oysters that were then transported onto the diked beds (Staricka 1975). Consequently, cultivation enhanced habitat but did not involve complete control over the entire life cycle. Hatchery techniques to provide a reliable source of seed oysters were not developed until the 1970s, long after natives had been largely replaced commercially (Chew 1991).
A substantial industry developed in Puget Sound based on dike cultivation of oysters. For many involved in the early industry in Puget Sound, O. lurida were "the chief food and livelihood" (Chase 1976). The Olympia Oyster Grower and Dealers Association was founded in 1914 to aid those involved in the industry and establish positive relationships among growers (Steele 1957).
In contrast, cultivation of native oysters was largely unsuccessful in Willapa Bay and other United States west coast estuaries. The cultivation efforts of the 1890s came too late after depletion of the population, and few dikes were built. In the absence of dikes, 60% of intertidal oysters were estimated to succumb to a severe freeze in January 1888 (Collins 1892). Early intertidal cultivation attempts were also destroyed by several cold winters in the mid-1890s (Staricka 1975, Driscoll 2003).
Control of Water Pollution Sources
Cultivation extended the window of production of O. lurida in Puget Sound, but not indefinitely. Circumstantial evidence implicates water pollution as a major factor in damaging oyster populations, although litigation never settled on this answer. Poor timber and mining practices in the upland areas of many United States west coast estuaries caused serious erosion, possibly smothering beds of O. lurida. For instance, historic photos of Netarts Bay, OR, show sailing schooners at anchor in parts of the estuary that today are vast mudflats at low tide. Tribal records indicate that water pollution from timber mills damaged oyster populations in Puget Sound as early as the 1860s (Steele 1957, Staricka 1975, Cook 2004). Growers in south Puget Sound were most concerned by a pulp mill that opened in Shelton, WA in 1927 (Chasan 1981), releasing sulfite waste liquor into a highly-productive oyster growing region.
Before the mill was opened, oceanographic predictions were made that tidal currents would move sulfite waste out of Oakland Bay and reach oyster beds within a few days of release (McKernan et al. 1949). In 1926 the Olympia Oyster Growers Association warned the Washington Department of Fisheries of the threat, the Association of Pacific Fisheries called for treatment of the waste, and a Washington State Fisheries Board committee reviewed pollution problems, but the discharge was approved (Chasan 1981, Gordon et al. 2001).
An investigation into the causes of decline of O. lurida by the Washington Department of Fisheries suggested that mortality of adults, poor condition, and lack of recruitment all occurred during the years of mill operation with no change in other environmental factors measured. Direct toxicity studies indicated that the effluent led to death, reduced quality, thinning of the shell, and reduced survival and setting of recruits. The effluent became less toxic with age, and impacts varied throughout south Puget Sound. Sulfite waste also produced indirect effects by reducing plankton abundances and fertilizing waterways. Based on the clear temporal correlation as well as studies demonstrating effluent toxicity to the oysters, department officials implicated the pulp mills as the primary cause of decline in Puget Sound (McKernan et al. 1949, Gunter & McKee 1960).
Oyster growers on Oakland Bay and Little Skookum sued the Rayonier Pulp and Paper Company for damages caused by pollution (Holt 2000). An out of court settlement was reached in 1931with the guarantee that no additional waste would be dumped into Mill Creek (Nelson 1990). Instead land at Goose Lake was purchased for discharge of liquor, but the waste seeped into water supplies (Bay 1975, Nelson 1990). The mill was closed periodically with corresponding improvements in oyster conditions, and in 1957 the mill shut down permanently for business purposes unrelated to the oysters (Chasan 1981, Holt 2000).
A final suit led by the Olympia Oyster Company went through the United States District Court in Tacoma, and in 1959 Judge George Boldt heard the case, but it was dismissed because of limited evidence (Chasan 1981, Holt 2000). The circumstantial and laboratory evidence available was insufficient to aid oyster growers in their 1959 suit. Interestingly, improvements in growth, fatness, and mortality rates were noted in the years after closure of the pulp mill at Shelton (WDFW 1961).
This segment of the Olympia oyster's history in Washington characterizes legal debates about a variety of environmental disasters, in which scientists from both sides weigh in with conflicting evidence (e.g., Paine et al. 1996). The difficulty in detecting environmental impact stems from the absence of baseline data and "control" sites without impacts.
Substitution of Species
Non-native species of oysters reduced the importance of O. lurida in two ways: at markets and in estuaries. The smaller size and stronger flavor of O. lurida made them less desirable than Crassostrea virginica for new immigrants from the United States east coast where C. virginica is native. Imports of C. virginica came by transcontinental rail in the 1870s, and accordingly the price in San Francisco for O. lurida fell dramatically from S16 a sack in the 1860s to S4 in the 1870s and S2.50 in the 1880s (Lloyd 1996).
C. virginica was also the first nonnative oyster to be planted directly in west coast estuaries. Both adult and seed oysters were planted in California in the 1870s, and transplants also occurred in Willapa Bay in 1894 (Townsend 1896). The transition to cultivation of C. virginica largely ended commercial interest in O. lurida in Willapa Bay (Woelke 1969). However, the growth of the C. virginica industry in Willapa Bay was short lived, as a dramatic population crash occurred in 1919. The cause of this decline is unclear, but may have been triggered by red tide, failed reproduction at east coast spat sources, or lack of local reproduction (Lloyd 1996, Gordon et al. 2001).
Crassostrea gigas (Pacific oyster) was imported from Japan to Washington as early as 1899 (Kincaid 1968). Concerted efforts to introduce this species began in 1919, when two Japanese-Americans (J. Emy Tsukimato and Joe Miyagi of the Pearl Oyster Company) planted C. gigas oysters that had been imported from Japan's Sendai Bay. Adult oysters were unable to withstand the long sea voyage across the Pacific Ocean, but attached recruits survived and flourished (Steele 1964). However, the Alien Land Law of 1921 forbade Japanese from owning land, and the men were forced to sell to Puget Sound oyster grower E.N. Steele (Steele 1964). Pacific oysters were subsequently imported to Willapa Bay in 1928 by Gerard Mogan, who had recently purchased 7,000 acres of unused tideland and attempted to revive the industry (Lloyd 1996).
Growing interest in C. gigas reduced cultivation of O. lurida, especially after declines in O. lurida reproduction and survival resulted from pulp mill pollution. C. gigas has proven to be a versatile generalist, introduced to at least 64 countries worldwide and established in 28% (Ruesink et al. 2005). It grows faster and larger than O. lurida (Steele 1957, Kincaid 1968). C. gigas requires little more than a layer of intertidal shell for cultivation, though stakes, long lines, anchors, baskets, or trays may be used (Nosho 1975). Likewise, they do not need dikes or grow in deep waters that require dredging, as is the case with the O. lurida and C. virginica (Steele 1964). C. gigas was not immediately embraced by United States consumers, but growers embarked on creative marketing strategies to increase sales (Steele 1964). To this extent commercial interests were redirected, and efforts to cultivate O. lurida fell to only a few growers (Cook et al. 1998).
Establishment of C. gigas occurred in a few locations in Washington by 1936 (Anonymous 1936), but natural recruitment was too small to support the needs of the industry for seed oysters. Thus, annual imports of seed came from Japan from the 1920s to 1977, when local hatcheries replaced imports (Steele 1964, Scholz & Tufts 1977, Figure 2). On average, 40,000 cases of seed were shipped each year, except for a brief hiatus during WWII. Several other species, including Ostrea edulis from Europe, Crassostrea sikamea from Japan, and Crassostrea ariakensis from Japan and China have been imported, reared in hatcheries, and outplanted for harvest. These and C. virginica continue to represent a small fraction of the total oysters harvested in Washington State, as C. gigas predominates (Gordon et al. 2001).
[FIGURE 2 OMITTED]
The ready availability of more economically-desirable, ecologically-robust oysters reduced the attention paid to O. lurida by shellfish growers. In addition, the nonindigenous oysters created ecological conditions that may thwart native oyster recovery. Large-scale imports of nonindigenous oysters were accompanied by numerous hitchhiking species (Ruiz et al. 2000), some of which now negatively affect O. lurida. Most attention has been paid to the Japanese oyster drill, Ocinebrinus (Ocinebrellus = Ceratostoma) inornata, an oyster predator that arrived in Washington by the 1930s (Hopkins 1937, Lindsay 1970). Some initial concern was expressed about the effects of drills on O. lurida, because its thinner shell would be easily penetrated (Hopkins 1937), but management has responded to the drill as a pest of cultivated bivalves, mostly nonindigenous themselves. Beginning in 1947, the Washington Department of Fisheries sent biologists to Japan to supervise the packing of oyster seed and limit further introductions of the snail (Steele 1964). Transfers of oysters are still regulated and quarantine areas established to reduce further spread of oyster drills (WDFW 1961, Nosho 1975). Many additional introduced species co-occur with O. lurida, including predators (flatworm Pseudostylochus ostreophagous from Japan; oyster drill Urosalpinx cinerea from the eastern United States), parasites (copepod Mytilicola orientalis from Japan), and competitors (slipper limpet Crepidula fornicata from the eastern United States, several compound ascidians) (Woelke 1956. Lindsay 1970). Anecdotal evidence suggests some strong effects, for instance, total loss of Olympia oyster spat from flatworms over two years (Terry Nosho, pers. comm.), however, empirical tests of the impacts of these species have only begun recently (Buhle & Ruesink 2009, Trimble et al. 2009).
Regulations and Marine Reserves
Rapid overexploitation of O. lurida prompted several conservation measures. Some of these regulations were self-imposed by oyster fishers. For instance, in 1855 in Willapa Bay, no oysters were collected during the major spawning period (June 1 to August 1). Prior to Washington's statehood, the territorial legislature promoted cultivation of oysters by allowing private ownership of tideflats outside natural beds, which could be used for growing and holding oysters before shipping. Individuals could own 10 acres in 1863, and this increased to 20 acres in 1877 (Woelke 1969).
Oyster conservation then became one of the first measures addressed after statehood was granted in 1889. Article 17 of the State Constitution gave the state ownership of all tidelands up to high water, but the first legislature passed an act regulating the appraisal and disposal of tidelands (Woelke 1969). The Callow Act of 1890 allowed oystermen to purchase up to 80 acres of tidelands that did not contain natural oyster beds (Woelke 1969). Later the Bush Act of 1895 established classes of land, including those tidelands with populations of oysters, and extended purchasing rights to anyone intending to cultivate oysters (Woelke 1969). A citizen could purchase a tract of up to 100 acres (Mogan 1931). These acts increased oyster farming and production.
Between 1890 and 1910, the Washington legislature also established numerous Oyster Reserves in locations overlapping natural oyster beds (Westley et al. 1985). Reserves initially totaled 11,239 acres in Willapa Bay and 4,500 acres in Puget Sound. In principle, these areas constituted a network of marine reserves, a strategy now widely touted to help protect fisheries and marine biodiversity (Allison et al. 1998). The Oyster Reserves, however, were not fully protected from exploitation. In 1897 the legislature directly specified the protection of natural oyster beds for the public use. In the early 1900s policy mandated that the reserves be improved to benefit the oyster industry and recreational harvesters, but they were also expected to bring in revenue to cover the cost of their management. Initially these funds were managed in the Oyster Reserve Fund, but in 1939 they were transferred to the Department of Fisheries. Oyster seed and adult oysters were sold early on, some efforts were made to improve the reserves (i.e., dike installation), and nonproductive parts of the reserves were gradually sold (Woelke 1969).
By the 1930s, as O. lurida production declined and the industry shifted to cultivation of C. gigas, the Oyster Reserves were of little importance. In the late 1930s and early 1940s C. gigas accumulated on the Willapa reserves (Kincaid 1968), and in 1949 the legislature officially changed reserve management goals to encourage the use of reserves for accumulating seed and supporting oyster growers and recreational harvesters of C. gigas (Woelke 1969). In 1995, a recent revision requested that reserve lands be assigned into five management categories, one of which is native Olympia oyster broodstock (RCW 77.60.030).
Today only 10,000 acres of reserves remain in Willapa Bay and 1,000 acres in Puget Sound caused by lack of effectiveness and interest (Westley et al. 1985, Cook et al. 1998). State Oyster Reserves are no longer actively managed for O. lurida, and rather clams and C. gigas are priorities (Westley et al. 1985, Cook et al. 1998). Willapa Bay oyster reserves are actively managed for commercial harvest of C. gigas, and at least 40% live oyster volume in shell must be returned to the beds for cultch (Dumbauld et al. 2003). No commercial harvest of any oyster species occurs on the Puget Sound reserves. Recreational harvesters target C. gigas, and O. lurida may be collected above a minimum size of 2.5 inches (Cook et al. 2000). Introduced Manila clams (Ruditapes philippinarum) are harvested commercially and recreationally from Oyster Reserves.
Recently interest has grown in O. lurida restoration among state agencies, tribes, oyster growers, and nonprofit organizations (Peter-Contesse & Peabody 2005). Reestablishment of populations could improve water quality, provide habitat for other native species, and potentially allow recreational or low levels of commercial harvest, serving the needs of a range of interests (Ruesink et al. 2005). Indeed, current restoration efforts are community projects involving and benefiting multiple stakeholders (Betsy Peabody, Puget Sound Restoration Fund, pets. comm.).
Since the 1970s Washington State has recognized O. lurida as a candidate threatened species, and in 1998 the Washington Department of Fish and Wildlife created a Stock Rebuilding Plan for the species. The plan aims to "restore and maintain O. lurida populations on public tidelands in their former range" (Cook et al. 1998). Considerations of water quality, habitat improvement, and predator and competitor presence are noted, with the caveat that additional research is needed into nearly all aspects of O. lurida biology and ecology. However, this plan never received a mandate or funding, and rather nonprofit organizations have worked to put the plan into action (Betsy Peabody, Puget Sound Restoration Fund, pers. comm.). Two strategies are common in current restoration efforts: habitat recovery by putting out shell and relying on natural recruitment; and planting juvenile hatchery-raised oysters. Little O. lurida shell is available for restoration, so C. gigas shell has typically been used.
Objectives of these restoration efforts include rehabilitating the species itself, for instance to allow future harvest, improving habitat for other species, for instance by providing water filtration and complex hard substrate in silty areas, and mitigating for coastal development (Yuasa 2003, Gear 2003, Rogers 2004). In Washington, restoration efforts include at least 41 sites out-planted by the Puget Sound Restoration Fund and a marine lease from Washington Department of Natural Resources to The Nature Conservancy (Pearson 2002, Blake 2003).
Successful restoration sites have a combination of characteristics that coincide with O. lurida biology. They are typically subtidal or in moist seeps, have low levels of predation (especially from introduced species), and prevent smothering of oysters by silt. The genetic structure of O. lurida populations is currently being studied, and small-scale genetic differentiation is expected based on the relatively short larval period and isolation of habitat in bays and estuaries (Baker 1995). Local broodstock is crucial for outplanting oysters, which limits some restoration efforts (Blake 2003).
Additionally, nonpoint pollution impacts water quality and creates human health risks associated with oyster consumption. Waste from septic and municipal sewage treatment systems, pet waste, and other storm water contaminants enter Puget Sound, and these sources have all contributed to shellfish harvest closures. About 30,000 acres of commercial shellfish growing area have been closed since the mid-1980s, and more areas are on the verge of closure (PSAT 2005). In some cases oysters grown in threatened areas must be transferred to clean waters to remove any contaminants before they go to market (Holt 2000).
LESSONS FOR EXPLOITED FISHERIES
It is clear from the story of O. lurida that because substitutes (C. virginica and C. gigas) existed to supply the oyster industry, there was little alarm at the loss of the native oyster. The oysters were introduced as pressure grew caused by shrinking natural stocks and increasing water pollution, and aquaculture shifted in response to these introductions. Because oystermen failed to defeat industry polluters, most growers either left the industry or took up cultivation of the heartier C. gigas. Management priorities, even for the State Oyster Reserves, conformed to industry interests. Further, concerns about introduction of species along with oysters, common today, did not exist at the time of the switch to C. gigas. Consequently, numerous pest species were also introduced. The timing and conditions of substitution made O. lurida easy to leave behind.
The establishment of protected areas also failed to prevent loss of native oysters. This system of protected areas was flawed and provided a lower level of protection than was necessary for success. Some extraction of oysters was permitted, and later management switched to C. gigas. In this way the initial goal of the reserves, to preserve the natural beds of O. lurida, was forgotten. In Willapa Bay where the fishery began before statehood, reserves came too late in the process of depletion. Puget Sound reserves, whereas established in sufficient time, suffered because of toxic pollutant inputs, because reserve borders do not prevent toxins from crossing. Altogether these protection level, timing, and pollution factors doomed the reserves to failure.
Decades later, the oysters have failed to recover, and the suite of current problems is somewhat unclear. Lack of hard substrate at appropriate tidal elevations, especially O. luridu shell, may limit recruitment and rebuilding of populations (White et al. 2009). Introduced species also act as competitors and predators (Buhle & Ruesink 2009, Trimble et al. 2009), and these species may be particularly damaging when oysters are already at low density. Currently it is too early to determine if restoration projects will be able to overcome these limitations.
The lessons of this nearly forgotten oyster hold meaning for species management techniques and marine protected areas. Overharvest went unmanaged early on in the O. lurida fishery, and running the oyster reserves based on industry interests hindered preservation of the species after alternative cultivatable species were made available. Further, without the progressive pollution laws created only recently, oyster growers faced tremendous opposition from timber industry stakeholders and were unable to prevent the destruction of their oysters. To this extent, the O. lurida story indicates that overharvest, introduced species, pollution, and habitat loss, some of the major environmental issues of the day, must be addressed in marine and estuarine systems to preserve unique native species and communities.
The authors thank the Washington Research Foundation for their support administered through the University of Washington Undergraduate Research Program's Research Fellowships for Advanced Undergraduates (JMW) and from the Andrew W. Mellon Foundation (JLR, ACT). Critical historic documents were available in libraries of the University of Washington and Washington State archives.
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JACQUELINE WHITE, JENNIFER L. RUESINK * AND ALAN C. TRIMBLE
Department of Biology, University of Washington, Box 351800, Seattle, Washington 98195-1800
* Corresponding author. E-mail: firstname.lastname@example.org
([dagger]) The taxonomy of the Olympia oyster has been in dispute since Harry (1985) proposed synonymy of Ostrea lurida Carpenter 1864 and Ostrea conchaphila Carpenter 1857. Poison et al. 2009 provide molecular evidence that the Olympia oyster refers to the nominal species, Ostrea lurida Carpenter 1864. In view of their genetic data, and for consistency, the original taxon. Ostrea lurida, is used throughout this volume to refer to the Olympia oyster, which is distributed from approximately Baja California (Mexico) to southeast Alaska.