P. Sean McDonald

The competitive and predatory impacts of the nonindigenous crab Carcinus maenas (L.) on early benthic phase Dungeness crab Cancer magister Dana

We evaluate the potential competitive and predatory impacts of nonindigenous European green crab Carcinus maenas on native Dungeness crab Cancer magister in the northeast Pacific. The coastal estuaries of Washington State, USA, provide appropriate habitat for recently introduced green crab, yet these areas are important nursery grounds for Dungeness crab and contribute greatly to the coastal crab fishery. Juvenile Dungeness crabs are dependent on limited intertidal epibenthic shell for refuge habitat during early benthic life and experience increased mortality on open sand and mud as a result of predation by fish and birds. Early juveniles throughout the subtidal are similarly at risk due to predation by fish and especially adult conspecifics. Laboratory experiments and infrared video observations revealed that juvenile green crab displace Dungeness crab of equal size from shelters during one-on-one competition. Green crab also consistently win nocturnal foraging trials in which the species compete for fresh, damaged clams. Field and laboratory enclosure experiments show that juvenile Dungeness crab emigrate from oyster shell habitat as a result of competition and predation by adult green crab. Depending on the extent to which the two species overlap, interactions with the dominant nonindigenous species could have a negative influence on juvenile Dungeness crab survival and could conceivably impact recruitment to the fishery. However, current evidence indicates that the distribution of green crab in Washington State is far removed from nursery areas of Dungeness crab.

Bioenergetics modeling to investigate habitat use by the nonindigenous crab,Carcinus maenas, in Willapa Bay, Washington

A bioenergetics model was developed and applied to questions of habitat use and migration behavior of nonindigenous European green crab (Carcinus maenas) in Willapa Bay, Washington, USA. The model was parameterized using existing data from published studies on the ecology and physiology ofC. maenas and allied brachyuran crabs., Simulations of the model were run describing four scenarios of habitat use and behavior during a 214-d simulation period (April–October) including crabs occupying mid littoral habitat, high littoral habitat, sublittoral habitat, and sublittoral habitat but undertaking intertidal migrations. Monthly trapping was done along an intertidal gradient in Willapa Bay to determine the actual distribution of crabs for the same time interval as the simulation period, and model results were compared to the observed pattern. Model estimates suggest no intrinsic energetic incentive for crabs to occupy littoral habitats since metabolic costs were c. 6% higher for these individuals than their sublittoral counterparts. Crabs in the littoral simulations were also less efficient than sublittoral crabs at converting consumed energy into growth. Monthly trapping revealed thatC. maenas are found predominantly in mid littoral habitats of Willapa Bay and there is no evidence of resident sublittoral populations. The discrepancy intimates the significance of other factors, including interspecific interactions, that are not incorporated into the model but nonetheless increase metabolic demand. Agonistic encounters with native Dungeness crabs (Cancer magister) may be chief among these additional costs, andC. maenas may largely avoid interactions by remaining in littoral habitats neglected by native crabs, such as meadows of nonindigenous smooth cordgrass (Spartina alterniflora). AdultC. maenas in Willapa Bay may occupy tidal elevations that minimize such encounters, and metabolic costs, while simultaneously maximizing submersion time and foraging opportunities.

Ecological Effects of the Harvest Phase Of Geoduck (Panopea generosa Gould, 1850) Aquaculture on Infaunal Communities in Southern Puget Sound, Washington

ABSTRACT Intertidal aquaculture for geoducks (Panopea generosa Gould, 1850) is expanding in southern Puget Sound, Washington, where gently sloping sandy beaches are used for field culture. Geoduck aquaculture contributes significantly to the regional economy, but has become controversial because of a range of unresolved questions involving potential biological impacts on marine ecosystems. From 2008 through 2012, the authors used a “before-after-control-impact” experimental design, emphasizing spatial scales comparable with those used by geoduck culturists to evaluate the effects of harvesting market-ready geoducks on associated benthic infaunal communities. Infauna were sampled at three different study locations in southern Puget Sound at monthly intervals before, during, and after harvests of clams, and along extralimital transects extending away from the edges of cultured plots to assess the effects of harvest activities in adjacent uncultured habitat. Using multivariate statistical approaches, strong seasonal and spatial signals in patterns of abundance were found, but there was scant evidence of effects on the community structure associated with geoduck harvest disturbances within cultured plots. Likewise, no indications of significant “spillover” effects of harvest on uncultured habitat adjacent to cultured plots were noted. Complementary univariate approaches revealed little evidence of harvest effects on infaunal biodiversity and indications of modest effects on populations of individual infaunal taxa. Of 10 common taxa analyzed, only three showed evidence of reduced densities, although minor, after harvests whereas the remaining seven taxa indicated either neutral responses to harvest disturbances or increased abundance either during or in the months after harvest events. It is suggested that a relatively active natural disturbance regime, including both small-scale and large-scale events that occur with comparable intensity but more frequently than geoduck harvest events in cultured plots, has facilitated assemblage-level infaunal resistance and resilience to harvest disturbances.

Effects of Geoduck (Panopea generosa Gould, 1850) Aquaculture Gear on Resident and Transient Macrofauna Communities of Puget Sound, Washington

ABSTRACT In Washington state, commercial culture of geoducks (Panopea generosa) involves large-scale out-planting of juveniles to intertidal habitats, and installation of PVC tubes and netting to exclude predators and increase early survival. Structures associated with this nascent aquaculture method are examined to determine whether they affect patterns of use by resident and transient macrofauna. Results are summarized from regular surveys of aquaculture operations and reference beaches in 2009 to 2011 at three sites during three phases of culture: (1) pregear (-geoducks, -structure), (2) gear present (+geoducks, +structures), and (3) postgear (+geoducks, -structures). Resident macroinvertebrates (infauna and epifauna) were sampled monthly (in most cases) using coring methods at low tide during all three phases. Differences in community composition between culture plots and reference areas were examined with permutational analysis of variance and homogeneity of multivariate dispersion tests. Scuba and shoreline transect surveys were used to examine habitat use by transient fish and macroinvertebrates. Analysis of similarity and complementary nonmetric multidimensional scaling were used to compare differences between species functional groups and habitat type during different aquaculture phases. Results suggest that resident and transient macrofauna respond differently to structures associated with geoduck aquaculture. No consistent differences in the community of resident macrofauna were observed at culture plots or reference areas at the three sites during any year. Conversely, total abundance of transient fish and macroinvertebrates were more than two times greater at culture plots than reference areas when aquaculture structures were in place. Community composition differed (analysis of similarity) between culture and reference plots during the gear-present phase, but did not persist to the next farming stage (postgear). Habitat complexity associated with shellfish aquaculture may attract some structure-associated transient species observed infrequently on reference beaches, and may displace other species that typically occur in areas lacking epibenthic structure. This study provides a first look at the effects of multiple phases of geoduck farming on macrofauna, and has important implications for the management of a rapidly expanding sector of the aquaculture industry.

The Impact of Yellow Shore Crabs, Hemigrapsus oregonensis, on Early Benthic Phase Dungeness Crabs, Cancer magister, in Intertidal Oyster Shell Mitigation Habitat

Dungeness crab (Cancer magister) megalopae recruit to northeastern Pacific coastal estuaries, and settle into intertidal and subtidal habitats where they molt into Early Benthic Phase (EBP) crabs, and are dependent on epibenthic structure for shelter from predation. Given the importance of shell refuge to their post-settlement ontogeny, the U.S. Army Corps of Engineers began constructing intertidal plots of oyster shell in the Grays Harbor estuary, Washington, to enhance recruitment and mitigate losses of subtidalC. magister entrained and killed during extensive dredging efforts. When shell habitat was newly constructed, settlement and survival ofC. magister were high, and expectations for the mitigation project were met. During the first several years (1992-1997), plots greater than 1 yr postconstruction were colonized by yellow shore crabs,Hemigrapsus oregonensis (often≥75 crabs m−2, and abundance of EBPC. magister was reduced to nearly zero. While some predation of settling megalopae byH. oregonensis does occur, the relationship between these species is characterized by density-dependent competitive interactions. Laboratory observations of competition for shell habitat indicate thatH. oregonensis are dominant over EBPC. magister and can evictC. magister from refuge spaces. Field experiments show that high densities of the former cause, the latter to emigrate from shell, and suggest detection and avoidance of areas with high densities ofH. oregonensis by settlingC. magister megalopae. More recently (1998-2001), abundance ofH. oregonensis has declined dramatically within plots of oyster shell, apparently due to recruitment failure, and patterns ofC. magister abundance and production have returned to levels consistent with original expectations of the mitigation project. Both intraspecific and interspecific competition for space are significant factors effecting population, regulation ofC. magister when they are strongly dependent on refuge from predation. Efficacy of constructed oyster shell plots as a mitigation tool may hinge on the population dynamics of a species previously considered of little consequence to the target species.

Aquaculture Disturbance Impacts the Diet but not Ecological Linkages of a Ubiquitous Predatory Fish

Aquaculture operations are a frequent and prominent cause of anthropogenic disturbance to marine and estuarine communities and may alter species composition and abundance. However, little is known about how such disturbances affect trophic linkages or ecosystem functions. In Puget Sound, Washington, aquaculture of the Pacific geoduck clam (Panopea generosa) is increasing and involves placing nets and polyvinyl chloride (PVC) tubes in intertidal areas to protect juvenile geoducks from predators. Initial studies of the structured phase of the farming cycle have documented limited impacts on the abundance of some species. To examine the effect of geoduck aquaculture on ecological linkages, the trophic relationships of a local ubiquitous consumer, Pacific staghorn sculpin (Leptocottus armatus), to its invertebrate prey were compared between geoduck aquaculture sites and nearby reference areas with no aquaculture. Mark-recapture data indicated that sculpin exhibit local site fidelity to cultured and reference areas. The stomach contents of sculpin and stable isotope signatures of sculpin and their prey were examined to study the trophic ecology of cultured and reference areas. Results showed that the structured phase of geoduck aquaculture initiated some changes to staghorn sculpin ecology, as reflected in sculpin diet through stomach content analysis. However, carbon and nitrogen stable isotopes revealed that the general food web function of sculpin remained unchanged. The source of carbon at the base of the food web and the trophic position of sculpin were not impacted by geoduck aquaculture. The study has important implications for geoduck aquaculture management and will inform regulatory decisions related to shellfish aquaculture policy.

Distribution, Abundance, and Habitat Associations of a Large Bivalve (Panopea generosa) in a Eutrophic Fjord Estuary

ABSTRACT Marine bivalves are important ecosystem constituents and frequently support valuable fisheries. In many nearshore areas, human disturbance—including declining habitat and water quality—can affect the distribution and abundance of bivalve populations, and complicate ecosystem and fishery management assessments. Infaunal bivalves, in particular, are frequently cryptic and difficult to detect; thus, assessing potential impacts on their populations requires suitable, scalable methods for estimating abundance and distribution. In this study, population size of a common benthic bivalve (the geoduck Panopea generosa) is estimated with a Bayesian habitat-based model fit to scuba and tethered camera data in Hood Canal, a fjord basin in Washington state. Densities declined more than two orders of magnitude along a north—south gradient, concomitant with patterns of deepwater dissolved oxygen, and intensity and duration of seasonal hypoxia. Across the basin, geoducks were most abundant in loose, unconsolidated, sand substrate. The current study demonstrates the utility of using scuba, tethered video, and habitat models to estimate the abundance and distribution of a large infaunal bivalve at a regional (385-km2) scale.