Artificial habitats host elevated densities of large reef-associated predators.

Large predators play important ecological roles, yet many are disproportionately imperiled. In marine systems, artificial reefs are often deployed to restore degraded reefs or supplement existing reefs, but it remains unknown whether these interventions benefit large predators. Comparative field surveys of thirty artificial and natural reefs across ~200 km of the North Carolina, USA coast revealed large reef-associated predators were more dense on artificial than natural reefs. This pattern was associated with higher densities of transient predators (e.g. jacks, mackerel, barracuda, sharks) on artificial reefs, but not of resident predators (e.g., grouper, snapper). Further analyses revealed that this pattern of higher transient predator densities on artificial reefs related to reef morphology, as artificial reefs composed of ships hosted higher transient predator densities than concrete reefs. The strength of the positive association between artificial reefs and transient predators increased with a fundamental habitat trait-vertical extent. Taller artificial reefs had higher densities of transient predators, even when accounting for habitat area. A global literature review of high trophic level fishes on artificial and natural habitats suggests that the overall pattern of more predators on artificial habitats is generalizable. Together, these findings provide evidence that artificial habitats, especially those like sunken ships that provide high vertical structure, may support large predators.

Artificial reefs facilitate tropical fish at their range edge.

Spatial planning increasingly incorporates theoretical predictions that artificial habitats assist species movement at or beyond range edges, yet evidence for this is uncommon. We conducted surveys of highly mobile fauna (fishes) on artificial habitats (reefs) on the southeastern USA continental shelf to test whether, in comparison to natural reefs, artificial reefs enhance local abundance and biomass of fishes at their poleward range margins. Here, we show that while temperate fishes were more abundant on natural reefs, tropical, and subtropical fishes exhibited higher abundances and biomasses on deep (25-35 m) artificial reefs. Further analyses reveal that this effect depended on feeding guilds because planktivorous and piscivorous but not herbivorous fishes were more abundant on artificial reefs. This is potentially due to heightened prey availability on and structural complexity of artificial reefs. Our findings demonstrate that artificial habitats can facilitate highly mobile species at range edges and suggest these habitats assist poleward species movement.

Living shorelines enhanced the resilience of saltmarshes to Hurricane Matthew (2016).

Nature-based solutions, such as living shorelines, have the potential to restore critical ecosystems, enhance coastal sustainability, and increase resilience to natural disasters; however, their efficacy during storm events compared to traditional hardened shorelines is largely untested. This is a major impediment to their implementation and promotion to policy-makers and homeowners. To address this knowledge gap, we evaluated rock sill living shorelines as compared to natural marshes and hardened shorelines (i.e., bulkheads) in North Carolina, USA for changes in surface elevation, Spartina alterniflora stem density, and structural damage from 2015 to 2017, including before and after Hurricane Matthew (2016). Our results show that living shorelines exhibited better resistance to landward erosion during Hurricane Matthew than bulkheads and natural marshes. Additionally, living shorelines were more resilient than hardened shorelines, as they maintained landward elevation over the two-year study period without requiring any repair. Finally, rock sill living shorelines were able to enhance S. alterniflora stem densities over time when compared to natural marshes. Our results suggest that living shorelines have the potential to improve coastal resilience while supporting important coastal ecosystems.

Flat and complex temperate reefs provide similar support for fish: Evidence for a unimodal species-habitat relationship.

Structural complexity, a form of habitat heterogeneity, influences the structure and function of ecological communities, generally supporting increased species density, richness, and diversity. Recent research, however, suggests the most complex habitats may not harbor the highest density of individuals and number of species, especially in areas with elevated human influence. Understanding nuances in relationships between habitat heterogeneity and ecological communities is warranted to guide habitat-focused conservation and management efforts. We conducted fish and structural habitat surveys of thirty warm-temperate reefs on the southeastern US continental shelf to quantify how structural complexity influences fish communities. We found that intermediate complexity maximizes fish abundance on natural and artificial reefs, as well as species richness on natural reefs, challenging the current paradigm that abundance and other fish community metrics increase with increasing complexity. Naturally occurring rocky reefs of flat and complex morphologies supported equivalent abundance, biomass, species richness, and community composition of fishes. For flat and complex morphologies of rocky reefs to receive equal consideration as essential fish habitat (EFH), special attention should be given to detecting pavement type rocky reefs because their ephemeral nature makes them difficult to detect with typical seafloor mapping methods. Artificial reefs of intermediate complexity also maximized fish abundance, but human-made structures composed of low-lying concrete and metal ships differed in community types, with less complex, concrete structures supporting lower numbers of fishes classified largely as demersal species and metal ships protruding into the water column harboring higher numbers of fishes, including more pelagic species. Results of this study are essential to the process of evaluating habitat function provided by different types and shapes of reefs on the seafloor so that all EFH across a wide range of habitat complexity may be accurately identified and properly managed.

Oyster reefs as carbon sources and sinks.

Carbon burial is increasingly valued as a service provided by threatened vegetated coastal habitats. Similarly, shellfish reefs contain significant pools of carbon and are globally endangered, yet considerable uncertainty remains regarding shellfish reefs' role as sources (+) or sinks (-) of atmospheric CO2 While CO2 release is a by-product of carbonate shell production (then burial), shellfish also facilitate atmospheric-CO2 drawdown via filtration and rapid biodeposition of carbon-fixing primary producers. We provide a framework to account for the dual burial of inorganic and organic carbon, and demonstrate that decade-old experimental reefs on intertidal sandflats were net sources of CO2 (7.1 ± 1.2 MgC ha-1 yr-1 (µ ± s.e.)) resulting from predominantly carbonate deposition, whereas shallow subtidal reefs (-1.0 ± 0.4 MgC ha-1 yr-1) and saltmarsh-fringing reefs (-1.3 ± 0.4 MgC ha-1 yr-1) were dominated by organic-carbon-rich sediments and functioned as net carbon sinks (on par with vegetated coastal habitats). These landscape-level differences reflect gradients in shellfish growth, survivorship and shell bioerosion. Notably, down-core carbon concentrations in 100- to 4000-year-old reefs mirrored experimental-reef data, suggesting our results are relevant over centennial to millennial scales, although we note that these natural reefs appeared to function as slight carbon sources (0.5 ± 0.3 MgC ha-1 yr-1). Globally, the historical mining of the top metre of shellfish reefs may have reintroduced more than 400 000 000 Mg of organic carbon into estuaries. Importantly, reef formation and destruction do not have reciprocal, counterbalancing impacts on atmospheric CO2 since excavated organic material may be remineralized while shell may experience continued preservation through reburial. Thus, protection of existing reefs could be considered as one component of climate mitigation programmes focused on the coastal zone.

Anomalously High Recruitment of the 2010 Gulf Menhaden (Brevoortia patronus) Year Class: Evidence of Indirect Effects from the Deepwater Horizon Blowout in the Gulf of Mexico.

Gulf menhaden (Brevoortia patronus) exhibited unprecedented juvenile recruitment in 2010 during the year of the Deepwater Horizon well blowout, exceeding the prior 39-year mean by more than four standard deviations near the Mississippi River. Abundance of that cohort remained exceptionally high for two subsequent years as recruits moved into older age classes. Such changes in this dominant forage fish population can be most parsimoniously explained as consequences of release from predation. Contact with crude oil induced high mortality of piscivorous seabirds, bottlenose dolphin (Tursiops truncatus), waders, and other fish-eating marsh birds, all of which are substantial consumers of Gulf menhaden. Diversions of fresh water from the Mississippi River to protect coastal marshes from oiling depressed salinities, impairing access to juvenile Gulf menhaden by aquatic predators that avoid low-salinity estuarine waters. These releases from predation led to an increase of Gulf menhaden biomass in 2011 to 2.4 million t, or more than twice the average biomass of 1.1 million t for the decade prior to 2010. Biomass increases of this magnitude in a major forage fish species suggest additional trophically linked effects at the population-, trophic-level and ecosystem scales, reflecting an heretofore little appreciated indirect effect that may be associated with major oil spills in highly productive marine waters.

Living shorelines can enhance the nursery role of threatened estuarine habitats.

Coastal ecosystems provide numerous services, such as nutrient cycling, climate change amelioration, and habitat provision for commercially valuable organisms. Ecosystem functions and processes are modified by human activities locally and globally, with degradation of coastal ecosystems by development and climate change occurring at unprecedented rates. The demand for coastal defense strategies against storms and sea-level rise has increased with human population growth and development along coastlines world-wide, even while that population growth has reduced natural buffering of shorelines. Shoreline hardening, a common coastal defense strategy that includes the use of seawalls and bulkheads (vertical walls constructed of concrete, wood, vinyl, or steel), is resulting in a "coastal squeeze" on estuarine habitats. In contrast to hardening, living shorelines, which range from vegetation plantings to a combination of hard structures and plantings, can be deployed to restore or enhance multiple ecosystem services normally delivered by naturally vegetated shores. Although hundreds of living shoreline projects have been implemented in the United States alone, few studies have evaluated their effectiveness in sustaining or enhancing ecosystem services relative to naturally vegetated shorelines and hardened shorelines. We quantified the effectiveness of (1) sills with landward marsh (a type of living shoreline that combines marsh plantings with an offshore low-profile breakwater), (2) natural salt marsh shorelines (control marshes), and (3) unvegetated bulkheaded shores in providing habitat for fish and crustaceans (nekton). Sills supported higher abundances and species diversity of fishes than unvegetated habitat adjacent to bulkheads, and even control marshes. Sills also supported higher cover of filter-feeding bivalves (a food resource and refuge habitat for nekton) than bulkheads or control marshes. These ecosystem-service enhancements were detected on shores with sills three or more years after construction, but not before. Sills provide added structure and may provide better refuges from predation and greater opportunity to use available food resources for nekton than unvegetated bulkheaded shores or control marshes. Our study shows that unlike shoreline hardening, living shorelines can enhance some ecosystem services provided by marshes, such as provision of nursery habitat.

Metrics to assess ecological condition, change, and impacts in sandy beach ecosystems.

Complexity is increasingly the hallmark in environmental management practices of sandy shorelines. This arises primarily from meeting growing public demands (e.g., real estate, recreation) whilst reconciling economic demands with expectations of coastal users who have modern conservation ethics. Ideally, shoreline management is underpinned by empirical data, but selecting ecologically-meaningful metrics to accurately measure the condition of systems, and the ecological effects of human activities, is a complex task. Here we construct a framework for metric selection, considering six categories of issues that authorities commonly address: erosion; habitat loss; recreation; fishing; pollution (litter and chemical contaminants); and wildlife conservation. Possible metrics were scored in terms of their ability to reflect environmental change, and against criteria that are widely used for judging the performance of ecological indicators (i.e., sensitivity, practicability, costs, and public appeal). From this analysis, four types of broadly applicable metrics that also performed very well against the indicator criteria emerged: 1.) traits of bird populations and assemblages (e.g., abundance, diversity, distributions, habitat use); 2.) breeding/reproductive performance sensu lato (especially relevant for birds and turtles nesting on beaches and in dunes, but equally applicable to invertebrates and plants); 3.) population parameters and distributions of vertebrates associated primarily with dunes and the supralittoral beach zone (traditionally focused on birds and turtles, but expandable to mammals); 4.) compound measurements of the abundance/cover/biomass of biota (plants, invertebrates, vertebrates) at both the population and assemblage level. Local constraints (i.e., the absence of birds in highly degraded urban settings or lack of dunes on bluff-backed beaches) and particular issues may require alternatives. Metrics - if selected and applied correctly - provide empirical evidence of environmental condition and change, but often do not reflect deeper environmental values per se. Yet, values remain poorly articulated for many beach systems; this calls for a comprehensive identification of environmental values and the development of targeted programs to conserve these values on sandy shorelines globally.

Multi-year persistence of beach habitat degradation from nourishment using coarse shelly sediments.

Beach nourishment is increasingly used to protect public beach amenity and coastal property from erosion and storm damage. Where beach nourishment uses fill sediments that differ in sedimentology from native beach sands, press disturbances to sandy beach invertebrates and their ecosystem services can occur. How long impacts persist is, however, unclear because monitoring after nourishment typically only extends for several months. Here, monitoring was extended for 3-4 years following each of two spatially separated, replicate nourishment projects using unnaturally coarse sediments. Following both fill events, the contribution to beach sediments of gravel-sized particles and shell fragments was enhanced, and although diminishing through time, remained elevated as compared to control sites at the end of 3-4 years of monitoring, including in the low intertidal and swash zones, where benthic macroinvertebrates concentrate. Consequently, two infaunal invertebrates, haustoriid amphipods and Donax spp., exhibited suppressed densities over the entire post-nourishment period of 3-4 years. Emerita talpoida, by contrast, exhibited lower densities on nourished than control beaches only in the early summer of the first and second years and polychaetes exhibited little response to nourishment. The overall impact to invertebrates of nourishment was matched by multi-year reductions in abundances of their predators. Ghost crab abundances were suppressed on nourished beaches with impacts disappearing only by the fourth summer. Counts of foraging shorebirds were depressed for 4 years after the first project and 2 years after the second project. Our results challenge the view that beach nourishment is environmentally benign by demonstrating that application of unnaturally coarse and shelly sediments can serve as a press disturbance to degrade the beach habitat and its trophic services to shorebirds for 2-4 years. Recognizing that recovery following nourishment can be slow, studies that monitor impacts for only several months are inadequate.

Integrating the invisible fabric of nature into fisheries management.

Overfishing and environmental change have triggered many severe and unexpected consequences. As existing communities have collapsed, new ones have become established, fundamentally transforming ecosystems to those that are often less productive for fisheries, more prone to cycles of booms and busts, and thus less manageable. We contend that the failure of fisheries science and management to anticipate these transformations results from a lack of appreciation for the nature, strength, complexity, and outcome of species interactions. Ecologists have come to understand that networks of interacting species exhibit nonlinear dynamics and feedback loops that can produce sudden and unexpected shifts. We argue that fisheries science and management must follow this lead by developing a sharper focus on species interactions and how disrupting these interactions can push ecosystems in which fisheries are embedded past their tipping points.

Prey distribution, physical habitat features, and guild traits interact to produce contrasting shorebird assemblages among foraging patches.

Worldwide declines in shorebird populations, driven largely by habitat loss and degradation, motivate environmental managers to preserve and restore the critical coastal habitats on which these birds depend. Effective habitat management requires an understanding of the factors that determine habitat use and value to shorebirds, extending from individuals to the entire community. While investigating the factors that influenced shorebird foraging distributions among neighboring intertidal sand flats, we built upon species-level understandings of individual-based, small-scale foraging decisions to develop more comprehensive guild- and community-level insights. We found that densities and community composition of foraging shorebirds varied substantially among elevations within some tidal flats and among five flats despite their proximity (all located within a 400-m stretch of natural, unmodified inlet shoreline). Non-dimensional multivariate analyses revealed that the changing composition of the shorebird community among flats and tidal elevations correlated significantly (ρ(s) = 0.56) with the spatial structure of the benthic invertebrate prey community. Sediment grain-sizes affected shorebird community spatial patterns indirectly by influencing benthic macroinvertebrate community compositions. Furthermore, combining sediment and macroinvertebrate information produced a 27% increase in correlation (ρ(s) = 0.71) with shorebird assemblage patterns over the correlation of the bird community with the macroinvertebrate community alone. Beyond its indirect effects acting through prey distributions, granulometry of the flats influenced shorebird foraging directly by modifying prey availability. Our study highlights the importance of habitat heterogeneity, showing that no single patch type was ideal for the entire shorebird community. Generally, shorebird density and diversity were greatest at lower elevations on flats when they became exposed; these areas are at risk from human intervention by inlet sand mining, construction of groins and jetties that divert sediments from flats, and installation of seawalls on inlet shorelines that induce erosion of flats.

Imprint of past environmental regimes on structure and succession of a deep-sea hydrothermal vent community.

Dramatic perturbations of ecological communities through rapid shifts in environmental regime do not always result in complete mortality of residents. Instead, legacy individuals may remain and influence the succession and composition of subsequent communities. We used a reciprocal transplant experiment to investigate whether a legacy effect is detectable in communities experiencing an abrupt increase or decrease in hydrothermal fluid flux at deep-sea vents. Vent habitats are characterized by strong gradients in productivity and physico-chemical stressors, both of which tend to increase with increasing vent fluid flux. In our experiments, many species survived transplantation from cool (water temperatures <2 degrees C above ambient) to warm (4-30 degrees C above ambient) habitats, resulting in significantly higher species richness on transplanted than remaining experimental substrata. A legacy effect was much less apparent in transplantation from warm to cool habitat, although a few vestimentiferan tubeworms, normally restricted to warm habitat, survived transplantation. The asymmetry in influence of legacy individuals suggests that productivity enhancement may outweigh potential physiological stress in setting limits to distributions of vent invertebrates. This influence of biological processes contrasts with theory developed in the rocky intertidal that predicts the predominance of physical control at the high-stress end of an environmental gradient. Prediction of successional transitions in vents and other habitats experiencing regime shifts in which remnant species may survive must take into account the possible influence of historical effects.

Practical proxies for tidal marsh ecosystem services: application to injury and restoration.

Tidal marshes are valued, protected and restored in recognition of their ecosystem services: (1) high productivity and habitat provision supporting the food web leading to fish and wildlife, (2) buffer against storm wave damage, (3) shoreline stabilization, (4) flood water storage, (5) water quality maintenance, (6) biodiversity preservation, (7) carbon storage and (8) socio-economic benefits. Under US law, federal and state governments have joint responsibility for facilitating restoration to compensate quantitatively for ecosystem services lost because of oil spills and other contaminant releases on tidal marshes. This responsibility is now met by choosing and employing metrics (proxies) for the suite of ecosystem services to quantify injury and scale restoration accordingly. Most injury assessments in tidal marshes are triggered by oil spills and are limited to: (1) documenting areas covered by heavy, moderate and light oiling; (2) estimating immediate above-ground production loss (based on stem density and height) of the dominant vascular plants within each oiling intensity category and (3) sampling sediments for chemical analyses and depth of contamination, followed by sediment toxicity assays if sediment contamination is high and likely to persist. The percentage of immediate loss of ecosystem services is then estimated along with the recovery trajectory. Here, we review potential metrics that might refine or replace present metrics for marsh injury assessment. Stratifying plant sampling by the more productive marsh edge versus the less accessible interior would improve resolution of injury and provide greater confidence that restoration is truly compensatory. Using microphytobenthos abundance, cotton-strip decomposition bioassays and other biogeochemical indicators, or sum of production across consumer trophic levels fails as a stand-alone substitute metric. Below-ground plant biomass holds promise as a potential proxy for resiliency but requires further testing. Under some conditions, like chronic contamination by organic pollutants that affect animals but not vascular plants, benthic infaunal density, toxicity testing, and tissue contamination, growth, reproduction and mortality of marsh vertebrates deserve inclusion in the assessment protocol. Additional metrics are sometimes justified to assay microphytobenthos, use by nekton, food and habitat for reptiles, birds and mammals, or support of plant diversity. Empirical research on recovery trajectories in previously injured marshes could reduce the largest source of uncertainty in quantifying cumulative service losses.

Interacting effects of temperature and salinity on Bonamia sp. parasitism in the Asian oyster Crassostrea ariakensis.

The proposition to introduce the Asian oyster Crassostrea ariakensis to the mid-Atlantic region of the USA is being considered with caution, particularly after the discovery of a novel microcell haplosporidian parasite, Bonamia sp., in North Carolina. Although this parasite was found to be pathogenic in C. ariakensis under warm euhaline conditions, its persistence in C. ariakensis exposed to various temperature and salinity combinations remained unresolved. In this laboratory experiment, we tested the influence of temperature in combination with a wide range of salinities (10, 20 and 30 psu) on Bonamia sp. Temperature was either changed from warm (>20 degrees C) to cold (6 degrees C for 6 weeks) and back to warm or maintained constant and warm. Warm temperature was associated with higher host mortality than cold temperature, suggesting that temperature influenced Bonamia sp. pathogenicity. The effect of salinity was revealed under warm temperature with highest mortality levels observed in infected C. ariakensis exposed to 30 psu. When temperature was increased following low-temperature exposure, Bonamia sp. was not detected; however sub-optimal experimental conditions may have contributed to this result, making it difficult to draw conclusions regarding the reemergence of the parasite after low-temperature exposure. Although the overwintering of Bonamia sp. in C. ariakensis will need to be further investigated, the results presented here suggest that Bonamia sp. may be able to persist in C. ariakensis under a combination of low temperature and meso- to euhaline salinities.

Strong seasonality of Bonamia sp. infection and induced Crassostrea ariakensis mortality in Bogue and Masonboro Sounds, North Carolina, USA.

Asian oyster Crassostrea ariakensis is being considered for introduction to Atlantic coastal waters of the USA. Successful aquaculture of this species will depend partly on mitigating impacts by Bonamia sp., a parasite that has caused high C. ariakensis mortality south of Virginia. To better understand the biology of this parasite and identify strategies for management, we evaluated its seasonal pattern of infection in C. ariakensis at two North Carolina, USA, locations in 2005. Small (<50 mm) triploid C. ariakensis were deployed to upwellers on Bogue Sound in late spring (May), summer (July), early fall (September), late fall (November), and early winter (December) 2005; and two field sites on Masonboro Sound in September 2005. Oyster growth and mortality were evaluated biweekly at Bogue Sound, and weekly at Masonboro, with Bonamia sp. prevalence evaluated using parasite-specific PCR. We used histology to confirm infections in PCR-positive oysters. Bonamia sp. appeared in the late spring Bogue Sound deployment when temperatures approached 25 degrees C, six weeks post-deployment. Summer- and early fall-deployed oysters displayed Bonamia sp. infections after 3-4 weeks. Bonamia sp. prevalences were 75% in Bogue Sound, and 60% in Masonboro. While oyster mortality reached 100% in late spring and summer deployments, early fall deployments showed reduced (17-82%) mortality. Late fall and early winter deployments, made at temperatures <20 degrees C, developed no Bonamia sp. infections at all. Seasonal Bonamia sp. cycling, therefore, is influenced greatly by temperature. Avoiding peak seasonal Bonamia sp. activity will be essential for culturing C. ariakensis in Bonamia sp.-enzootic waters.

Cascading effects of the loss of apex predatory sharks from a coastal ocean.

Impacts of chronic overfishing are evident in population depletions worldwide, yet indirect ecosystem effects induced by predator removal from oceanic food webs remain unpredictable. As abundances of all 11 great sharks that consume other elasmobranchs (rays, skates, and small sharks) fell over the past 35 years, 12 of 14 of these prey species increased in coastal northwest Atlantic ecosystems. Effects of this community restructuring have cascaded downward from the cownose ray, whose enhanced predation on its bay scallop prey was sufficient to terminate a century-long scallop fishery. Analogous top-down effects may be a predictable consequence of eliminating entire functional groups of predators.

Bonamia perspora n. sp. (Haplosporidia), a parasite of the oyster Ostreola equestris, is the first Bonamia species known to produce spores.

Examination of the oyster Ostreola equestris as a potential reservoir host for a species of Bonamia discovered in Crassostrea ariakensis in North Carolina (NC), USA, revealed a second novel Bonamia sp. Histopathology, electron microscopy, and molecular phylogenetic analysis support the designation of a new parasite species, Bonamia perspora n. sp., which is the first Bonamia species shown to produce a typical haplosporidian spore with an orifice and hinged operculum. Spores were confirmed to be from B. perspora by fluorescent in situ hybridization. Bonamia perspora was found at Morehead City and Wilmington, NC, with an overall prevalence of 1.4% (31/2,144). Uninucleate, plasmodial, and sporogonic stages occurred almost exclusively in connective tissues; uninucleate stages (2-6 microm) were rarely observed in hemocytes. Spores were 4.3-6.4 microm in length. Ultrastructurally, uninucleate, diplokaryotic, and plasmodial stages resembled those of other spore-forming haplosporidians, but few haplosporosomes were present, and plasmodia were small. Spore ornamentation consisted of spore wall-derived, thin, flat ribbons that emerged haphazardly around the spore, and which terminated in what appeared to be four-pronged caps. Number of ribbons per spore ranged from 15 to 30, and their length ranged from 1.0 to 3.4 microm. Parsimony analysis identified B. perspora as a sister species to Bonamia ostreae.

Depletion, degradation, and recovery potential of estuaries and coastal seas.

Estuarine and coastal transformation is as old as civilization yet has dramatically accelerated over the past 150 to 300 years. Reconstructed time lines, causes, and consequences of change in 12 once diverse and productive estuaries and coastal seas worldwide show similar patterns: Human impacts have depleted >90% of formerly important species, destroyed >65% of seagrass and wetland habitat, degraded water quality, and accelerated species invasions. Twentieth-century conservation efforts achieved partial recovery of upper trophic levels but have so far failed to restore former ecosystem structure and function. Our results provide detailed historical baselines and quantitative targets for ecosystem-based management and marine conservation.

When r-selection may not predict introduced-species proliferation: predation of a nonnative oyster.

Predicting outcomes of species introductions may be enhanced by integrating life-history theory with results of contained experiments that compare ecological responses of exotic and analogue native species to dominant features of the recipient environment. An Asian oyster under consideration for introduction to the Chesapeake Bay, USA, the rapidly growing Suminoe oyster (Crassostrea ariakensis), may not be as successful an invader as its r-selected life history suggests if the trade-off for rapid growth and maturation is lower investment in defenses against blue crab (Callinectes sapidus) predation than the native Eastern oyster (Crassostrea virginica). In laboratory trials, blue crabs simultaneously offered equal numbers of Suminoe and Eastern oysters consumed more nonnatives, irrespective of whether the crabs had previous experience with Suminoe oysters as prey. Satiated blue crabs consumed nearly three times as many Suminoe oysters as Eastern oysters of 25-mm shell height, and eight times as many of 35-mm shell height. Despite blue crabs consuming small (30 mm) Suminoe oysters at twice the rate of large (40 mm) Suminoe oysters, when 40-mm Suminoe were paired with 30-mm Eastern oysters, seven times as many of the larger (Suminoe) oysters were consumed. The greater susceptibility of C. ariakensis than C. virginica to blue crab predation appears to be based upon the biomechanics of shell strength rather than active selection of a more attractive food. Much less force was required to crush shells of Suminoe than Eastern oysters of similar shell height. Tissue transplant experiments demonstrated greater predation on oyster tissues in weaker C. ariakensis shells independent of tissue identity, and duration of handling time before rejection of C. virginica exceeded the time to crush C. ariakensis. These results, coupled with the present importance of blue crab predation in limiting recovery of native Eastern oysters, imply a role for blue crabs in inhibiting Suminoe oysters, if introduced, from attaining high adult densities required to restore a fishery, provide appreciable reef habitat, and reduce turbidity through filtration. Thus, in high-predation environments, allocation of resources to rapid growth and development rather than to predation defenses reflects a life-history trade-off that may promote early stages of invasion, yet prevent attainment of dense adult populations.