As in other eutrophied estuaries and coastal embayments, persistent hypoxia now routinely develops during summer in the mesohaline portion of the Neuse River estuary (North Carolina, USA). In response to interannual differences in hydrography, summer 1997 exhibited much more intense and widespread hypoxia than summer 1998, permitting inferences about impacts of hypoxia on food web dynamics by comparing system changes across these two summers. The trophic structure of the Neuse estuary now resembles the generic pattern for a degraded temperate estuary with (1) intense planktonic algal blooms and similarly high production of free-living bacteria, (2) trivial levels of abundance of rooted aquatic plants and benthic macroalgae, (3) depleted apex predators, and (4) functional extinction of the historically dominant benthic grazer, eastern oysters. Detailed carbon-flow models, based on comprehensive field data, demonstrated large differences between the two summers in trophic transfers and system dynamics. Largely because of greater mortality of benthic invertebrates from more intense hypoxia, total biomass of heterotrophs declined over summer by 51% in 1997 as compared to only 17% in 1998. Because net primary production increased over summer and herbivory in this system is predominantly benthic, the fraction of primary production consumed by herbivores declined over summer by 35% in 1997 and 29% in 1998. Influx of juvenile fishes and their rapid growth in the estuarine nursery over summer led to increases in energy demand by demersal fishes of 380% and 507% in the successive summers. Thus, hypoxia-enhanced diversion of energy flows into microbial pathways away from consumers and mass mortality of benthic invertebrates from bottom hypoxia occurred at the season of greatest demand by predatory fishes and crabs using the estuary as nursery. Average residence time of carbon in the ecosystem declined by 51% in 1997 and 29% in 1998. Total system throughput declined over summer 1997 while increasing in 1998, indicating the reduced capacity of the system to transfer carbon to higher trophic levels in the more hypoxic summer. Late-summer trophic pathways were characterized by greater numbers of cycles, but flows became increasingly dominated by microbial loops rather than transfers to consumers. Ecosystem trophic efficiency was only ϳ4%, lower than other estuaries similarly analyzed. System properties indicative of resiliency of system function including development capacity, ascendancy, and flow diversity declined over summer 1997, while increasing or declining less in 1998. Thus, intensification of hypoxia caused dramatic reduction in the ecosystem's ability to transfer energy to higher trophic levels and rendered the ecosystem potentially less resilient to other stressors.

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The structure and dynamics of natural communities result from the interplay of abiotic and biotic factors. We used manipulative field experiments to determine the relative roles of abiotic conditions and biotic interactions in structuring deep-sea (2500 m depth) communities along environmental gradients around hydrothermal vents of the eastern tropical Pacific Ocean (East Pacific Rise, at 9Њ50Ј N). We tested (1) whether predation by crabs and fishes affects the recruitment of benthic species and subsequent community structure and (2) whether the effects of predation vary along the steep gradients of temperature, oxygen, sulfide, and metal concentrations near vents. Recruitment substrates (basalt cubic blocks, roughly 10 cm on a side), both uncaged and caged to exclude predators (crabs, fishes, whelks, and octopi), were deployed along a decreasing vent fluid-flux gradient. The exclusion of predators for 8 mo increased the abundance of small mobile gastropods and amphipod crustaceans but decreased the abundance of sessile invertebrates, including juvenile vestimentiferan worms, tubiculous polychaetes, and mussels. Effects of predation were strongest nearest to hydrothermal vents, where abiotic environmental conditions were most extreme but productivity and the overall abundances of benthic invertebrates and mobile predators were the greatest. Additional 5-mo experiments conducted at three different locations showed similar trends at all sites, indicating that these effects of predation on benthic community structure are repeatable. Stomach-content analyses of the most abundant predators found at vents indicated that the zoarcid fish (Thermarces cerberus) primarily feeds on the vent snail Cyathermia naticoides, the limpet Lepetodrilus elevatus, and the amphipod crustacean Ventiella sulfuris, the very species that showed the greatest increase following predator exclusion. In contrast, brachyuran (Bythograea thermydron) and galatheid (Munidopsis subsquamosa) crab stomachs did not contain small mobile grazers, and crabs presented with arrays of the most common vent invertebrate species preferred mussels and vestimentiferans over limpets. Our results indicate that predation by large mobile predators influences the structure of hydrothermal vent communities, directly by reducing the abundance of gastropod prey species, and indirectly by reducing gastropod grazing and by bulldozing of recruits of sessile invertebrates.

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The structure and dynamics of natural communities result from the interplay of abiotic and biotic factors. We used manipulative field experiments to determine the relative roles of abiotic conditions and biotic interactions in structuring deep‐sea (2500 m depth) communities along environmental gradients around hydrothermal vents of the eastern tropical Pacific Ocean (East Pacific Rise, at 9°50′ N). We tested (1) whether predation by crabs and fishes affects the recruitment of benthic species and subsequent community structure and (2) whether the effects of predation vary along the steep gradients of temperature, oxygen, sulfide, and metal concentrations near vents. Recruitment substrates (basalt cubic blocks, roughly 10 cm on a side), both uncaged and caged to exclude predators (crabs, fishes, whelks, and octopi), were deployed along a decreasing vent fluid‐flux gradient. The exclusion of predators for 8 mo increased the abundance of small mobile gastropods and amphipod crustaceans but decreased the abundance of sessile invertebrates, including juvenile vestimentiferan worms, tubiculous polychaetes, and mussels. Effects of predation were strongest nearest to hydrothermal vents, where abiotic environmental conditions were most extreme but productivity and the overall abundances of benthic invertebrates and mobile predators were the greatest. Additional 5‐mo experiments conducted at three different locations showed similar trends at all sites, indicating that these effects of predation on benthic community structure are repeatable. Stomach‐content analyses of the most abundant predators found at vents indicated that the zoarcid fish (Thermarces cerberus) primarily feeds on the vent snail Cyathermia naticoides, the limpet Lepetodrilus elevatus, and the amphipod crustacean Ventiella sulfuris, the very species that showed the greatest increase following predator exclusion. In contrast, brachyuran (Bythograea thermydron) and galatheid (Munidopsis subsquamosa) crab stomachs did not contain small mobile grazers, and crabs presented with arrays of the most common vent invertebrate species preferred mussels and vestimentiferans over limpets. Our results indicate that predation by large mobile predators influences the structure of hydrothermal vent communities, directly by reducing the abundance of gastropod prey species, and indirectly by reducing gastropod grazing and by bulldozing of recruits of sessile invertebrates.

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