The structure of ecological communities reflects a tension among forces that alter populations. Marine ecologists previously emphasized control by locally operating forces (predation, competition, and disturbance), but newer studies suggest that inputs from large-scale oceanographically modulated subsidies (nutrients, particulates, and propagules) can strongly influence community structure and dynamics. On New Zealand rocky shores, the magnitude of such subsidies differs profoundly between contrasting oceanographic regimes. Community structure, and particularly the pace of community dynamics, differ dramatically between intermittent upwelling regimes compared with relatively persistent downwelling regimes. We suggest that subsidy rates are a key determinant of the intensity of species interactions, and thus of structure in marine systems, and perhaps also nonmarine communities. Many ecological processes determine the structure and dynamics of communities and ecosystems. Theoretical and experimental advances have led to a growing awareness that different processes operate at different spatial and temporal scales (1, 2). Capturing the full richness of ecosystem dynamics thus requires studies ranging across a wide range of scales and allowing effective evaluation of the relative impact of the relevant factors. Doing so can be challenging. The logistics are daunting, especially in using the power of experimentation over large spatial scales. One solution, the ''comparative-experimental'' approach (3), involves replicated local-scale experimentation at multiple sites spanning larger scales, coupled with local-scale repeated sampling of factors that vary at characteristically larger scales.In its early development, marine community ecology focused on the dynamic consequences of primarily local-scale processes such as species interactions and physical disturbance (4). More recently, marine and nonmarine ecologists alike have documented the influence on communities of larger-scale phenomena including subsidies of materials and propagules transferred between adjacent ecosystems (5-8). Although evidence for the importance of subsidies is growing, questions remain about their impact, generality, magnitude, and interdependence and the physical and biotic mechanisms that underlie them.Here we use the comparative-experimental approach to address the role of large-scale oceanographic phenomena in structuring communities on New Zealand rocky shores. Based on earlier results (6, 9), we predicted that intertidal community structure and dynamics would reflect the coastal oceanographic regime. We hypothesized that the influences of oceanographically modulated subsidies (propagules, as a cause of increases in population density of benthic species, and the concentration of phytoplankton and detritus, as food for filter feeders) would be high with upwelling and low with downwelling. Study SystemPrevious research on the west and east coasts of the South Island of New Zealand (pairs of sites 100-500 m apart on each coast) revealed strik...

A recent model predicts that species interactions in benthic marine communities vary predictably with upwelling regimes. To test this model, we studied the Pisaster-Mytilus interaction at 14 rocky intertidal sites distributed among three oceanographic regions along a 1300-km stretch of the U.S. West Coast. Regions included an intermittentupwelling region (northern), a persistent-upwelling region (central), and a region of weak and infrequent upwelling (southern). We quantified predation rates by the sea star Pisaster ochraceus on its main prey Mytilus californianus by transplanting mussels into the sea star's low-zone foraging range and comparing the rate of mussel loss in ϩPisaster plots to those in ϪPisaster plots. To evaluate the relation between predation rates and key ecological processes and conditions, we quantified phytoplankton concentration and rates of mussel recruitment, mussel growth, mussel abundance, and sea star abundance.Predictions of the model are expressed as responses of predator and prey abundance, and species interaction strength (per capita and per population or total impact at the population level). As predicted by theory, per capita predation rates were independent of upwelling regime, with no variation with region. Contrary to expectation however, perpopulation predation rates were similar between intermittent-and persistent-upwelling regions but were greater under strong upwelling than under weak upwelling conditions. The greatest variation in per-population predation rates was at the level of site within region. Also contrary to theory, average abundances of prey (mussel cover) and predators (sea stars) were similar among oceanographic regions and varied mostly at the site level.As expected from theory, predation rate was high where sea star density was high, a condition that often coincided with a high food supply (phytoplankton) for filter feeders, including larvae, and high recruitment. With the exception of two sites having dense sea star populations and thus high predation, low values of either or both were associated with low predation, suggesting that the supply of prey often depended on conditions that favored subsidies of both phytoplankton and new larvae to prey populations. The occurrence of high predator density and high predation at sites of low inputs of particulate food and propagules suggests that understanding sea star life history is a key to a fuller understanding of variation in predation on a coastal scale. Evidence suggests that often sporadic recruitment of sea stars along the coast is balanced by great longevity, which tends to even out predation impact on coastal intertidal communities.

Activated defenses against herbivores and predators are defenses whereby a precursor compound is stored in an inactive or mildly active form. Upon damage to the prey, the precursor is enzymatically converted to a more potent toxin or feeding deterrent. In marine systems, activated defenses are only known to exist in a few species of tropical macroalgae. In this study, we examined an activated defense system in temperate marine macroalgae in which the osmolyte dimethylsulfoniopropionate (DMSP) is converted to acrylic acid or acrylate, depending upon the pH, and dimethyl sulfide (DMS) by the enzyme DMSP lyase upon damage to the alga. We surveyed 39 species of red, green, and brown algae from the Washington and Oregon coasts, and found high concentrations of DMSP in the chlorophytes Acrosiphonia coalita, Codium fragile, Enteromorpha intestinalis, E. linza, Ulva californica, U. fenestrata, and U. taeniata, and in the rhodophyte Polysiphonia hendryi. Concentrations of DMSP ranged from 0.04% of the alga's fresh mass (FM) to 1.8% FM. We found significant DMSP lyase activity in 1 green alga, U. fenestrata, and 1 red alga, P. hendryi, with DMSP cleavage rates approaching 300 mmol kg-1 FM min-1. Loss of DMSP and the production of DMS when the tissues of U. californica and P. hendryi were crushed suggested that physical damage results in DMSP cleavage. In laboratory feeding preference experiments, acrylic acid deterred feeding by the sea urchin Strongylocentrotus droebachiensis at concentrations of 0.1 to 2% FM and by S. purpuratus at 0.25 to 2% FM, while the precursor DMSP functioned as a feeding attractant to both sea urchins. In contrast, feeding by the isopod Idotea wosnesenskii was not deterred by acrylic acid even at concentrations as high as 8% FM. Our data suggest that DMSP may function as a precursor in an activated defense system in diverse species of temperate macroalgae and may possibly contribute to the widespread success of the Ulvophyceae. This chemical system is also found in unicellular phytoplankton, and presents an opportunity to compare and contrast the ecological role of chemical defense among micro-and macroorganisms.

The comparative‐experimental approach uses identically designed, replicated experiments at different sites along environmental gradients in order to gain insight into the changing dynamics of communities with changing environmental conditions. Such studies reveal how ecological processes vary in intensity and interact to produce community structure. Early emphases were on the community consequences of shifting top‐down impacts, competition and disturbance with environmental stress. Recent advances include the more precise quantification of gradients and thus a better understanding of species responses to the environment, and the revelation that bottom‐up forces can vary significantly on within‐region scales, with major consequences for the impact of top‐down forces and thus community dynamics. Here the use of the method to examine the role of geographic location (coastal ecosystems in different hemispheres) and oceanographic conditions (upwelling vs downwelling) on these bottom‐up/top‐down linkages is advanced. We show that a bottom‐up factor (prey recruitment) and a top‐down effect (predation rate) vary consistently with oceanographic conditions within each coastal ecosystem, and also between geographic locations (New Zealand, Oregon). In general, both recruitment and predation rates are higher in Oregon. It is postulated that these differences are common responses to oceanographic variation, and that between‐hemisphere differences result from the stronger and more persistent upwelling in the California Current ecosystem.

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