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Structural complexity mediates ecological processes such as predation, competition, and recruitment in marine systems, but relatively little is known about its effects on herbivory. In temperate seagrasses, such as eelgrass (Zostera marina), the primary herbivores are small crustacean and gastropod mesograzers that promote seagrass persistence by preferentially consuming competing epiphytic algae. We used a laboratory grazing experiment, a field colonization experiment, and stable isotope analysis to determine whether one component of eelgrass structural complexity, shoot density, dictates the strength of mesograzer top-down effects on epiphytic algae, and whether this is influenced by mesograzer community composition. Our results suggest that increasing structural complexity shifted eelgrass communities from a bottom-up to a top-down controlled system. In the lab, mesograzers reduced epiphyte standing stock only in high-shoot density experimental communities, though grazing impact varied among different combinations of dominant mesograzer taxa. In our field experiment, epiphyte biomass was inversely correlated with mesograzer density in high but not in low-shoot density eelgrass plots. High-shoot density plots contained lower epiphyte biomass despite housing lower densities of mesograzers, when compared to low-density plots, suggesting potential effects of mesograzer behavior, community composition, or self-shading on epiphyte growth. Our results suggest that structural complexity can strongly influence rates of top-down and bottom-up processes in eelgrass habitat, and should be incorporated into future experiments on the role of herbivores in seagrass ecosystems.

Predator-prey interactions are strongly influenced by habitat structure, particularly in coastal marine habitats such as seagrasses in which structural complexity (SC) may vary over small spatial scales. For seagrass mesopredators such as juvenile fishes, optimality models predict that fitness will be maximized at levels of SC that enhance foraging but minimize predation risk, both of which are functions of body size. We tested the hypothesis that in eelgrass (Zostera marina) habitat, optimal SC for juvenile giant kelpfish (Heterostichus rostratus), an abundant eelgrass mesopredator in southern California, changes through ontogeny. To do this, we quantified eelgrass SC effects on habitat associations, relative predation risk, and foraging efficiency for three size classes of juvenile giant kelpfish. We found that habitat selection differed with fish size: small fish selected dense eelgrass, whereas larger fish selected sparse eelgrass. Small kelpfish experienced the lowest relative predation risk in dense eelgrass but also had higher foraging efficiency in dense eelgrass, suggesting that dense eelgrass is selected by these fish because it minimizes risk and maximizes potential for growth. Surprisingly, larger kelpfish did not experience lower predation risk than small kelpfish. However, larger kelpfish experienced higher foraging efficiency in sparse eelgrass vs. dense eelgrass, suggesting that they select sparse eelgrass to maximize foraging efficiency. Our study highlights that trade-offs between predation risk and foraging can occur within a single habitat type, that studies should consider how habitat value changes through ontogeny, and that seagrass habitat value may be maximal when within-patch variability in SC is high.

Biotic resistance is the ability of native communities to repel the establishment of invasive species. Predation by native species may confer biotic resistance to communities, but the environmental context under which this form of biotic resistance occurs is not well understood. We evaluated several factors that influence the distribution of invasive Asian mussels (Musculista senhousia) in Mission Bay, a southern California estuary containing an extensive eelgrass (Zostera marina) habitat. Asian mussels exhibit a distinct spatial pattern of invasion, with extremely high densities towards the back of Mission Bay (up to 4,000 m(-2)) in contrast with near-complete absence at sites towards the front of the bay. We established that recruits arrived at sites where adult mussels were absent and found that dense eelgrass does not appear to preclude Asian mussel growth and survival. Mussel survival and growth were high in predator-exclusion plots throughout the bay, but mussel survival was low in the front of the bay when plots were open to predators. Additional experiments revealed that consumption by spiny lobsters (Panulirus interruptus) and a gastropod (Pteropurpura festiva) likely are the primary factors responsible for resistance to Asian mussel invasion. However, biotic resistance was dependent on location within the estuary (for both species) and also on the availability of a hard substratum (for P. festiva). Our findings indicate that biotic resistance in the form of predation may be conferred by higher order predators, but that the strength of resistance may strongly vary across estuarine gradients and depend on the nature of the locally available habitat.