Local-scale nutrient regeneration facilitates seaweed growth on wave-exposed rocky shores in an upwelling system

Abstract This study shows that, even on exposed, wave-swept, rocky shores in a nutrient-replete upwelling ecosystem, mussels (Mytilus californianus) facilitate the growth of the seaweed Porphyra perforata by enhancing nutrient concentrations in the nearby water column. In field surveys on emergent substrate in the mid-intertidal zone, we found ten times greater abundance of P. perforata on mussels than on adjacent rock. In field experiments, P. perforata accumulated and grew more quickly on mussels than on bare rock or on mussel mimics, suggesting that nutrients excreted by mussels might be responsible for greater P. perforata cover. At high tide, water column ammonium concentrations over mussel beds were nearly double those found over bare rock. Correspondingly, tissue nitrogen concentrations were higher, and carbon-to-nitrogen ratios were lower in P. perforata growing on mussels compared to bare rock. Given the dominance of mussels in mid-intertidal regions of temperate coasts worldwide, ammonium regeneration could be a general contributor to local-scale nutrient availability, even in high-flow systems characterized by high nutrient concentrations

Seaweed richness and herbivory increase rate of community recovery from disturbance.

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities

Grazer diversity interacts with biogenic habitat heterogeneity to accelerate intertidal algal succession.

Environmental heterogeneity contributes to coexistence by allowing species with different traits to persist when different species perform best at different times or places. This interaction between niche differences and environmental variability may also help explain relationships between biodiversity and ecosystem functioning, but few data are available to rigorously evaluate this hypothesis. We assessed how a biologically relevant aspect of environmental heterogeneity interacts with species diversity to determine ecosystem processes in a natural rocky intertidal community. We used field removals to factorially manipulate biogenic habitat heterogeneity (barnacles, bare rock, and plots that were 50/50 mixes of the two habitat types) and gastropod grazer species richness and then tracked algal community succession and recovery over the course of 1 yr. We found that herbivore diversity, substrate heterogeneity, and their interaction played unique roles in the peak abundance and timing of occurrence of different algal functional groups. Early successional microalgae were most heavily grazed in diverse herbivore assemblages and those with barnacles present, which was likely due to complementary feeding strategies among all three grazers. In contrast, late successional macroalgae were strongly influenced by the presence of a habitat generalist limpet. In this herbivore's absence, heterogeneous habitats (i.e., mixtures of bare rock and barnacles) experienced the greatest algal accumulation, which was partly a result of complementary habitat use by the remaining herbivores. The complex way habitat identity and heterogeneity altered grazer-algal interactions in our study suggests species' differences and environmental heterogeneity both separately and interactively contribute to the relationship between biodiversity and ecosystem functions

Appendix C. Tables showing the occurrence and abundance of algal and mobile invertebrate species across treatments.

Tables showing the occurrence and abundance of algal and mobile invertebrate species across treatments

Appendix B. A figure showing percent cover of perennial algae, Shannon diversity, and density of perennial algae by algal species treatment, and a table of the estimated slopes and standard errors of percent cover of perennial algal species during the recovery period in the presence and absence of herbivores.

A figure showing percent cover of perennial algae, Shannon diversity, and density of perennial algae by algal species treatment, and a table of the estimated slopes and standard errors of percent cover of perennial algal species during the recovery period in the presence and absence of herbivores

Local-scale nutrient regeneration facilitates seaweed growth on wave-exposed rocky shores in an upwelling system

This study shows that, even on exposed, wave-swept, rocky shores in a nutrient-replete upwelling ecosystem, mussels (Mytilus californianus) facilitate the growth of the seaweed Porphyra perforata by enhancing nutrient concentrations in the nearby water column. In field surveys on emergent substrate in the mid-intertidal zone, we found ten times greater abundance of P. perforata on mussels than on adjacent rock. In field experiments, P. perforata accumulated and grew more quickly on mussels than on bare rock or on mussel mimics, suggesting that nutrients excreted by mussels might be responsible for greater P. perforata cover. At high tide, water column ammonium concentrations over mussel beds were nearly double those found over bare rock. Correspondingly, tissue nitrogen concentrations were higher, and carbon-to-nitrogen ratios were lower in P. perforata growing on mussels compared to bare rock. Given the dominance of mussels in mid-intertidal regions of temperate coasts worldwide, ammonium regeneration could be a general contributor to local-scale nutrient availability, even in high-flow systems characterized by high nutrient concentrations. © 2009, by the American Society of Limnology and Oceanography, Inc

Prey diversity is associated with weaker consumer effects in a meta-analysis of benthic marine experiments.

A rapidly accumulating body of research has shown that species diversity consistently affects the functioning of ecosystems. The incorporation of trophic complexity and the extension of this research to larger scales and natural ranges in species diversity remain as important challenges for understanding the true magnitude of these effects in natural systems. Here, we test whether the diversity of prey communities affects the magnitude of aggregate consumer effects. We conducted a meta-analysis of 57 consumer removal field experiments from a range of intertidal and subtidal hard substrate marine communities. We found that the richness of the prey community was the strongest predictor of the magnitude of consumer effects while controlling for habitat type, taxonomic composition, and other variables. Consumer removal increased aggregate prey abundance on average by 1200% at the lower limit of prey diversity (two species), but only 200% at the upper limit of 37 species. Importantly, compositional change was substantial at both high and low prey diversity, suggesting predation intensity did not vary with prey richness. Rather diverse prey communities appear to be more capable of maintaining abundance via compensatory responses, by containing prey species that are resistant to (or tolerant of) predators. These results suggest that the effects of species diversity on trophic interactions may scale consistently from small-scale manipulations to cross-community comparisons

Evolved differences in energy metabolism and growth dictate the impacts of ocean acidification on abalone aquaculture.

Ocean acidification (OA) poses a major threat to marine ecosystems and shellfish aquaculture. A promising mitigation strategy is the identification and breeding of shellfish varieties exhibiting resilience to acidification stress. We experimentally compared the effects of OA on two populations of red abalone (Haliotis rufescens), a marine mollusc important to fisheries and global aquaculture. Results from our experiments simulating captive aquaculture conditions demonstrated that abalone sourced from a strong upwelling region were tolerant of ongoing OA, whereas a captive-raised population sourced from a region of weaker upwelling exhibited significant mortality and vulnerability to OA. This difference was linked to population-specific variation in the maternal provisioning of lipids to offspring, with a positive correlation between lipid concentrations and survival under OA. This relationship also persisted in experiments on second-generation animals, and larval lipid consumption rates varied among paternal crosses, which is consistent with the presence of genetic variation for physiological traits relevant for OA survival. Across experimental trials, growth rates differed among family lineages, and the highest mortality under OA occurred in the fastest growing crosses. Identifying traits that convey resilience to OA is critical to the continued success of abalone and other shellfish production, and these mitigation efforts should be incorporated into breeding programs for commercial and restoration aquaculture

Seawater carbonate chemistry and survival and growth of two populations of red abalone (Haliotis rufescens)

Ocean acidification (OA) poses a major threat to marine ecosystems and shellfish aquaculture. A promising mitigation strategy is the identification and breeding of shellfish varieties exhibiting resilience to acidification stress. We experimentally compared the effects of OA on two populations of red abalone (Haliotis rufescens), a marine mollusc important to fisheries and global aquaculture. Results from our experiments simulating captive aquaculture conditions demonstrated that abalone sourced from a strong upwelling region were tolerant of ongoing OA, whereas a captive-raised population sourced from a region of weaker upwelling exhibited significant mortality and vulnerability to OA. This difference was linked to population-specific variation in the maternal provisioning of lipids to offspring, with a positive correlation between lipid concentrations and survival under OA. This relationship also persisted in experiments on second-generation animals, and larval lipid consumption rates varied among paternal crosses, which is consistent with the presence of genetic variation for physiological traits relevant for OA survival. Across experimental trials, growth rates differed among family lineages, and the highest mortality under OA occurred in the fastest growing crosses. Identifying traits that convey resilience to OA is critical to the continued success of abalone and other shellfish production, and these mitigation efforts should be incorporated into breeding programs for commercial and restoration aquaculture