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Foundation species are vital to the maintenance of biodiversity and ecosystem functioning in many systems. On rocky shores, rockweeds (large brown algae in the Order Fucales) have the potential to provide habitat and ameliorate stress for mobile invertebrates. To determine the relative role of 2 rockweeds (Silvetia compressaandPelvetiopsisspp.) as foundation species at sites along a latitudinal gradient, we conducted observational surveys and then initiated a 12 mo removal experiment. We found that richness and abundance of mobile invertebrates declined over time when rockweeds were removed, but only at the southernmost site. In contrast, at our other sites, there was no change in the richness and abundance of mobile invertebrates following rockweed removal. At the southern site, rockweeds played an important role in maintaining mobile invertebrate diversity. At our central and northern sites, rockweeds were less important in maintaining the diversity of mobile invertebrates. At these sites, alternative species, including bladed and branching taxa in the generaMastocarpus,Mazzaella,Corallina, andEndocladia, co-occur with rockweeds and can buffer the system against their loss. However, these alternative foundation species are rare to absent at the southern site, potentially due to greater physical stress. The loss of rockweed foundation species, which are declining at our southern site, can have cascading effects by causing local co-extinctions of associated species. This study highlights the importance of foundation species, especially in areas where their functional redundancy is low, and how the loss of foundation species can alter diversity, leading to potential changes in ecosystem functioning.

Foundation species provide physical structure that enhances the diversity and abundance of associated organisms. Canopy-forming seaweeds are known to act as foundation species on rocky shores by lowering temperature and desiccation stress. Direct solar radiation, including ultraviolet (UV) light, can also reduce photosynthetic rates in algae, cause oxidative stress and DNA damage. The reduction in UV exposure provided by an algal canopy could therefore be important for understory organisms, including the red alga Chondrus crispus on New England's (USA) rocky shores, and this relationship may be more important at higher tidal elevations with increased low-tide exposure time. In field experiments, we investigated the relationship between tidal elevation and an index of C. crispus UV exposure, the concentration of UV-absorbing pigments. Low on the shore, C. crispus grew without a canopy. Higher on the shore, in the mid-intertidal zone, C. crispus was found under the canopy-forming rockweed, Fucus distichus subsp. evanescens. At this elevation, C. crispus was shaded (>50%; >1 m above MLLW). We performed a canopy removal experiment that spanned the mid-zone where C. crispus and F. distichus subsp. evanescens co-occur and the low-zone (no canopy). Following canopy removal in the mid-zone, UV pigment concentrations increased with tidal elevation. After accounting for the effect of elevation, removal of the algal canopy resulted in UV-protective pigment concentrations 2-fold higher than in un-manipulated control plots. These results suggest that amelioration of solar UV exposure might be another mechanism by which canopy seaweeds, acting as foundation species, facilitate understory seaweeds on rocky shores.

The co-occurrence of several species, all of which share similar resource requirements, remains a paradox in ecology. Here, I evaluated resource use along multiple environmental axes to understand the potential for niche partitioning and complementarity in a guild of suspension-feeding rocky shore invertebrates. I focused on the mussels Perna canaliculus, Mytilus galloprovincialis, Aulacomya maoriana, and Xenostrobus pulex, all of which coexist along the coastline of the South Island of New Zealand. I quantified the mussels’ distributions, both vertically on the shore and within the three-dimensional mussel bed matrix, and used carbon (δ13C) and nitrogen (δ15N) stable isotope ratios to compare species’ diets. Mussels exhibited niche partitioning along all resource axes. Given the mussels’ use of different spatial and food resources, I evaluated the potential for complementarity with respect to mussels’ roles as mediators of an important spatial subsidy, carbon inputs from the nearshore ocean into rocky-shore ecosystems. In these systems, mussels are basal species, capturing and consuming particulate organic matter in the ocean and making it available for local consumption within the benthic community. I found that mussel diversity matters; even the most productive species—Perna canaliculus—only contributed about half of the mussel-mediated carbon that accumulated over a year. Multiple co-occurring species are, therefore, likely to provide more carbon at the base of the intertidal food web than any one species living independently, and complementarity in resource use along multiple environmental axes is likely an important mechanism underlying this relationship.

A recommendation of: Thibault Androuin, Stanislas F. Dubois, Cedric Hubas, Gwendoline Lefebvre, Fabienne Le Grand, Gauthier Schaal, Antoine Carlier Trophic niche of the invasive gregarious species Crepidula fornicata, in relation to ontogenic changes 10.1101/2020.07.30.229021

Susan Lynn Williams (1951–2018) was an exceptional marine ecologist whose research focused broadly on the ecology of benthic nearshore environments dominated by seagrasses, seaweeds, and coral reefs. She took an empirical approach founded in techniques of physiological ecology. Susan was committed to applying her research results to ocean management through outreach to decision-makers and resource managers. Susan’s career included research throughout the USA in tropical, temperate, and polar regions, but she specialized in tropical marine ecology. Susan’s scholarship, leadership, and friendship touched many people, leading to this multi-authored paper. Susan’s scholarship was multi-faceted, and she excelled in scientific discovery, integration of scientific results, application of science for conservation, and teaching, especially as a mentor to undergraduate and graduate students and postdoctoral scholars. Susan served in a variety of leadership positions throughout her career. She embodied all facets of leadership; leading by example, listening to others, committing to the “long haul,” maintaining trust, and creating a platform for all to shine. Susan was an important role model for women in science. Susan was also a loyal friend, maintaining friendships for many decades. Susan loved cooking and entertaining with friends. This paper provides an overview of the accomplishments of Susan in the broad categories of scholarship, leadership, and friendship.

Some sedentary marine invertebrates have the potential to modify the environments they experience by moving, even as adults. Of particular interest are sea anemones, which, despite appearing immobile, can move throughout their lives. Individual locomotion may mitigate changes in environment conditions and, therefore, play an important role in the natural history of sea anemones, especially in naturally variable and/or stochastic environments. Sea anemones that associate with algal endosymbionts may respond to changes in nutrition, both autotrophic (from algae) and heterotrophic (from prey). Here, we describe the adult movement behaviors and asexual reproduction of the sea anemone Exaiptasia diaphana in response to changes in food availability and photosymbiont density. Anemones were collected from mangrove roots in the Florida Keys USA (24° 49′ 21.91″ N, 80° 48′ 37.95″ W) during January 2016 and exposed to a factorial experiment in which food availability and exposure to temperature shock were manipulated. Sea anemones exhibited a variety of responses, including (1) increased crawling along the substrate in response to starvation, (2) increased detachment from the substrate and reattachment in a new location in response to starvation, and (3) increased production of motile asexual clones in response to both starvation and temperature-induced changes in symbiont density. These responses are shaped not only by the direct consequences to the sea anemone, but also by the effects on the symbiotic algae, which exchange sugars, lipids, and oxygen for nutrients within the host. Observed patterns of movement and reproduction are likely advantageous for life in the dynamic mangrove root fouling communities where this anemone species occurs. The ability to disperse as an adult may give this otherwise sedentary invertebrate an advantage in naturally stochastic conditions or in rapidly changing environments.

Intraspecific differentiation across a steep environmental gradient depends on the relative influences of evolutionary, organismal, and environmental processes. But steep environmental gradients may be nested within larger-scale, regional conditions that could influence these processes at the local scale. Therefore, we hypothesized that phenotypic differentiation along a steep environmental gradient would vary among regions. To test this hypothesis, we conducted a reciprocal transplant experiment on rocky intertidal shores, a habitat characterized by gradients in abiotic and biotic stress, in three regions of the Gulf of Maine. We used the ubiquitous and ecologically important rockweed species Fucus vesiculosus to quantify differentiation in growth, tissue nitrogen, and nitrogen productivity between upper and lower intertidal individuals. We found that phenotypic differentiation between tide heights varied among traits and regions. Although tissue nitrogen did not vary among any treatment combinations, growth and nitrogen productivity response were region specific. A strong effect of transplant height was found in all regions; however, an effect of home (source) height was only detectable in the central Gulf of Maine. Our study reveals that intraspecific responses to steep environmental gradients vary among populations, but the mechanisms underlying these patterns remain unknown. Given the roles that rockweeds play as food and habitat, these in situ patterns of growth and nitrogen productivity could have important community- and ecosystem-level consequences.

Climatic warming is a primary driver of change in ecosystems worldwide. Here, we synthesize responses of species richness and evenness from 187 experimental warming studies in a quantitative meta-analysis. We asked 1) whether effects of warming on diversity were detectable and consistent across terrestrial, freshwater and marine ecosystems, 2) if effects on diversity correlated with intensity, duration, and experimental unit size of temperature change manipulations, and 3) whether these experimental effects on diversity interacted with ecosystem types. Using multilevel mixed linear models and model averaging, we also tested the relative importance of variables that described uncontrolled environmental variation and attributes of experimental units. Overall, experimental warming reduced richness across ecosystems (mean log-response ratio = -0.091, 95% bootstrapped CI: -0.13, -0.05) representing an 8.9% decline relative to ambient temperature treatments. Richness did not change in response to warming in freshwater systems, but was more strongly negative in terrestrial (-11.8%) and marine (-10.5%) experiments. In contrast, warming impacts on evenness were neutral overall and in aquatic systems, but weakly negative on land (7.6%). Intensity and duration of experimental warming did not explain variation in diversity responses, but negative effects on richness were stronger in smaller experimental units, particularly in marine systems. Model-averaged parameter estimation confirmed these main effects while accounting for variation in latitude, ambient temperature at the sites of manipulations, venue (field versus lab), community trophic type, and whether experiments were open or closed to colonization. These analyses synthesize extensive experimental evidence showing declines in local richness with increased temperature, particularly in terrestrial and marine communities. However, the more variable effects of warming on evenness were better explained by the random effect of site identity, suggesting that effects on species' relative abundances were contingent on local species composition.

Biodiversity is changing on both global and local scales, motivating research to understand the consequences of these changes for how communities and ecosystems function. Here, we explore the role of life history strategies in mediating biodiversity and ecosystem functioning. In particular, we evaluate how the composition, biomass (% cover), and richness of perennial (persistence ≥ 1 year) and ephemeral (persistence < 1 year) species change along a gradient of increasing seaweed species richness on a rocky shoreline. We show that the majority of biomass is comprised of perennial species, especially where overall richness is low, whereas the majority of species are ephemeral, especially where overall richness is high. We then present and discuss the results of an 18-month field manipulation quantifying the factorial effects of tidal elevation, wave exposure, herbivore removals, thermal and desiccation stress amelioration, and nutrient additions on perennial versus ephemeral species. In particular, the diversity of ephemeral species was strongly affected, relative to perennial species, by tidal elevation, wave exposure, and herbivory; herbivores reduced diversity of ephemeral species relative to perennials. Relative to perennial cover, ephemeral cover was greater higher on the shore, in more wave-exposed habitats, and where herbivores were removed, plots were unscreened, and/or nutrients were added. Thus, perennials and ephemerals responded differently to environmental context and experimental manipulation. We compared nitrate uptake and photosynthesis rates of ephemeral and perennial species and found that maximum nitrate uptake and photosynthesis rates of ephemerals were twice as high as those of perennials. These results highlight the disproportionate roles that ephemeral species play in mediating ecosystem-level processes. In combination with our comparisons of the diversity and cover of perennial and ephemeral species along a biodiversity gradient, these results demonstrate the utility of incorporating life history traits into our efforts to understand the functional consequences of biodiversity change.

Species invasions have been increasing in frequency worldwide, yet critical gaps remain in our understanding of how invaders affect community structure and ecosystem functioning, particularly during the initial stages of invasion. Even less is known about changes in the invader that may take place immediately following an invasion. This study examined the recent invasion of the red macroalga Dasysiphonia (formerly, Heterosiphonia) japonica to the western North Atlantic Ocean with the aim of filling in gaps in our understanding of the impacts that invasive seaweeds have at the species, community and ecosystem levels immediately following their establishment. Within 5 years of invasion, community composition had changed and biodiversity had decreased to nearly half of pre-invasion levels. In addition, the relative proportion of Dasysiphonia decreased by 35% over our four-year study from initially high levels shortly after establishment. We found evidence that functional traits of this initially aggressive invader changed over time, as it ultimately became a less aggressive, co-inhabiting member of the local algal community, particularly with respect to nutrient uptake and relative abundances, although native diversity remained low relative to pre-invasion levels. Using these realistic changes in community structure, including decreases in biodiversity, we also showed that nutrient uptake of algal assemblages changed over time, suggesting changes in the functional characteristics of invaded communities, with implications for ecosystem-level processes such as nutrient fluxes. This study provides rare empirical evidence about the successional stages occurring at the individual, community, and ecosystem levels during the first 5 years of an invasion.

Intertidal seaweeds must cope with a suite of stressors imposed by aerial exposure at low tide, including nutrient limitation due to emersion. Seaweeds can access nutrients only when submerged, so individuals living higher compared to lower on the shore may have adaptations allowing them to acquire sufficient amounts of nutrients to survive and maintain growth. Using a combination of observations and experiments, we aimed to identify intraspecific variation in nitrate uptake rates across the intertidal distribution of F. vesiculosus, as well as test for acclimation in response to a change in tide height. We replicated our study at sites spanning nearly the entire Gulf of Maine coastline, to examine how local environmental variability may alter intraspecific variation in nitrate uptake. We found that average nitrate uptake rates were ~18% higher in upper compared to lower intertidal Fucus vesiculosus. Furthermore, we found evidence for both acclimation and adaptation to tide height during a transplant experiment. F. vesiculosus transplanted from the lower to the upper intertidal zone was characterized by increased nitrate uptake, but individuals transplanted from the upper to the lower intertidal zone retained high uptake rates. Our observations differed among Gulf of Maine regions and among time points of our study. Importantly, these differences may reflect associations between nitrate uptake rates and abiotic environmental conditions and seaweed nutrient status. Our study highlights the importance of long-term variation in ambient nutrient supply in driving intraspecific variation of seaweeds across the intertidal gradient and local and seasonal variation in ambient nutrient levels in mediating intraspecific differences.

Mutualistic symbioses are common, especially in nutrient-poor environments where an association between hosts and symbionts can allow the symbiotic partners to persist and collectively out-compete non-symbiotic species. Usually these mutualisms are built on an intimate transfer of energy and nutrients (e.g. carbon and nitrogen) between host and symbiont. However, resource availability is not consistent, and the benefit of the symbiotic association can depend on the availability of resources to mutualists. We manipulated the diets of two temperate sea anemone species in the genusAnthopleurain the field and recorded the responses of sea anemones and algal symbionts in the family Symbiodiniaceae to our treatments. Algal symbiont density, symbiont volume and photosynthetic efficiency of symbionts responded to changes in sea anemone diet, but the responses depended on the species of sea anemone. We suggest that temperate sea anemones and their symbionts can respond to changes in anemone diet, modifying the balance between heterotrophy and autotrophy in the symbiosis. Our data support the hypothesis that symbionts are upregulated or downregulated based on food availability, allowing for a flexible nutritional strategy based on external resources.

Author(s): Wallingford, Piper D.; Pandori, Lauren L.M.; Bedgood, Samuel A.; Bracken, Matthew E.S.; Elsberry, Laura A.; Henry, Amy K.; Mahanes, Samuel A.; Sorte, Cascade J.B. | Abstract

Herbivores mediate the abundances of primary producers both from the top-down, by consuming them, and from the bottom-up, by recycling nutrients. Whereas the top-down effects of herbivores on algae in marine ecosystems are well-documented, less is known about their roles as mediators of local-scale nutrient availability. We conducted a series of surveys and measurements of tide pools and the grazers in those pools between October of 2016 and June of 2017 at an intertidal site on the coast of Southern California, USA (33°35′16.3″N, 117°52′1.5″W). We surveyed grazer abundances in the field, measured biomass of representatives from four different grazer groups (littorine snails, limpets, chitons, and turban snails), measured ammonium excretion rates, and quantified ammonium accumulation rates in tide pools at our study site. We found that different grazer groups were characterized by different per-biomass ammonium excretion rates. Some grazer groups—turban snails and chitons—contributed more ammonium than predicted by their biomass, whereas other grazer groups—littorine snails and limpets—contributed less ammonium than predicted by biomass. Because of these differences between grazer groups, ammonium accumulation rates in tide pools at our study site were effectively predicted based on the ammonium excretion rates of the different grazer groups. However, ammonium accumulation rates were not related to total herbivore biomass. Our results highlight the importance of grazer identity—and particularly the role of species such as turban snails that contribute disproportionately to nutrient recycling—in understanding the contributions of grazers as mediators of bottom-up processes in marine systems.

Predicting the impacts of ocean acidification in coastal habitats is complicated by bio-physical feedbacks between organisms and carbonate chemistry. Daily changes in pH and other carbonate parameters in coastal ecosystems, associated with processes such as photosynthesis and respiration, often greatly exceed global mean predicted changes over the next century. We assessed the strength of these feedbacks under projected elevated CO2 levels by conducting a field experiment in 10 macrophyte-dominated tide pools on the coast of California, USA. We evaluated changes in carbonate parameters over time and found that under ambient conditions, daytime changes in pH, pCO2, net ecosystem calcification (NEC), and O2 concentrations were strongly related to rates of net community production (NCP). CO2 was added to pools during daytime low tides, which should have reduced pH and enhanced pCO2. However, photosynthesis rapidly reduced pCO2 and increased pH, so effects of CO2 addition were not apparent unless we accounted for seaweed and surfgrass abundances. In the absence of macrophytes, CO2 addition caused pH to decline by ∼0.6 units and pCO2 to increase by ∼487 µatm over 6 hr during the daytime low tide. As macrophyte abundances increased, the impacts of CO2 addition declined because more CO2 was absorbed due to photosynthesis. Effects of CO2addition were, therefore, modified by feedbacks between NCP, pH, pCO2, and NEC. Our results underscore the potential importance of coastal macrophytes in ameliorating impacts of ocean acidification.

Predicting the impacts of ocean acidification in coastal habitats is complicated by bio-physical feedbacks between organisms and carbonate chemistry. Daily changes in pH and other carbonate parameters in coastal ecosystems, associated with processes such as photosynthesis and respiration, often greatly exceed global mean predicted changes over the next century. We assessed the strength of these feedbacks under projected elevated CO2 levels by conducting a field experiment in 10 macrophyte-dominated tide pools on the coast of California, USA. We evaluated changes in carbonate parameters over time and found that under ambient conditions, daytime changes in pH, pCO2, net ecosystem calcification (NEC), and O2 concentrations were strongly related to rates of net community production (NCP). CO2 was added to pools during daytime low tides, which should have reduced pH and enhanced pCO2. However, photosynthesis rapidly reduced pCO2 and increased pH, so effects of CO2 addition were not apparent unless we accounted for seaweed and surfgrass abundances. In the absence of macrophytes, CO2 addition caused pH to decline by ∼0.6 units and pCO2 to increase by ∼487 µatm over 6 hr during the daytime low tide. As macrophyte abundances increased, the impacts of CO2 addition declined because more CO2 was absorbed due to photosynthesis. Effects of CO2addition were, therefore, modified by feedbacks between NCP, pH, pCO2, and NEC. Our results underscore the potential importance of coastal macrophytes in ameliorating impacts of ocean acidification.

Author(s): Elsberry, Laura A.; Fales, Robin J.; Bracken, Matthew E.S. | Abstract: Biodiversity is currently threatened at local, regional, and global scales, and identifying the species that are vulnerable to these changes is essential for conservation efforts. For example, the breadth of species’ ranges may offer important clues to their susceptibility to loss, as widespread species may be more resistant to loss than species with a narrower range breadth. To determine the potential for shifts in community interactions along the latitudinal geographic ranges of species, we examined pair-wise associations between narrow-range endemic and widespread rocky intertidal species. We surveyed rocky intertidal species composition at eight sites along the California coast from San Diego to Cape Mendocino. Four sites were south of Point Conception, and four sites were north of Point Conception. Point Conception is a major biogeographic feature for coastal marine species, where sea surface temperatures transition from cool temperate waters in the north to warm temperate waters in the south. To determine whether pair-wise species associations were significant, we compared the observed communities’ standardized effect size to a null model to determine which species occurred together more or less often than by chance. Across all sites, widespread species were considerably more abundant than narrow-range endemic species, and the majority of species were widespread. However, total species richness was unrelated to the number of widespread species, and was, instead, determined by the number of narrow-range endemic species present at a site. Our analyses suggest that species are more aggregated than segregated south of Point Conception, but the opposite is true north of Point Conception. Additionally, we found that species associations between narrow-range endemics drove the overall patterns in species associations. One possible explanation for these patterns is that positive interactions, especially those involving narrow-range endemic species, are more important in southern California’s more thermally stressful intertidal habitats.

Most studies evaluating the effects of biodiversity on ecosystem functioning are conducted at a single location, limiting our understanding of how diversity-function relationships may change when measured across different spatial scales. We conducted a species-removal experiment at three sites nested in each of three regions along the rocky intertidal coastline of the Gulf of Maine, USA, to evaluate the potential for scale-dependent effects of species loss on the biomass of intertidal seaweed assemblages. We randomly assigned 50 plots in the mid-intertidal zone at each site to one of five treatments (n = 10 plots each): an unmanipulated control, a polyculture plot that contained our three target seaweed species, and three monoculture plots. We manipulated diversity by removing all non-target species from monoculture and polyculture plots, then removing additional biomass from polyculture plots, proportionate to species' relative abundances, so that the average amount removed from monoculture and polyculture plots was equivalent at each site. At the largest spatial scale, all sites considered together, after accounting for region and site nested within region seaweed diversity had consistent, positive effects on seaweed cover. Diverse polyculture plots always had higher cover than was predicted by the average performance of the component seaweed species and usually had higher cover than even the best-performing component species. Diversity effects weakened and became less consistent at smaller spatial scales, so that at the scale of individual sites, diverse polycultures only performed better than the average of monocultures ~40% of the time. Hence, our results indicate that weak and/or inconsistent biodiversity effects at the level of individual sites may scale up to stronger, more consistent effects at larger spatial scales. Quantitative summaries of biodiversity experiments conducted at the scale of individual sites do not capture this spatial aspect of biodiversity effects and may therefore underestimate the functional consequences of biodiversity loss.

Humans are modifying the availability of nutrients such as nitrogen (N) and phosphorus (P), and it is therefore important to understand how these nutrients, independently or in combination, influence the growth and nutrient content of primary producers. Using meta-analysis of 118 field and laboratory experiments in freshwater, marine and terrestrial ecosystems, we tested hypotheses about co-limitation of N and P by comparing the effects of adding N alone, P alone, and both N and P together on internal N (e.g. %N, C:N) and P (e.g. %P, C:P) concentrations in autotroph communities. In particular, we tested the following predictions. First, if only one nutrient was limiting, addition of that nutrient should decrease the concentration of the other nutrient, but addition of the non-limiting nutrient would have no effect on the internal concentration of the limiting nutrient. If community co-limitation was occurring then addition of either nutrient should result in a decrease in the internal concentration of the other nutrient. Community co-limitation could also result in no change – or even an increase – in N concentrations in response to P addition if P stimulated growth of N fixers. Finally, if biochemically dependent co-limitation was occurring, addition of a limiting nutrient would not decrease, and could even increase, the concentration of the other, co-limited nutrient. We found no general evidence for the decrease in the internal concentration of one nutrient due to addition of another nutrient. The one exception to this overall pattern was marine systems, where N addition decreased internal P concentrations. In contrast, P addition increased internal N concentrations across all experiments, consistent with co-limitation. These results have important implications for understanding the roles that N and P play in controlling producer growth and internal nutrient accumulation as well as for managing the effects of nutrient enrichment in ecosystems. Synthesis On a global scale, humans have doubled nitrogen (N) inputs and quadrupled phosphorus (P) inputs relative to pre-industrial levels. N and P fertilization influences autotroph internal nutrient concentrations and ratios and thereby affects a variety of community and ecosystem processes, including decomposition and consumer population dynamics. It is therefore critical to understand the effects of nutrient additions on the growth and nutrient concentrations of primary producers. We used meta-analysis to evaluate the responses of autotroph internal N and P concentrations to additions of N, P, and N+P and make inferences about limitation and co-limitation of N and P across marine, terrestrial, and freshwater ecosystems. We found little evidence for single-nutrient limitation, highlighting the fact that multiple nutrients generally limit primary production.

Interactions between grazers and autotrophs are complex, including both top-down consumptive and bottom-up facilitative effects of grazers. Thus, in addition to consuming autotrophs, herbivores can also enhance autotroph biomass by recycling limiting nutrients, thereby increasing nutrient availability. Here, we evaluated these consumptive and facilitative interactions between snails (Littorina littorea) and seaweeds (Fucus vesiculosus and Ulva lactuca) on a rocky shore. We partitioned herbivores' total effects on seaweeds into their consumptive and facilitative effects and evaluated how community context (the presence of another seaweed species) modified the effects of Littorina on a focal seaweed species. Ulva, the more palatable species, enhanced the facilitative effects of Littorina on Fucus. Ulva did not modify the consumptive effect of Littorina on Fucus. Taken together, the consumptive and facilitative effects of snails on Fucus in the presence of Ulva balanced each other, resulting in no net effect of Littorina on Fucus. In contrast, the only effect of Fucus on Ulva was to enhance consumptive effects of Littorina on Ulva. Our results highlight the necessity of considering both consumptive and facilitative effects of herbivores on multiple autotroph species in order to gain a mechanistic understanding of grazers' top-down and bottom-up roles in structuring communities.

The successful establishment and spread of invasive species may be determined by intrinsic traits of the invader, characteristics of the invaded community, and interactions between the two. The red filamentous macroalga Heterosiphonia japonica has recently invaded and expanded its range in the northwestern Atlantic. We surveyed and compared macroalgal communities at sites with different levels of invasion in Nahant, Massachusetts, and examined potential factors contributing to Heterosiphonia’s successful invasion by assessing its growth rates and rates of herbivory by native grazers in the laboratory. We evaluated growth and herbivory for isolated individuals and for individuals within a macroalgal assemblage comprised of the most abundant species in the native community. We also measured macroalgal nitrate uptake rates to examine a potential mechanism underlying observed differences in growth rate. High abundances of Heterosiphonia were associated with lower native macroalgal species richness, evenness, and diversity and with differences in species composition in the field. Experiments showed that rates of growth and grazing were context dependent. In isolation, Heterosiphonia grew comparably and was grazed similarly to the two native filamentous macroalgae. Within the context of the native macroalgal assemblage, it grew much faster than all common native species, experienced reduced herbivory from one native grazer, and increased herbivory from another. Consistent with its faster growth, Heterosiphonia was the most efficient at nitrate uptake. Our results suggest that multiple interactions between intrinsic traits (e.g., competitiveness for nutrients) and extrinsic factors (e.g., presence of native macroalgal species, identity and relative abundances of native grazers) contribute to Heterosiphonia’s invasion success.

Nutrients such as nitrogen (N) and phosphorus (P) limit primary productivity, and recent anthropogenic activities are changing the availability of these nutrients, leading to alterations in the type and magnitude of nutrient limitation. Recent work highlights the potential for N and P to interact to limit primary production in terrestrial and freshwater systems. However, mechanisms underlying co-limitation are not well described. Documentation of ambient nutrient levels and tissue nutrients of the intertidal macroalga Fucus vesiculosus for 2 years in the southern Gulf of Maine, USA, indicates that N availability may be impacting the ability of F. vesiculosus to access P, despite relatively high ambient P concentrations. To experimentally validate these observations, F. vesiculosus individuals were enriched with N or P for 6 weeks, followed by an uptake experiment to examine how the interactions between these nutrients affected macroalgal N and P uptake efficiency. Results illustrate that exposure of seaweed to different nutrient regimes influenced nutrient uptake efficiency. Notably, seaweeds enriched with N displayed the highest P uptake efficiency at low, biologically relevant, P concentrations. Our results confirm that N availability may be mediating the ability of primary producers to access P. These interactions between limiting nutrients have implications for not only the growth and functioning of primary producers who rely directly on these nutrients but also the entire communities that they support.

Synthesis The tissue chemistry of plants can influence ecosystem processes including growth, herbivory, and decomposition. Our comparison of nitrogen and phosphorus in over 1700 autotroph taxa demonstrates that latitudinal trends in tissue chemistry are consistent across non-vascular and vascular species in freshwater, terrestrial, and marine ecosystems. Tissue chemistry varies most within species and taxonomic lineages, yet the nitrogen to phosphorus ratio within individuals is strikingly similar among species in different ecosystems. These results shed new light on existing hypotheses, suggesting that light (e.g. photon flux) and growing season duration are primary drivers of latitudinal gradients in tissue chemistry, but providing little support for temperature, nutrient supply, or soil substrate age. Photoautotroph nitrogen (N) and phosphorus (P) tissue concentrations can influence ecosystem function via processes including growth, decomposition, and consumption, and may reflect traits maintaining coexistence. Studies in terrestrial systems have led to hypotheses that latitudinal trends in the N and P content of leaves may be driven by soil substrate age, environmental temperature, or season length; however, terrestrial patterns alone cannot differentiate these mechanisms. Here, we demonstrate that broad geographical patterns of N and P in freshwater and marine multicellular photoautotrophs are concordant with those in terrestrial ecosystems. Our > 6800 record database reveals that mean tissue N and P increase with latitude in all ecosystems, but P increases more rapidly, causing N:P to decline; mean N:P scaling within individuals also is identical among systems, despite very different evolutionary environments. A partitioning of the variance in these data suggests that species composition and local environmental context likely lead to the variation observed within a latitudinal band. However, the consistency of trends in photosynthetic tissue chemistry across Earth’s ecosystems suggests that biogeographical gradients in insolation and growing season length may constrain tissue N and P, whereas global trends in temperature, nutrient supply, and soil substrate age are unlikely to generate the consistent latitudinal trends among ecosystems. Thus, this cross-ecosystem comparison suggests a new hypothesis, global patterns of insolation, while also providing a new perspective on other mechanisms that have been hypothesized to underlie latitudinal trends in photosynthetic tissue chemistry.

Patterns in rocky intertidal seaweed biodiversity influence the resilience and functioning of these important primary producer communities. In turn, seaweed biodiversity patterns are the result of many ecological factors. We determined the influences of thermal and desiccation stress, herbivory, and nutrients on seaweed biodiversity on a northern California rocky shoreline. In a fully crossed design at two tidal heights at wave-protected and exposed sites, we deployed screens to reduce stress, removed herbivores, and added nutrients for 18 months. The treatments reduced temperature, increased relative humidity, decreased herbivore abundances, and increased nitrogen in both seawater and seaweeds. Seaweed abundance and biodiversity (cover, biomass, species richness, diversity, evenness, and community composition) were influenced by tidal height, physical stress, and herbivores. Wave exposure affected all response variables except biomass and evenness. Stress and herbivores had independent additive effects on seaweed abundance and diversity. Physical stress did not make the community as a whole more susceptible to herbivores, and screens had overarching positive effects on seaweed biodiversity even though they also had positive effects on herbivore abundance. Nutrients had virtually no effect on seaweed biodiversity, and we observed no bottom-up effects of nutrient addition on herbivore density or biomass. Small green algae and diatoms were important contributors to overall algal cover and to changes in composition across treatments, but larger macroalgae dominated the species richness response. The striking absence of interactions between stress and herbivory highlights how seaweed communities can respond independently to important drivers of biodiversity. Thus, nonadditive, potentially synergistic effects do not necessarily complicate the understanding of how seaweed biodiversity responds to environmental change.

Experimental tests of the hypothesis that ecological communities can exist in “multiple stable states” are rare, and some argue, impossible, because of the unlikelihood that any system will meet the necessary criteria. These are that alternative states (1) are in the same location, (2) experience the same environment, (3) persist for multiple generations, (4) resist repeated perturbations, and (5) result from a pulse manipulation. In 1974, we initiated an experiment testing the ability of Ascophyllum nodosum-dominated rocky intertidal communities to recover from complete canopy removal. Manipulations were monitored frequently for 5 yr after clearance and resurveyed again after a 35-yr hiatus. After clearance, Ascophyllum was replaced immediately by another fucoid alga, Fucus spp, which continued to dominate the space through 1979 despite regular annual recruitment by Ascophyllum. Observations in 2009 revealed that Fucus spp. still dominated the cleared plots. Surveys in 2011 and 2013 demonstrated that Fucus persisted in experimental plots and Ascophyllum persisted in adjacent, unmanipulated plots. All criteria for testing an alternative state were met. Fucus persisted through multiple generations of both fucoids, a steady annual rain of recruits of both species, a high frequency of storm-driven perturbations, and it resulted from a pulse manipulation. Likely mechanisms include poor Ascophyllum recruitment directly under the Fucus spp. canopy despite abundant recruitment in adjacent areas, self-maintenance by Fucus spp. through high recruitment and fast growth, and recruitment facilitation of Fucus by the barnacle Semibalanus balanoides. Several lines of evidence indicate that other possible mechanisms including indirect facilitation of fucoids through predation on competitors (mussels), and positive or negative effects of littorine grazing, are unlikely. Although prior results in Maine suggested that the mussel Mytilus edulis was an alternative stable state, the new results suggest that Fucus spp. was the alternative state to Ascophyllum. During the nearly four decades of this experiment, a number of important species including fucoids, littorine grazers, and mussels all declined in abundance, most likely due to climate change. The presumed impact of climate change makes prediction of the long-term response of this system difficult, but it already differs dramatically from its structure in the 1970s.

Filter-feeding invertebrates consume phytoplankton and detritus and therefore serve as important mediators of the exchange of materials from nearshore pelagic to intertidal benthic ecosystems. Here, we evaluated the linkages between nearshore and intertidal systems on tem- perate rocky reefs on the coasts of Oregon, USA, and New Zealand's South Island. We used dif- ferences in the concentrations of both nearshore particulate organic carbon and chlorophyll a (chl a), a proxy for phytoplankton availability, at different sites in Oregon and New Zealand to evaluate the influences of suspended particulate organic material (POM) quality and quantity on the rates of carbon inputs associated with intertidal mussels (Mytilus californianus in Oregon and Mytilus galloprovincialis in New Zealand). We also analyzed the carbon stable isotope ratios (δ 13 C) of intertidal mussels and nearshore POM to examine changes in mussel growth in carbon relative to changes in their potential food sources along gradients of POM quality (i.e. carbon-to- chlorophyll ratios, C:chl a). In both Oregon and New Zealand, the δ 13 C in mussel tissues did not change along a gradient of food quality, whereas the δ 13 C of the POM declined as food quality declined (i.e. C:chl a increased), suggesting that mussels were selectively consuming high-quality food. We also found that the availability of phytoplankton, a high-quality component of the POM, was a better predictor of mussel growth in carbon (mg C g �1 d �1 ) than the total concentration of particulate organic carbon, which includes both higher-quality phytoplankton and lower-quality detrital material. Our results highlight the necessity of considering POM quality while evaluating the role of filter-feeders as mediators of carbon inputs into intertidal systems.

Ecology Letters (2012) 15: 461–467 Abstract Predicting the consequences of changes in biodiversity requires understanding both species’ susceptibility to extirpation and their functional roles in ecosystems. However, few studies have evaluated the effects of realistic, non-random biodiversity losses, severely limiting the applicability of biodiversity research to conservation. Here, we removed sessile species from a rocky shore community in a way that deliberately mimicked natural patterns of species loss. We found that the rarest species in the system act from the bottom up to disproportionately impact the diversity and abundance of consumers. Realistic losses of rare species in a diverse assemblage of seaweeds and sessile invertebrates, collectively comprising

Ecology Letters (2011) 14: 852–862 Abstract Synergistic interactions between multiple limiting resources are common, highlighting the importance of co-limitation as a constraint on primary production. Our concept of resource limitation has shifted over the past two decades from an earlier paradigm of single-resource limitation towards concepts of co-limitation by multiple resources, which are predicted by various theories. Herein, we summarise multiple-resource limitation responses in plant communities using a dataset of 641 studies that applied factorial addition of nitrogen (N) and phosphorus (P) in freshwater, marine and terrestrial systems. We found that more than half of the studies displayed some type of synergistic response to N and P addition. We found support for strict definitions of co-limitation in 28% of the studies: i.e. community biomass responded to only combined N and P addition, or to both N and P when added separately. Our results highlight the importance of interactions between N and P in regulating primary producer community biomass and point to the need for future studies that address the multiple mechanisms that could lead to different types of co-limitation.

A recommendation of: Aline Migne, Gaspard Delebecq, Dominique Davoult, Nicolas Spilmont, Dominique Menu, Marie-Andree Janquin and Francois Gevaert Photosynthesis of Laminaria digitata during the immersion and emersion periods of spring tidal cycles during hot, sunny weather https://hal.sorbonne-universite.fr/hal-01827565v4

Extreme events, such as heat waves, are predicted to increase in frequency, duration, and severity as a consequence of climate change. However, global change research generally focuses on increases in mean temperatures and fails to address the potential impacts of increasingly severe heat waves. In addition, climate change may interact with another primary threat to biodiversity, non-native species invasions. We assessed the impacts of a short-term heat wave on the marine epibenthic fouling community of Bodega Harbor, California, USA, by exposing experimental mesocosms to a simulated heat wave in the laboratory and then monitoring community development in the fi eld. We hypothesized that (1) juveniles would be more susceptible to heat waves than adults, (2) native species would be more susceptible than non-native species, and (3) non-native species would recover more quickly than native species. We observed no eff ect of the heat wave on juvenile species richness, either initially or during the recovery period, relative to communities at ambient seawater temperatures. In contrast, total adult species richness initially declined in response to the heat wave. Adult community composition also changed in heat-wave treatments, with non-natives representing the majority of species and occupying more cover than native species. Th e reduction in native richness associated with the heat wave persisted through the recovery period, whereas invasive richness was actually higher on heat-wave versus ambient plates at 95 days. Heat waves have the potential to alter the composition of this community because of species-, taxon-, and/or origin-specifi c responses; for example, nonnative bryozoans displayed greater resistance than native and non-native tunicates. Recovery from the heat wave occurred via growth of resistant individuals and larval recruitment. Our study highlights the importance of considering species ’ and community responses to heat waves, and not just mean predicted temperature increases, to evaluate the consequences of climate change.

Plant-herbivore interactions mediate the trophic structure of ecosystems. We use a comprehensive data set extracted from the literature to test the relative explanatory power of two contrasting bodies of ecological theory, the metabolic theory of ecology (MTE) and ecological stoichiometry (ES), for per-capita and population-level rates of herbivory across ecosystems. We found that ambient temperature and herbivore body size (MTE) as well as stoichiometric mismatch (ES) both constrained herbivory, but at different scales of biological organization. Herbivore body size, which varied over 11 orders of magnitude, was the primary factor explaining variation in per-capita rates of herbivory. Stoichiometric mismatch explained more variation in population-level herbivory rates and also in per-capita rates when we examined data from within functionally similar trophic groups (e.g. zooplankton). Thus, predictions from metabolic and stoichiometric theories offer complementary explanations for patterns of herbivory that operate at different scales of biological organization.

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.

Declines in biodiversity have prompted concern over the consequences of species loss for the goods and services provided by natural ecosystems. However, relatively few studies have evaluated the functional consequences of realistic, nonrandom changes in biodiversity. Instead, most designs have used randomly selected assemblages from a local species pool to construct diversity gradients. It is therefore difficult, based on current evidence, to predict the functional consequences of realistic declines in biodiversity. In this study, we used tide pool microcosms to demonstrate that the effects of real-world changes in biodiversity may be very different from those of random diversity changes. Specifically, we measured the relationship between the diversity of a seaweed assemblage and its ability to use nitrogen, a key limiting nutrient in nearshore marine systems. We quantified nitrogen uptake using both experimental and model seaweed assemblages and found that natural increases in diversity resulted in enhanced rates of nitrogen use, whereas random diversity changes had no effect on nitrogen uptake. Our results suggest that understanding the real-world consequences of declining biodiversity will require addressing changes in species performance along natural diversity gradients and understanding the relationships between species' susceptibility to loss and their contributions to ecosystem functioning.

The cycles of the key nutrient elements nitrogen (N) and phosphorus (P) have been massively altered by anthropogenic activities. Thus, it is essential to understand how photosynthetic production across diverse ecosystems is, or is not, limited by N and P. Via a large-scale meta-analysis of experimental enrichments, we show that P limitation is equally strong across these major habitats and that N and P limitation are equivalent within both terrestrial and freshwater systems. Furthermore, simultaneous N and P enrichment produces strongly positive synergistic responses in all three environments. Thus, contrary to some prevailing paradigms, freshwater, marine and terrestrial ecosystems are surprisingly similar in terms of N and P limitation.

Consumer and resource control of diversity in plant communities have long been treated as alternative hypotheses. However, experimental and theoretical evidence suggests that herbivores and nutrient resources interactively regulate the number and relative abundance of coexisting plant species. Experiments have yielded divergent and often contradictory responses within and among ecosystems, and no effort has to date reconciled this empirical variation within a general framework. Using data from 274 experiments from marine, freshwater, and terrestrial ecosystems, we present a cross-system analysis of producer diversity responses to local manipulations of resource supply and/or herbivory. Effects of herbivory and fertilization on producer richness differed substantially between systems: ( i ) herbivores reduced species richness in freshwater but tended to increase richness in terrestrial systems; ( ii ) fertilization increased richness in freshwater systems but reduced richness on land. Fertilization consistently reduced evenness, whereas herbivores increased evenness only in marine and terrestrial ecosystems. Producer community evenness and ecosystem productivity mediated fertilization and herbivore effects on diversity across ecosystems. Herbivores increased producer richness in more productive habitats and in producer assemblages with low evenness. These same assemblages also showed the strongest reduction in richness with fertilization, whereas fertilization increased (and herbivory decreased) richness in producer assemblages with high evenness. Our study indicates that system productivity and producer evenness determine the direction and magnitude of top-down and bottom-up control of diversity and may reconcile divergent empirical results within and among ecosystems.

Although nutrient availability and oceanographic context can influence top-down processes in intertidal communities, the reciprocal effects of consumers on bottom-up processes, such as nitrogen uptake by macroalgae, are less well-documented. We used a combination of field obser- vations, mesocosm experiments, and laboratory nitrate-uptake measurements to evaluate the inter- action between the kelp crab Pugettia producta and its preferred food source, the kelp Egregia menziesii, in northern California, USA. We found that P. producta fed selectively on E. menziesii, removing tissues with a high surface area to volume ratio (SA:V), which resulted in a 66% decrease in overall SA:V. Because these high SA:V tissues are disproportionately more responsible for nitrate uptake, selective herbivory by P. producta decreased E. menziesii's biomass-specific uptake of nitro- gen, the primary limiting nutrient in this ecosystem, by 65% per gram of remaining algal tissue. In field surveys, where P. producta abundances were high, E. menziesii thalli more frequently exhibited signs of intense grazing and thus had lower SA:V ratios, suggesting that grazing is sufficiently intense in the field to generate differences in algal morphology and nitrate uptake. P. producta there- fore influences E. menziesii biomass and productivity from the top down, by removing substantial amounts of biomass, and from the bottom up, by reducing the kelp's subsequent ability to acquire nutrients. Because of selective grazing on specialized tissues, the effects of mesoherbivores, like P. producta, on the structure and dynamics of marine communities may often be greater than predicted simply by the biomass they consume.

Many ecological phenomena are characterized by context dependency, and the relationship between diversity and productivity is no exception. We examined the re- lationship between macroalgal diversity and nutrient availability by evaluating the effects of reduced nutrients and their subsequent replacement via local-scale nutrient loading in tide pools. Macroalgae in Oregon coast high-intertidal pools have evolved in a nitrate-rich upwelling ecosystem, but instead of settling on low-intertidal reefs (where algae are often immersed in nutrient-rich nearshore waters) these individuals have colonized high-zone pools, where they are isolated from the ocean for extended periods of time and are subjected to extended periods of nitrate depletion. In some pools, this nutrient stress was ameliorated by a positive interaction: the excretion of ammonium by invertebrates. We conducted ex- perimental manipulations to quantify invertebrate-mediated ammonium loading and ma- croalgal ammonium uptake in high-intertidal pools. Variation in tide pool volume and invertebrate biomass created a gradient of local-scale nutrient inputs, allowing us to address the relationship between nitrogen loading and algal diversity. Slow-growing species tolerant of low nitrogen availability were joined by fast-growing species with higher nitrogen re- quirements in pools with higher ammonium loading rates. A fourfold increase in the am- monium loading rate was associated with a doubling in the number of macroalgal species, and macroalgal assemblages in more species-rich pools were characterized by higher rates of biomass-specific ammonium uptake. These patterns contrast with productivity-diversity relationships in terrestrial systems, where local-scale nutrient enrichments generally result in reduced producer diversity due to displacement of subordinate species by aggressive competitors. Our data suggest that the effect of enrichment on diversity is context-depen- dent. Each ecosystem has a critical level of nutrient availability, determined by the level of nutrients typically available in that system. Below this critical level, local-scale nutrient additions increase diversity, but above it, diversity declines with enrichment.

Predicting the outcome of future climate change requires an understanding of how alterations in multiple environmental factors manifest in natural communities and affect ecosystem functioning. We conducted an in situ, fully factorial field manipulation of CO2 and temperature on a rocky shoreline in southeastern Alaska, USA. Warming strongly impacted functioning of tide pool systems within one month, with the rate of net community production (NCP) more than doubling in warmed pools under ambient CO2 levels relative to initial NCP values. However, in pools with added CO2, NCP was unaffected by warming. Productivity responses paralleled changes in the carbon-to-nitrogen ratio of a red alga, the most abundant primary producer species in the system, highlighting the direct link between physiology and ecosystem functioning. These observed changes in algal physiology and community productivity in response to our manipulations indicate the potential for natural systems to shift rapidly in response to changing climatic conditions and for multiple environmental factors to act antagonistically.

Considerable research has examined the influence of herbivores on the maintenance of plant diversity, but fewer studies have examined the reciprocal effect of plant diversity on the animals that use the plant community for food and shelter, particularly in marine systems. Several mechanisms could underlie such effects. Animal diversity and abundance could be increased by complementary use of different plants by different animals, or by an indirect effect of plant diversity on plant production that results in more total plant biomass in high plant-diversity communities. Alternatively, plant species identity could play a dominant role leading to sampling effects or no effect of diversity at all. We conducted a sixyear field manipulation of the richness of rocky shore seaweeds in northern California and measured the effects of algal richness and identity on the invertebrate community, from meiofauna to macrofauna. We found that diverse algal communities hosted more species of both large and small invertebrates than the average algal monoculture but that the mechanisms underlying this pattern differed substantially for organisms of different size. More species of macrofauna occurred in the polycultures than in any of the monocultures, likely due to the greater total cover of algae produced in polycultures. Rare and common macrofaunal taxa responded to host plant species richness in opposite ways, with more occurrences of rare taxa and lower abundance of very common taxa in the polycultures. In contrast, meiofaunal richness in polycultures was no different than that of monocultures of finely branched species, leading to strong effects of algal identity. Our findings are similar to those from terrestrial systems in that the effects of plant diversity we observed were most related to the greater amount of habitat in polycultures as a result of overyielding in algal biomass. However, our findings differ from those in terrestrial systems in that the primary mechanisms for both richness and identity effects appear related to the value of plants as shelter from harsh abiotic conditions or predation rather than food, and in that animal body size altered the mechanisms underlying diversity effects. © 2014 by the Ecological Society of America.

Synergistic interactions between multiple limiting resources are common, highlighting the importance of co-limitation as a constraint on primary production. Our concept of resource limitation has shifted over the past two decades from an earlier paradigm of single-resource limitation towards concepts of co-limitation by multiple resources, which are predicted by various theories. Herein, we summarise multiple-resource limitation responses in plant communities using a dataset of 641 studies that applied factorial addition of nitrogen (N) and phosphorus (P) in freshwater, marine and terrestrial systems. We found that more than half of the studies displayed some type of synergistic response to N and P addition. We found support for strict definitions of co-limitation in 28% of the studies: i.e. community biomass responded to only combined N and P addition, or to both N and P when added separately. Our results highlight the importance of interactions between N and P in regulating primary producer community biomass and point to the need for future studies that address the multiple mechanisms that could lead to different types of co-limitation.

In order for research into the consequences of biodiversity changes to be more applicable to real-world ecosystems, experiments must be conducted in the field, where a variety of factors other than diversity can affect the rates of key biogeochemical and physiological processes. Here, we experimentally evaluate the effects of two factors known to affect the diversity and composition of intertidal seaweed assemblages--tidal elevation and herbivory--on nitrate uptake by those assemblages. Based on surveys of community composition at the end of a 1.5-year press experiment, we found that both tide height and herbivores affected seaweed community structure. Not surprisingly, seaweed species richness was greater at lower tidal elevations. Herbivores did not affect richness, but they altered the types of species that were present; seaweed species characterized by higher rates of nitrate uptake were more abundant in herbivore-removal plots. Both tide height and herbivores affected nitrate uptake by seaweed assemblages. Individual seaweed species, as well as entire seaweed assemblages, living higher on the shore had greater rates of biomass-specific nitrate uptake, particularly at high ambient nitrate concentrations. Grazed seaweed assemblages exhibited reduced nitrate uptake, but only at low nitrate concentrations. We evaluated the effect of seaweed richness on nitrate uptake, both alone and after accounting for effects of tidal elevation and herbivores. When only richness was considered, we found no effect on uptake. However, when simultaneous effects of richness, tide height, and herbivores on uptake were evaluated, we found that all three had relatively large and comparable effects on nitrate uptake coefficients and that there was a negative relationship between seaweed richness and nitrate uptake. Particularly because effects of richness on uptake were not apparent unless the effects of tide height and herbivory were also considered, these results highlight the importance of considering the effects of environmental context when evaluating the consequences of biodiversity change in more realistic systems.