A.: Interhemispheric comparison of recruitment to intertidal communities: Pattern persistence and scales of variation

Abstract. Recruitment variation can be a major source of fluctuation in populations and communities, making it difficult to generalize results. Determining the scales of variation and whether spatial patterns in the supply of individuals are persistent over time can provide insight into spatial generality and the application of conservation and metacommunity models. We examined these issues using eight-year-long data sets of monthly recruitment of intertidal mussels (Mytilus spp., Perumytilus purpuratus, Semimytilus algosus, Brachidontes granulata) and barnacles (Balanus glandula, Chthamalus dalli, Jehlius cirratus, Notochthamalus scabrosus) at sites spanning .900 km along the coasts of Oregon-northern California (OR-NCA, 45.47-39.438 N) and central Chile (CC, 29.5-34.658 S). We evaluated four general ''null'' hypotheses: that despite different phylogenies and great spatial separation of these taxa, their similar life history strategies and environmental settings lead to similar patterns of recruitment (1) between hemispheres, (2) in time, (3) in space, and (4) at larger and smaller spatial scales. Hypothesis 1 was rejected: along the OR-NCA coast, rates of recruitment were between two and three orders of magnitude higher, and patterns of seasonality were generally stronger and more coherent across space and time than along CC. Surprisingly, however, further analysis revealed regularities in both time and space for all species, supporting hypotheses 2 and 3. Temporal decorrelation scales were 1-3 months, and characteristic spatial scales of recruitment were ;250 km. Contrary to hypothesis 4, for the ecologically dominant species in both hemispheres, recruitment was remarkably persistent at larger mesoscales (kilometers) but was highly stochastic at smaller microscales (meters). Across species, increased recruitment variation at large scales was positively associated with increased persistence. Our results have several implications. Although the two regions span distinct latitudinal ranges, potential forcing processes behind these patterns include similar large-scale climates and topographically locked hydrographic features, such as upwelling. Further, spatial persistence of the recruitment patterns of most species at the mesoscale supports the view that marine protected areas can be powerful conservation and management tools. Finally, persistent and yet contrasting spatial patterns of recruitment among competing species suggest that recent metacommunity models might provide useful representations of the mechanisms involved in species coexistence

Designing effective reserve networks for nonequilibrium metacommunities

The proliferation of efficient fishing practices has promoted the depletion of commercial stocks around the world and caused significant collateral damage to marine habitats. Recent empirical studies have shown that marine reserves can play an important role in reversing these effects. Equilibrium metapopulation models predict that networks of marine reserves can provide similar benefits so long as individual reserves are sufficiently large to achieve self-sustainability, or spaced based on the extent of dispersal of the target species in order to maintain connectivity between neighboring reserves. However, these guidelines have not been tested in nonequilibrium metacommunity models that exhibit the kinds of complex spatiotemporal dynamics typically seen in natural marine communities. Here, we used a spatially explicit predator–prey model whose predictions have been validated in a marine system to show that current guidelines are not optimal for metacommunities. In equilibrium metacommunities, there is a community-level trade-off for designing effective reserves: Networks whose size and spacing are smaller than the extent of dispersal maximize global predator abundance but minimize global prey abundance because of trophic cascades, whereas the converse is true for reserve networks whose size and spacing are larger than the extent of dispersal. In nonequilibrium metacommunities, reserves whose size and spacing match the extent of spatial autocorrelation in adult abundance (i.e., the extent of patchiness) escape this community-level trade-off by maximizing global abundance and persistence of both the prey and the predator. Overall, these results suggest that using the extent of adult patchiness instead of the extent of larval dispersal as the size and spacing of reserve networks is critical for designing community-based management strategies. By emphasizing patchiness over dispersal distance, our results show how the apparent complexity of nonequilibrium communities can actually simplify management guidelines and reduce uncertainty associated with the assessment of dispersal in marine environments.Keywords: metacommunities, dynamic resources, spatial management, patchiness, reserve networks, nonequilibrium, trophic cascade

Oceanographic and climatic variation drive top-down/bottom-up coupling in the Galápagos intertidal meta-ecosystem

The impact of herbivores on primary producers in differing oceanographic regimes is a matter of intense ecological interest due to ongoing changes in their abundance, that of their predators, and anthropomorphic alteration of nutrient cycles and climatic patterns. Interactions between productivity and herbivory in marine habitats have been studied on temperate rocky shores, coral reefs, mangroves, and salt marshes, but less so at tropical latitudes. To determine how herbivore–alga dynamics varied with oceanographic regime, we used the comparative-experimental approach in rocky intertidal communities on the Galápagos Islands from January 2006 to January 2009. This setting was selected because strongly contrasting oceanographic conditions occurred within a range of ~181 km, with significant differences in temperature, nutrients, phytoplankton productivity, and intertidal communities, and in abundance of macro-herbivores, including marine iguanas. Experiments and measurements were conducted at two sites in each of three oceanographic regimes characterized by low, intermediate, and high bottom-up inputs. At sites of low inputs, macro-herbivores (fish, crabs, iguanas) had a consistent top-down effect, reducing algal abundance, and leaving a few grazer-resistant varieties. At sites of intermediate and high inputs, consumer impacts were stronger during La Niña (cool phase) than during El Niño (warm phase). At sites of high inputs, algal biomass was naturally relatively high and was dominated by the edible algae Ulva spp. Macro-grazers reduced algal biomass, but their primary effect was indirect, as articulated corallines displaced other species of algae in their absence. Prior results from the tropics had revealed dominant effects of top-down interactions and recruitment in structuring intertidal communities. Our results suggest that, when a broader oceanographic scenario is taken into account, the relative importance of top-down and bottom-up forces are context dependent, varying with oceanographic regime and climatic variability.Keywords: Galápagos, Algae, Tropics, Bottom-up, Rocky shores, Climate, Productivity, Top-down, Oceanographic conditions, Herbivory, ENSO, Marine iguana

The Dynamics and Impact of Ocean Acidification and Hypoxia: Insights from Sustained Investigations in the Northern California Current Large Marine Ecosystem

Coastal upwelling ecosystems around the world are defined by wind-generated currents that bring deep, nutrient-rich waters to the surface ocean where they fuel exceptionally productive food webs. These ecosystems are also now understood to share a common vulnerability to ocean acidification and hypoxia (OAH). In the California Current Large Marine Ecosystem (CCLME), reports of marine life die-offs by fishers and resource managers triggered research that led to an understanding of the risks posed by hypoxia. Similarly, unprecedented losses from shellfish hatcheries led to novel insights into the coastal expression of ocean acidification. Partnership for Interdisciplinary Studies of Coastal Oceans (PISCO) scientists and other researchers in the CCLME responded to the rise of OAH with new ocean observations and experiments. This work revealed insights into the expression of OAH as coupled environmental stressors, their temporal and spatial variability, and impacts on species, ecological communities, and fisheries. Sustained investigations also deepened the understanding of connections between climate change and the intensification of hypoxia, and are beginning to inform the ecological and eco-evolutionary processes that can structure responses to the progression of ocean acidification and other pathways of global change. Moreover, because the severity of the die-offs and hatchery failures and the subsequent scientific understanding combined to galvanize public attention, these scientific advances have fostered policy advances. Across the CCLME, policymakers are now translating the evolving scientific understanding of OAH into new management actions

Wind-driven inner-shelf circulation off central Oregon during summer

Velocity measurements from 17 deployments of moored acoustic Doppler current profilers obtained during four summer upwelling seasons are used to describe the crossshelf divergence of Ekman transport in the inner shelf off Oregon. For each deployment the measured surface and bottom cross-shelf transports were compared with estimates of the theoretical Ekman transports to find the fraction of full theoretical Ekman transport present. In general, in 15 m of water at 1–2 km offshore, measured transport was 25% of the full Ekman transport. Measured transports reached full Ekman transport 5–6 km offshore in 50 m of water. This result indicates that the region of active upwelling marked by the divergence of Ekman transport was limited to a narrow region along the coast. With small wind stress curl and no major headlands in the region, no along-shelf trends in the transport fractions were observed. Average transport fractions at each station were similar from year to year with one exception. The interannual variability seen at this particular site was most likely a result of local along-shelf bathymetric features. In addition, a weak linear relationship was found between the ambient stratification and the fraction of full Ekman transport. Reduced cross-shelf transport occurred at times of decreased stratification. This type of ‘‘shutdown’’ of the inner-shelf cross-shelf circulation has significant biological implications, sequestering production in the nearshore and reducing larval cross-shelf transport

The complex net effect of reciprocal interactions and recruitment facilitation maintains an intertidal kelp community

1. Theoretical and empirical ecology has transitioned from a focus on the role of negative interactions in species coexistence to a more pluralistic view that acknowledges that coexistence in natural communities is more complex, and depends on species interactions that vary in strength, sign, and reciprocity, and such contexts as the environment and life-history stage. 2. We used a whole-community approach to examine how species interactions contribute to the maintenance of a rocky intertidal macroalgal canopy–understorey assemblage. We determined both the types of interactions in this network, and whether interactions were sensitive to environmental gradients. 3. Focusing on a structurally dominant canopy kelp Saccharina sessilis, and its diverse co-occurring understorey assemblage, we evaluated the role of the understorey in controlling S. sessilis recruitment and quantified the reciprocal effect of the S. sessilis canopy and understorey on one another using a removal experiment replicated across 600 km of coastline. We determined the sensitivity of interactions to natural variation in light and nutrient availability (replicated among four regions on the N.E. Pacific coast), and under different wave conditions (three wave regimes). 4. Surprisingly, species interactions were similar across sites and thus not context-dependent. Unexpectedly, the understorey community had a strong positive effect on the S. sessilis canopy, whereby the adult canopy decreased dramatically following understorey removal. Additionally, S. sessilis recruitment depended on the presence of understorey coralline algal turf. In turn, the canopy had a neutral effect on the coralline understorey, but a negative effect on non-calcifying algal turfs, likely eventually generating positive indirect canopy effects on the coralline understorey. Density-dependent intraspecific competition between S. sessilis adults and recruits may moderate this positive feedback between the S. sessilis canopy and coralline understorey. 5. Synthesis. Our research highlights the importance of positive interactions for coexistence in natural communities, and the necessity of studying multiple life-history stages and reciprocal species interactions in order to elucidate the mechanisms that maintain diversity.Keywords: marine, plant-plant interactions, aquatic plant ecology, positive interactions, environmental gradients, benthic, macroalga

Benthic-Pelagic Links and Rocky Intertidal Communities: Bottom-Up Effects on Top-Down Control?

Insight into the dependence of benthic communities on biological and physical processes in nearshore pelagic environments, long considered a ``black box,'' has eluded ecologists. In rocky intertidal communities at Oregon coastal sites 80 km apart, differences in abundance of sessile invertebrates, herbivores, carnivores, and macrophytes in the low zone were not readily explained by local scale differences in hydrodynamic or physical conditions (wave forces, surge flow, or air temperature during low tide). Field experiments employing predator and herbivore manipulations and prey transplants suggested top-down (predation, grazing) processes varied positively with bottom-up processes (growth of filter-feeders, prey recruitment), but the basis for these differences was unknown. Shore-based sampling revealed that between-site differences were associated with nearshore oceanographic conditions, including phytoplankton concentration and productivity, particulates, and water temperature during upwelling. Further, samples taken at 19 sites along 380 km of coastline suggested that the differences documented between two sites reflect broader scale gradients of phytoplankton concentration. Among several alternative explanations, a coastal hydrodynamics hypothesis, reflecting mesoscale (tens to hundreds of kilometers) variation in the interaction between offshore currents and winds and continental shelf bathymetry, was inferred to be the primary underlying cause. Satellite imagery and offshore chlorophyll-a samples are consistent with the postulated mechanism. Our results suggest that benthic community dynamics can be coupled to pelagic ecosystems by both trophic and transport linkages.KEYWORDS: scale, productivity, currents, upwelling, community structureThis is the publisher’s final pdf. The published article is copyrighted by the National Academy of Sciences of the United States of America and can be found at: http://www.pnas.org

Are meta-ecosystems organized hierarchically? A model and test in rocky intertidal habitats

Ecosystems are shaped by processes occurring and interacting over multiple temporal and spatial scales. Theory suggests such complexity can be simplified by focusing on processes sharing the same scale as the pattern of interest. This scale-dependent approach to studying communities has been challenged by multiscale meta-ecosystem theory, which recognizes that systems are interconnected by the movement of ‘‘ecological subsidies’’ and suggests that cross-scale feedbacks between local and regional processes can be equally important for understanding community structure. We reconcile these two perspectives by developing and testing a hierarchical meta-ecosystem model. The model predicts local community responses to connectivity over multiple oceanographic spatial scales, defined as macro- (100s of km), meso- (10s of km), and local scale (100s of m). It assumes that local communities occur in distinct regions and that connectivity effects are strongest among local sites. Predictions are that if macroscale processes dominate, then regardless of mesoscale differences, (1) local communities will be similar, and (2) will be even more so with increased connectivity. With dominance of mesoscale (i.e., regional) processes, (3) local structure will be similar within but distinct between regions, and (4) with increased connectivity similar both within and among regions. With dominance of local-scale processes, (5) local communities will differ both within and among regions, and (6) with increased connectivity be similar within but not between regions. We tested the model by evaluating rocky intertidal community structure patterns with variation in ecological subsidies and environmental conditions at 13 sites spanning 725 km of the northern California Current system. External factors operating at meso- and local scales had strong effects, explaining 52% and 27% of the variance, respectively, in community structure. Sessile invertebrate and predator dominance was associated with weaker upwelling, higher phytoplankton abundance, and higher recruitment, and the opposite was true for macrophyte dominance. Overall, our results support the theory that meta-ecosystems are organized hierarchically, with environmental processes dominating at meso- to macroscales and ecological processes playing a more important role at local scales, but with important bidirectional cross-scale interactions.Keywords: variation partitioning, meta-ecosystems, ecological subsidies, relative importance, spatial scale, oceanographic conditions, coastal ecosystems, rocky intertidal communities, ecosystem dynamics, northern California Current large marine ecosyste

Robert Treat Paine III, 1933–2016

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Sea Star Wasting Disease in the Keystone Predator Pisaster ochraceus in Oregon: Insights into Differential Population Impacts, Recovery, Predation Rate, and Temperature Effects from Long-Term Research

Sea star wasting disease (SSWD) first appeared in Oregon in April 2014, and by June had spread to most of the coast. Although delayed compared to areas to the north and south, SSWD was initially most intense in north and central Oregon and spread southward. Up to 90% of individuals showed signs of disease from June-August 2014. In rocky intertidal habitats, populations of the dominant sea star Pisaster ochraceus were rapidly depleted, with magnitudes of decline in density among sites ranging from -2x to -9x (59 to 84%) and of biomass from -2.6x to -15.8x (60 to 90%) by September 2014. The frequency of symptomatic individuals declined over winter and persisted at a low rate through the spring and summer 2015 (~5–15%, at most sites) and into fall 2015. Disease expression included six symptoms: initially with twisting arms, then deflation and/or lesions, lost arms, losing grip on substrate, and final disintegration. SSWD was disproportionally higher in orange individuals, and higher in tidepools. Although historically P. ochraceus recruitment has been low, from fall 2014 to spring 2015 an unprecedented surge of sea star recruitment occurred at all sites, ranging from ~7x to 300x greater than in 2014. The loss of adult and juvenile individuals in 2014 led to a dramatic decline in predation rate on mussels compared to the previous two decades. A proximate cause of wasting was likely the “Sea Star associated Densovirus” (SSaDV), but the ultimate factors triggering the epidemic, if any, remain unclear. Although warm temperature has been proposed as a possible trigger, SSWD in Oregon populations increased with cool temperatures. Since P. ochraceus is a keystone predator that can strongly influence the biodiversity and community structure of the intertidal community, major community-level responses to the disease are expected. However, predicting the specific impacts and time course of change across west coast meta-communities is difficult, suggesting the need for detailed coast-wide investigation of the effects of this outbreak