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Abstract Structure‐forming foundation species facilitate consumers by providing habitat and refugia. In return, consumers can benefit foundation species by reducing top‐down pressures and increasing the supply of nutrients. Consumer‐mediated nutrient dynamics (CND) fuel the growth of autotrophic foundation species and generate more habitat for consumers, forming reciprocal feedbacks. Such feedbacks are threatened when foundation species are lost to disturbances, yet testing these interactions requires long‐term studies, which are rare. Here, we experimentally evaluated how disturbance to giant kelp, a marine foundation species, affects (1) CND of the forest animal community and (2) nutrient feedbacks that help sustain forest primary production during extended periods of low nitrate. Our experiment involved removing giant kelp annually during the winter for 10 years at four sites to mimic frequent wave disturbance. We paired temporal changes in the forest community in kelp removal and control plots with estimates of taxon‐specific ammonium excretion rates (reef fishes and macroinvertebrates) and nitrogen (N) demand (giant kelp and understory macroalgae) to determine the effects of disturbance on CND as measured by ammonium excretion, N demand by kelp forest macroalgae, and the percentage of nitrogen demand met by ammonium excretion. We found that disturbance to giant kelp decreased ammonium excretion by 66% over the study, mostly due to declines in fishes. Apart from a few fish species that dominated CND, most reef‐associated consumers were unaffected by disturbance. Disturbance to giant kelp reduced its N demand by 56% but increased that of the understory by 147% due to its increased abundance in the absence of a kelp canopy. Overall, disturbance had little effect on the fraction of N demand of macroalgae met by consumer excretion due to the offsetting responses of giant kelp, understory macroalgae, and consumers to disturbance. Across both disturbance regimes, on average, consumers supported 11%–12% of the N required by all kelp forest macroalgae and 48% of N demand by the understory macroalgae (which are confined to the benthos where most reef‐associated consumers reside). Our findings suggest that CND constitutes a considerable contribution of N required in kelp forests, yet nutrient inputs decrease following reductions in essential habitat perpetuated by frequent disturbances. 

Abstract Marine protected areas (MPAs) are widely implemented tools for long‐term ocean conservation and resource management. Assessments of MPA performance have largely focused on specific ecosystems individually and have rarely evaluated performance across multiple ecosystems either in an individual MPA or across an MPA network. We evaluated the conservation performance of 59 MPAs in California's large MPA network, which encompasses 4 primary ecosystems (surf zone, kelp forest, shallow reef, deep reef) and 4 bioregions, and identified MPA attributes that best explain performance. Using a meta‐analytic framework, we evaluated the ability of MPAs to conserve fish biomass, richness, and diversity. At the scale of the network and for 3 of 4 regions, the biomass of species targeted by fishing was positively associated with the level of regulatory protection and was greater inside no‐take MPAs, whereas species not targeted by fishing had similar biomass in MPAs and areas open to fishing. In contrast, species richness and diversity were not as strongly enhanced by MPA protection. The key features of conservation effectiveness included MPA age, preimplementation fisheries pressure, and habitat diversity. Important drivers of MPA effectiveness for single MPAs were consistent across MPAs in the network, spanning regions and ecosystems. With international targets aimed at protecting 30% of the world's oceans by 2030, MPA design and assessment frameworks should consider conservation performance at multiple ecologically relevant scales, from individual MPAs to MPA networks. 

The unusual blue color polymorphism of lingcod ( Ophiodon elongatus ) is the subject of much speculation but little empirical research; ~20% of lingcod individuals exhibit this striking blue color morph, which is discrete from and found within the same populations as the more common brown morph. In other species, color polymorphisms are intimately linked with host–parasite interactions, which led us to ask whether blue coloration in lingcod might be associated with parasitism, either as cause or effect. To test how color and parasitism are related in this host species, we performed parasitological dissection of 89 lingcod individuals collected across more than 26 degrees of latitude from Alaska, Washington, and California, USA. We found that male lingcod carried 1.89 times more parasites if they were blue than if they were brown, whereas there was no difference in parasite burden between blue and brown female lingcod. Blue individuals of both sexes had lower hepatosomatic index (i.e., relative liver weight) values than did brown individuals, indicating that blueness is associated with poor body condition. The immune systems of male vertebrates are typically less effective than those of females, due to the immunocompromising properties of male sex hormones; this might explain why blueness is associated with elevated parasite burdens in males but not in females. What remains to be determined is whether parasites induce physiological damage that produces blueness or if both blue coloration and parasite burden are driven by some unmeasured variable, such as starvation. Although our study cannot discriminate between these possibilities, our data suggest that the immune system could be involved in the blue color polymorphism–an exciting jumping-off point for future research to definitively identify the cause of lingcod blueness and a hint that immunocompetence and parasitism may play a role in lingcod population dynamics. 

Abstract Understanding species’ responses to upwelling may be especially important in light of ongoing environmental change. Upwelling frequency and intensity are expected to increase in the future, while ocean acidification and deoxygenation are expected to decrease the pH and dissolved oxygen (DO) of upwelled waters. However, the acute effects of a single upwelling event and the integrated effects of multiple upwelling events on marine organisms are poorly understood. Here, we use in situ measurements of pH, temperature, and DO to characterize the covariance of environmental conditions within upwelling‐dominated kelp forest ecosystems. We then test the effects of acute (0–3 days) and chronic (1–3 months) upwelling on the performance of two species of kelp forest grazers, the echinoderm,Mesocentrotus franciscanus, and the gastropod,Promartynia pulligo. We exposed organisms to static conditions in a regression design to determine the shape of the relationship between upwelling and performance and provide insights into the potential effects in a variable environment. We found that respiration, grazing, growth, and net calcification decline linearly with increasing upwelling intensity forM.francicanusover both acute and chronic timescales.Promartynia pulligoexhibited decreased respiration, grazing, and net calcification with increased upwelling intensity after chronic exposure, but we did not detect an effect over acute timescales or on growth after chronic exposure. Given the highly correlated nature of pH, temperature, and DO in the California Current, our results suggest the relationship between upwelling intensity and growth in the 3‐month trial could potentially be used to estimate growth integrated over long‐term dynamic oceanographic conditions forM.franciscanus. Together, these results indicate current exposure to upwelling may reduce species performance and predicted future increases in upwelling frequency and intensity could affect ecosystem function by modifying the ecological roles of key species. 

Abstract Calls for using marine protected areas (MPAs) to achieve goals for nature and people are increasing globally. While the conservation and fisheries impacts of MPAs have been comparatively well‐studied, impacts on other dimensions of human use have received less attention. Understanding how humans engage with MPAs and identifying traits of MPAs that promote engagement is critical to designing MPA networks that achieve multiple goals effectively, equitably and with minimal environmental impact.In this paper, we characterize human engagement in California's MPA network, the world's largest MPA network scientifically designed to function as a coherent network (124 MPAs spanning 16% of state waters and 1300 km of coastline) and identify traits associated with higher human engagement. We assemble and compare diverse indicators of human engagement that capture recreational, educational and scientific activities across California's MPAs.We find that human engagement is correlated with nearby population density and that site “charisma” can expand human engagement beyond what would be predicted based on population density alone. Charismatic MPAs tend to be located near tourist destinations, have long sandy beaches and be adjacent to state parks and associated amenities. In contrast, underutilized MPAs were often more remote and lacked both sandy beaches and parking lot access.Synthesis and applications: These results suggest that achieving MPA goals associated with human engagement can be promoted by developing land‐based amenities that increase access to coastal MPAs or by locating new MPAs near existing amenities during the design phase. Alternatively, human engagement can be limited by locating MPAs in areas far from population centres, coastal amenities or sandy beaches. Furthermore, managers may want to prioritize monitoring, enforcement, education and outreach programmes in MPAs with traits that predict high human engagement. Understanding the extent to which human engagement impacts the conservation performance of MPAs is a critical next step to designing MPAs that minimize tradeoffs among potentially competing objectives. Read the freePlain Language Summaryfor this article on the Journal blog. 

Abstract Marine protected areas (MPAs) have gained attention as a conservation tool for enhancing ecosystem resilience to climate change. However, empirical evidence explicitly linking MPAs to enhanced ecological resilience is limited and mixed. To better understand whether MPAs can buffer climate impacts, we tested the resistance and recovery of marine communities to the 2014–2016 Northeast Pacific heatwave in the largest scientifically designed MPA network in the world off the coast of California, United States. The network consists of 124 MPAs (48 no‐take state marine reserves, and 76 partial‐take or special regulation conservation areas) implemented at different times, with full implementation completed in 2012. We compared fish, benthic invertebrate, and macroalgal community structure inside and outside of 13 no‐take MPAs across rocky intertidal, kelp forest, shallow reef, and deep reef nearshore habitats in California's Central Coast region from 2007 to 2020. We also explored whether MPA features, including age, size, depth, proportion rock, historic fishing pressure, habitat diversity and richness, connectivity, and fish biomass response ratios (proxy for ecological performance), conferred climate resilience for kelp forest and rocky intertidal habitats spanning 28 MPAs across the full network. Ecological communities dramatically shifted due to the marine heatwave across all four nearshore habitats, and MPAs did not facilitate habitat‐wide resistance or recovery. Only in protected rocky intertidal habitats did community structure significantly resist marine heatwave impacts. Community shifts were associated with a pronounced decline in the relative proportion of cold water species and an increase in warm water species. MPA features did not explain resistance or recovery to the marine heatwave. Collectively, our findings suggest that MPAs have limited ability to mitigate the impacts of marine heatwaves on community structure. Given that mechanisms of resilience to climate perturbations are complex, there is a clear need to expand assessments of ecosystem‐wide consequences resulting from acute climate‐driven perturbations, and the potential role of regulatory protection in mitigating community structure changes. 

Abstract The changing global climate is having profound effects on coastal marine ecosystems around the world. Structure, functioning, and resilience, however, can vary geographically, depending on species composition, local oceanographic forcing, and other pressures from human activities and use. Understanding ecological responses to environmental change and predicting changes in the structure and functioning of whole ecosystems require large‐scale, long‐term studies, yet most studies trade spatial extent for temporal duration. We address this shortfall by integrating multiple long‐term kelp forest monitoring datasets to evaluate biogeographic patterns and rates of change of key functional groups (FG) along the west coast of North America. Analysis of data from 469 sites spanning Alaska, USA, to Baja California, Mexico, and 373 species (assigned to 18 FG) reveals regional variation in responses to both long‐term (2006–2016) change and a recent marine heatwave (2014–2016) associated with two atmospheric and oceanographic anomalies, the “Blob” and extreme El Niño Southern Oscillation (ENSO). Canopy‐forming kelps appeared most sensitive to warming throughout their range. Other FGs varied in their responses among trophic levels, ecoregions, and in their sensitivity to heatwaves. Changes in community structure were most evident within the southern and northern California ecoregions, while communities in the center of the range were more resilient. We report a poleward shift in abundance of some key FGs. These results reveal major, ongoing region‐wide changes in productive coastal marine ecosystems in response to large‐scale climate variability, and the potential loss of foundation species. In particular, our results suggest that coastal communities that are dependent on kelp forests will be more impacted in the southern portion of the California Current region, highlighting the urgency of implementing adaptive strategies to sustain livelihoods and ensure food security. The results also highlight the value of multiregional integration and coordination of monitoring programs for improving our understanding of marine ecosystems, with the goal of informing policy and resource management in the future.