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  1. Abstract It is critical to understand how human modifications of Earth’s ecosystems are influencing ecosystem functioning, including net and gross community production ( NCP and GCP , respectively) and community respiration ( CR ). These responses are often estimated by measuring oxygen production in the light ( NCP ) and consumption in the dark ( CR ), which can then be combined to estimate GCP . However, the method used to create “dark” conditions—either experimental darkening during the day or taking measurements at night—could result in different estimates of respiration and production, potentially affecting our ability to make integrative predictions. We tested this possibility by measuring oxygen concentrations under daytime ambient light conditions, in darkened tide pools during the day, and during nighttime low tides. We made measurements every 1–3 months over one year in southeastern Alaska. Daytime respiration rates were substantially higher than those measured at night, associated with higher temperature and oxygen levels during the day and leading to major differences in estimates of GCP calculated using daytime versus nighttime measurements. Our results highlight the potential importance of measuring respiration rates during both day and night to account for effects of temperature and oxygen—especially in shallow-water, constrained systems—with implications for understanding the impacts of global change on ecosystem metabolism. 
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  2. Abstract

    High community diversity may either prevent or promote the establishment of exotic species. The biotic resistance hypothesis holds that species-rich communities are more resistant to invasion than species-poor communities due to mechanisms including greater interspecific competition. Conversely, the invasional meltdown hypothesis proposes that greater exotic diversity increases invasibility via facilitative interactions between exotic species. To evaluate the degree to which biotic resistance or invasional meltdown influences marine community structure during the assembly period, we studied the development of marine epibenthic “fouling” communities at two southern California harbors. With a focus on sessile epibenthic species, we found that fewer exotic species established as total and exotic richness increased during community assembly and that this effect remained after accounting for space availability. We also found that changes in exotic abundance decreased over time. Throughout the assembly period, gains in exotic abundance were greatest when space was abundant and richness was low. Altogether, we found greater support for biotic resistance than invasional meltdown, suggesting that both native and exotic species contribute to biotic resistance during early development of these communities. However, our results indicate that biotic resistance may not always reduce the eventual dominance of exotic species.

     
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  3. Abstract

    Globally, species are undergoing range shifts in response to climate change. However, the potential impacts of climate-driven range shifts are not well understood. In southern California, the predatory whelkMexacanthina lugubrishas undergone a northward range shift of more than 100 km in the past four decades. We traced the history of the whelk’s range shift and assessed potential effects using an integrated approach, consisting of field surveys, as well as feeding and thermotolerance experiments. We found that at sites whereMexacanthinaand native species co-occurred, native whelks distributions peaked lower in the intertidal. In laboratory experiments, we found that the presence ofMexacanthinaled to reduced growth in native whelks (Acanthinucella spirata). Additionally, the range-shifting whelk was able to tolerate higher temperatures than common native species (A. spirataandNucella emarginata), suggesting further impacts as a result of climate warming. Many species are likely to undergo range shifts as a coping mechanism for changing climatic conditions. However, communities are unlikely to shift as a whole due to species-specific responses. By studying the impacts of range-shifting species, likeMexacanthina, we can better understand how climate change will alter existing community structure and composition.

     
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  4. Abstract

    Biological processes play important roles in determining how global changes manifest at local scales. Primary producers can absorb increased CO2via daytime photosynthesis, modifying pH in aquatic ecosystems. Yet producers and consumers also increase CO2via respiration. It is unclear whether biological modification of pH differs across the year, and, if so, what biotic and abiotic drivers underlie temporal differences. We addressed these questions using the intensive study of tide pool ecosystems in Alaska, USA, including quarterly surveys of 34 pools over 1 year and monthly surveys of five pools from spring to fall in a second year. We measured physical conditions, community composition, and changes in pH and dissolved oxygen during the day and night. We detected strong temporal patterns in pH dynamics. Our measurements indicate that pH modification varies spatially (between tide pools) and temporally (across months). This variation in pH dynamics mirrored changes in dissolved oxygen and was associated with community composition, including both relative abundance and diversity of benthic producers and consumers, whose role differed across the year, particularly at night. These results highlight the importance of the time of year when considering the ways that community composition influences pH conditions in aquatic ecosystems.

     
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  5. Species introduced through human-related activities beyond their native range, termed alien species, have various impacts worldwide. The IUCN Environmental Impact Classification for Alien Taxa (EICAT) is a global standard to assess negative impacts of alien species on native biodiversity. Alien species can also positively affect biodiversity (for instance, through food and habitat provisioning or dispersal facilitation) but there is currently no standardized and evidence-based system to classify positive impacts. We fill this gap by proposing EICAT+, which uses 5 semiquantitative scenarios to categorize the magnitude of positive impacts, and describes underlying mechanisms. EICAT+ can be applied to all alien taxa at different spatial and organizational scales. The application of EICAT+ expands our understanding of the consequences of biological invasions and can inform conservation decisions. 
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  6. Abstract

    Impacts of increases in extreme heat events under climate change may differ across ontogeny for species with complex life cycles. Different life stages may (1) experience unequal levels of environmental stress that vary across space and time (exposure) and (2) have different stress tolerances (sensitivity). We used a field experiment to investigate whether exposure, sensitivity, and overall mortality risk differed between life stages of a marine foundation species (the musselMytilus californianus) across thermal conditions that vary in space (habitat, elevation, and site) and time (season) in southern California, USA. We deployed temperature loggers to document habitat‐specific exposure patterns, conducted laboratory thermal tolerance assays to calculate sensitivity, and performed field surveys to determine whether risk patterns were reflected in distributions. Exposure to extreme temperatures was highest in solitary habitats and during spring. Juvenile mussels were more sensitive to extreme heat than adults, and sensitivity for both life stages was highest in December and March. Risk was largely seasonal for juveniles but was more temporally variable for adults. Spatial occurrence patterns were congruent with risk assessments for both life stages (i.e., higher occurrence in lower risk habitats). These results highlight the importance of incorporating life stage and temporal dynamics when predicting impacts of climate change.

     
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  7. Abstract

    Invasive species science has focused heavily on the invasive agent. However, management to protect native species also requires a proactive approach focused on resident communities and the features affecting their vulnerability to invasion impacts. Vulnerability is likely the result of factors acting across spatial scales, from local to regional, and it is the combined effects of these factors that will determine the magnitude of vulnerability. Here, we introduce an analytical framework that quantifies the scale‐dependent impact of biological invasions on native richness from the shape of the native species–area relationship (SAR). We leveraged newly available, biogeographically extensive vegetation data from the U.S. National Ecological Observatory Network to assess plant community vulnerability to invasion impact as a function of factors acting across scales. We analyzed more than 1000 SARs widely distributed across the USA along environmental gradients and under different levels of non‐native plant cover. Decreases in native richness were consistently associated with non‐native species cover, but native richness was compromised only at relatively high levels of non‐native cover. After accounting for variation in baseline ecosystem diversity, net primary productivity, and human modification, ecoregions that were colder and wetter were most vulnerable to losses of native plant species at the local level, while warmer and wetter areas were most susceptible at the landscape level. We also document how the combined effects of cross‐scale factors result in a heterogeneous spatial pattern of vulnerability. This pattern could not be predicted by analyses at any single scale, underscoring the importance of accounting for factors acting across scales. Simultaneously assessing differences in vulnerability between distinct plant communities at local, landscape, and regional scales provided outputs that can be used to inform policy and management aimed at reducing vulnerability to the impact of plant invasions.

     
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  8. null (Ed.)
    Abstract 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. 
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  9. Abstract

    Ocean warming, acidification, deoxygenation and reduced productivity are widely considered to be the major stressors to ocean ecosystems induced by emissions ofCO2. However, an overlooked stressor is the change in ocean circulation in response to climate change. Strong changes in the intensity and position of the western boundary currents have already been observed, and the consequences of such changes for ecosystems are beginning to emerge. In this study, we address climatically induced changes in ocean circulation on a global scale but relevant to propagule dispersal for species inhabiting global shelf ecosystems, using a high‐resolution global ocean model run under theIPCC RCP8.5 scenario. The ¼ degree model resolution allows improved regional realism of the ocean circulation beyond that of availableCMIP5‐class models. We use a Lagrangian approach forced by modelled ocean circulation to simulate the circulation pathways that disperse planktonic life stages. Based on trajectory backtracking, we identify present‐day coastal retention, dominant flow and dispersal range for coastal regions at the global scale. Projecting into the future, we identify areas of the strongest projected circulation change and present regional examples with the most significant modifications in their dominant pathways. Climatically induced changes in ocean circulation should be considered as an additional stressor of marine ecosystems in a similar way to ocean warming or acidification.

     
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