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Creators/Authors contains: "Burkepile, Deron E"

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  1. The data included in this data package were collected in the lagoon of Moorea, French Polynesia. Data on coral mortality were collected in July 2019. Data on nitrogen content (%N) in the long-lived brown macroalga Turbinaria ornata were collected during six sampling campaigns between January 2016 and May 2021 to characterize nitrogen availability at each site. Data on seawater temperatures at six LTER sites in the back reef were collected from 2005-2019, but temperature loggers at two sites (LTER 1 and LTER 6) only recorded partial ocean temperature records during the heatwave from 2018-2019. We used our entire time series (spanning from 2005 to 2018) of temperature across our four long-term sites for which we have data in 2019 to predict ocean temperature data for LTER 1 and LTER 6 in 2019. These data were used for analyses in the manuscript entitled "Effects of nitrogen enrichment on coral mortality depend on the intensity of heat stress". 
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  2. Rodil, Ivan (Ed.)
    Abstract Understanding the mechanisms underlying nutrient (nitrogen and phosphorus) and carbon cycling in reefs is critical for effective management. Research on reef nutrient and carbon cycling needs to account for (i) the contributions of multiple organisms, (ii) abiotic and biotic drivers, and (iii) a social-ecological perspective. In this paper, we review the mechanisms underlying nutrient and carbon cycling in reef social-ecological systems and analyse them using causal loop analysis. We identify direct and indirect pathways and feedback loops through nutrient and carbon cycles that shape the dominant benthic state of reefs: coral, algal, and sponge-dominated states. We find that two of three anthropogenic impact scenarios (size-selective fishing and land use change) have primarily negative consequences for coral and macroalgae via the nutrient and carbon cycles. A third scenario (runoff) has fewer negative impacts on sponges compared to other benthos. In all scenarios, frequent positive feedback loops (size-selective fishing: 7 of 12 loops; runoff: 6 of 9 loops; land use change: 8 of 11 loops) lead to system destabilization; however, the presence of multiple loops introduces avenues whereby reefs may retain coral dominance despite anthropogenic pressures. Context-specific information on the relative strength of loops will be necessary to predict future reef state. 
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  3. 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. 
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  4. Abstract Cryptic species (evolutionarily distinct lineages that do not align with morphologically defined species) are being increasingly discovered but are poorly integrated into ecological theory. In particular, we still lack a useful understanding of if and how cryptic species differ in ways that affect community recovery from disturbances and responses to anthropogenic stressors, such as the removal of consumers and pollution from nutrients. On coral reefs, nutrient pollution increases the growth of macroalgae that displace corals. Reductions in herbivorous fishes reduce the suppression of macroalgae, while reductions in coralivorous fishes reduce predation on corals. An unresolved question is if and how cryptic coral species respond differently to these impacts, thereby differing in their ability to influence coral community dynamics and maintain coral dominance. Therefore, we assessed how the response of crypticPocilloporaspecies over a period of three years following a simulated disturbance from a cyclone depended on the experimental reduction of fish consumer pressure and nutrient addition. After three years, five morphologically cryptic, but genetically distinct,Pocilloporaspecies recruited to the reef. However, recruitment was dominated by two species:Pocillopora tuahiniensis(46%) andPocillopora meandrina(43%). Under ambient conditions, recruitment ofP. tuahiniensisandP. meandrinawas similar, but experimentally reducing consumer pressure increased recruitment ofP. tuahiniensisby up to 73% and reduced recruitment ofP. meandrinaby up to 49%. In both species, nutrient enrichment increased recruitment and colony growth rates equally, but colonies ofP. tuahiniensisgrew faster and were up to 25% larger after three years than those ofP. meandrina,and growth was unaffected by reduced consumer pressure. Predation by excavating corallivorous fish was higher forP. meandrinathan forP. tuahiniensis, especially under nutrient enrichment. In contrast, polyp extension (an indicator of elevated heterotrophic feeding as well as susceptibility and attractiveness to corallivores) was lower forP. meandrinathan forP. tuahiniensis, especially under low to medium consumer pressure. Overall, we uncovered ecological differences in the response of morphologically cryptic foundation species to two pervasive stressors on coral reefs. Our results demonstrate how cryptic species respond differently to key anthropogenic stressors, which may contribute to response diversity that can support ecological resilience or increase extinction risk. 
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  5. Abstract Understanding how foundation species recover from disturbances is key for predicting the future of ecosystems in the Anthropocene. Coral reefs are dynamic ecosystems that can undergo rapid declines in coral abundance following disturbances. Understanding why some reefs recover quickly from these disturbances whereas others recover slowly (or not at all) gives insight into the drivers of community resilience. From 2006 to 2010 coral reefs on the fore reef of Moorea, French Polynesia, experienced severe disturbances that reduced coral cover from ~46% in 2005 to <1% in 2010. Following these disturbances, coral cover increased from 2010 to 2018. Although there was a rapid and widespread recovery of corals, reefs at 17 m depth recovered more slowly than reefs at 10 m depth. We investigated the drivers of different rates of coral recovery between depths from 2010 to 2018 using a combination of time‐series data on coral recruitment, density, growth, and mortality in addition to field experiments testing for the effects of predation. Propagule abundance did not influence recovery, as the density of coral recruits (spat <6 months old) did not differ between depths. However, mortality of juvenile corals (≤5 cm diameter) was higher at 17 m, leading to densities of juvenile corals 3.5 times higher at 10 m than at 17 m depth. Yet, there were no differences in the growth of corals between depths. These results point to an early life stage bottleneck after settlement, resulting in greater mortality at 17 m than at 10 m as the likely driver of differential coral recovery between depths. We used experiments and time‐series data to test mechanisms that could drive different rates of juvenile coral mortality across depths, including differences in predation, competition, and the availability of suitable substratum. The results of these experiments suggested that increased coral mortality at 17 m may have been influenced by higher intensity of fish predation, and higher mortality of corals attached to unfavorable substratum. In contrast, the abundance of macroalgae, a coral competitor, did not explain differences in coral survival. Our work suggests that top‐down processes and substratum quality can create bottlenecks in corals that can drive rates of coral recovery after disturbance. 
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  6. Abstract Marine heatwaves are increasing in frequency and duration, threatening tropical reef ecosystems through intensified coral bleaching events. We examined a strikingly variable spatial pattern of bleaching in Moorea, French Polynesia following a heatwave that lasted from November 2018 to July 2019. In July 2019, four months after the onset of bleaching, we surveyed > 5000 individual colonies of the two dominant coral genera,PocilloporaandAcropora, at 10 m and 17 m water depths, at six forereef sites around the island where temperature was measured. We found severe bleaching increased with colony size for both coral genera, butAcroporableached more severely thanPocilloporaoverall. Acroporableached more at 10 m than 17 m, likely due to higher light availability at 10 m compared to 17 m, or greater daily temperature fluctuation at depth. Bleaching inPocilloporacorals did not differ with depth but instead varied with the interaction of colony size and Accumulated Heat Stress (AHS), in that larger colonies (> 30 cm) were more sensitive to AHS than mid-size (10–29 cm) or small colonies (5–9 cm). Our findings provide insight into complex interactions among coral taxa, colony size, and water depth that produce high spatial variation in bleaching and related coral mortality. 
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  7. Abstract Climate change is intensifying extreme weather events, including marine heatwaves, which are prolonged periods of anomalously high sea surface temperature that pose a novel threat to aquatic animals. Tropical animals may be especially vulnerable to marine heatwaves because they are adapted to a narrow temperature range. If these animals cannot acclimate to marine heatwaves, the extreme heat could impair their behavior and fitness. Here, we investigated how marine heatwave conditions affected the performance and thermal tolerance of a tropical predatory fish, arceye hawkfish (Paracirrhites arcatus), across two seasons in Moorea, French Polynesia. We found that the fish’s daily activities, including recovery from burst swimming and digestion, were more energetically costly in fish exposed to marine heatwave conditions across both seasons, while their aerobic capacity remained the same. Given their constrained energy budget, these rising costs associated with warming may impact how hawkfish prioritize activities. Additionally, hawkfish that were exposed to hotter temperatures exhibited cardiac plasticity by increasing their maximum heart rate but were still operating within a few degrees of their thermal limits. With more frequent and intense heatwaves, hawkfish, and other tropical fishes must rapidly acclimate, or they may suffer physiological consequences that alter their role in the ecosystem. 
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  8. Abstract. African elephants (Loxodonta africana) are the largest extant terrestrial mammals, with bodies containing enormous quantities of nutrients. Yet, we know little about how these nutrients move through the ecosystem after an elephant dies. Here, we investigated the initial effects (1–26 months postmortem) of elephant megacarcasses on savanna soil and plant nutrient pools in the Kruger National Park, South Africa. We hypothesized that (H1) elephant megacarcass decomposition would release nutrients into soil, resulting in higher concentrations of soil nitrogen (N), phosphorus (P), and micronutrients near the center of carcass sites; (H2) carbon (C) inputs into the soil would stimulate microbial activity, resulting in increased soil respiration potential near the center of carcass sites; and (H3) carcass-derived nutrients would be absorbed by plants, resulting in higher foliar nutrient concentrations near the center of carcass sites. To test our hypotheses, we identified 10 elephant carcass sites split evenly between nutrient-poor granitic and nutrient-rich basaltic soils. At each site, we ran transects in the four cardinal directions from the center of the carcass site, collecting soil and grass (Urochloa trichopus, formerly U. mosambicensis) samples at 0, 2.5, 5, 10, and 15 m. We then analyzed samples for C, N, P, and micronutrient concentrations and quantified soil microbial respiration potential. We found that concentrations of soil nitrate, ammonium, δ15N, phosphate, and sodium were elevated closer to the center of carcass sites (H1). Microbial respiration potentials were positively correlated with soil organic C, and both respiration and organic C decreased with distance from the carcass (H2). Finally, we found evidence that plants were readily absorbing carcass-derived nutrients from the soil, with foliar %N, δ15N, iron, potassium, magnesium, and sodium significantly elevated closer to the center of carcass sites (H3). Together, these results indicate that elephant megacarcasses release ecologically consequential pulses of nutrients into the soil which stimulate soil microbial activity and are absorbed by plants into the above-ground nutrient pools. These localized nutrient pulses may drive spatiotemporal heterogeneity in plant diversity, herbivore behavior, and ecosystem processes. 
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  9. Ecosystems are intrinsically linked, such that management actions in one ecosystem can influence adjacent ecosystems. However, adequate data, and even protocols, for monitoring cross-ecosystem responses to conservation initiatives are lacking. Here, we evaluate potential indicators, operating on different spatial, temporal, and biological scales, for measuring the effects of island-based restoration on coral reef ecosystems. We show that island restoration status had consistent effects on populations of tropical seabirds across spatial scales from 100 m to entire islands. Seabirds, in turn, provided nutrient subsidies that were incorporated by marine algae and coral-reef fishes, with the most pronounced effects closer to shore, at leeward sites, and at low trophic levels. Microbes and macroalgae exhibited assemblage-level responses to seabird-derived nutrients entering the marine environment, but there were few differences in coral reef benthic and fish assemblages. By identifying and focusing on specific indicators such as macroalgal nutrients, managers can better monitor cross-ecosystem responses to conservation interventions with limited resources. 
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