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Dramatic changes in populations of fishes living on coral reefs have been documented globally and, in response, the research community has initiated efforts to assess and monitor reef fish assemblages. A variety of visual census techniques are employed, however results are often incomparable due to differential methodological performance. Although comparability of data may promote improved assessment of fish populations, and thus management of often critically important nearshore fisheries, to date no standardized and agreed-upon survey method has emerged. This study describes the use of methods across the research community and identifies potential drivers of method selection. An online survey was distributed to researchers from academic, governmental, and non-governmental organizations internationally. Although many methods were identified, 89% of survey-based projects employed one of three methods-belt transect, stationary point count, and some variation of the timed swim method. The selection of survey method was independent of the research design (i.e., assessment goal) and region of study, but was related to the researcher's home institution. While some researchers expressed willingness to modify their current survey protocols to more standardized protocols (76%), their willingness decreased when methodologies were tied to long-term datasets spanning five or more years. Willingness to modify current methodologies was also less common among academic researchers than resource managers. By understanding both the current application of methods and the reported motivations for method selection, we hope to focus discussions towards increasing the comparability of quantitative reef fish survey data.

Upwelling is an important source of inorganic nutrients in marine systems, yet little is known about how gradients in upwelling affect primary producers on coral reefs. The Southern Line Islands span a natural gradient of inorganic nutrient concentrations across the equatorial upwelling region in the central Pacific. We used this gradient to test the hypothesis that benthic autotroph ecophysiology is enhanced on nutrient-enriched reefs. We measured metabolism and photophysiology of common benthic taxa, including the algae Porolithon, Avrainvillea, and Halimeda, and the corals Pocillopora and Montipora. We found that temperature (27.2-28.7°C) was inversely related to dissolved inorganic nitrogen (0.46-4.63 μM) and surface chlorophyll a concentrations (0.108-0.147 mg m-3), which increased near the equator. Contrary to our prediction, ecophysiology did not consistently track these patterns in all taxa. Though metabolic rates were generally variable, Porolithon and Avrainvillea photosynthesis was highest at the most productive and equatorial island (northernmost). Porolithon photosynthetic rates also generally increased with proximity to the equator. Photophysiology (maximum quantum yield) increased near the equator and was highest at northern islands in all taxa. Photosynthetic pigments also were variable, but chlorophyll a and carotenoids in Avrainvillea and Montipora were highest at the northern islands. Phycobilin pigments of Porolithon responded most consistently across the upwelling gradient, with higher phycoerythrin concentrations closer to the equator. Our findings demonstrate that the effects of in situ nutrient enrichment on benthic autotrophs may be more complex than laboratory experiments indicate. While upwelling is an important feature in some reef ecosystems, ancillary factors may regulate the associated consequences of nutrient enrichment on benthic reef organisms.

Human impacts can homogenize and simplify ecosystems, favoring communities that are no longer naturally coupled with (or reflective of) the background environmental regimes in which they are found. Such a process of biophysical decoupling has been explored little in the marine environment due to a lack of replication across the intact-to-degraded ecosystem spectrum. Coral reefs lacking local human impacts provide critical baseline scenarios in which to explore natural biophysical relationships, and provide a template against which to test for their human-induced decoupling. Using 39 Pacific islands, 24 unpopulated (relatively free from local human impacts) and 15 populated (with local human impacts present), spanning 45° of latitude and 65° of longitude, we ask, what are ‘natural’ biophysical relationships on coral reefs and do we see evidence for their human-induced decoupling? Estimates of the percent cover of benthic groups were related to multiple physical environmental drivers (sea surface temperature, irradiance, chlorophyll-a, and wave energy) using mixed-effects models and island mean condition as the unit of replication. Models across unpopulated islands had high explanatory power, identifying key physical environmental drivers of variations in benthic cover in the absence of local human impacts. These same models performed poorly and lost explanatory power when fitted anew to populated (human impacted) islands; biophysical decoupling was clearly evident. Furthermore, key biophysical relationships at populated islands (i.e. those relationships driving benthic variation across space in conjunction with chronic human impact) bore little resemblance to the baseline scenarios identified from unpopulated islands. Our results highlight the ability of local human impacts to decouple biophysical relationships in the marine environment and fundamentally restructure the natural rules of nature.

An increase in the intensity and frequency of extreme environmental conditions due to anthropogenic climate change impacts coral reefs through myriad stressors, from elevated sea-surface temperatures to increased storm activity. A reef’s response to these disturbances can be influenced by factors including taxonomic composition, life history strategies, or spatial patterning of the reef community members. We explored the disturbance-specific responses of coral reefs by following changes in benthic cover of major functional groups, community assemblage, and the response of common coral taxa at six islands across the central Pacific over the course of two years. We observed a decrease in average coral cover at four of the six islands, with differing underlying shifts in assemblage structure. Reefs with the highest pre-disturbance benthic coverage of Montipora spp. displayed significant increases in average hard coral cover compared to those where Acropora spp. was in the highest abundance; Acropora -dominated reefs showed significant declines in coral cover especially when exposed to the physical stress associated with a cyclone. Change in total coral cover was variable between islands within the same region, even among adjacent islands facing similar disturbance. These results highlight the importance of assemblage composition in influencing how benthic communities respond to major disturbance events such as thermal stress or cyclones. Improving our understanding of the drivers of differential community responses to disturbance will be important in predicting future changes in reef structure under changing ocean conditions.

For sessile organisms such as reef-building corals, differences in the degree of dispersion of individuals across a landscape may result from important differences in life-history strategies or may reflect patterns of habitat availability. Descriptions of spatial patterns can thus be useful not only for the identification of key biological and physical mechanisms structuring an ecosystem, but also by providing the data necessary to generate and test ecological theory. Here, we used an in situ imaging technique to create large-area photomosaics of 16 plots at Palmyra Atoll, central Pacific, each covering 100 m 2 of benthic habitat. We mapped the location of 44,008 coral colonies and identified each to the lowest taxonomic level possible. Using metrics of spatial dispersion, we tested for departures from spatial randomness. We also used targeted model fitting to explore candidate processes leading to differences in spatial patterns among taxa. Most taxa were clustered and the degree of clustering varied by taxon. A small number of taxa did not significantly depart from randomness and none revealed evidence of spatial uniformity. Importantly, taxa that readily fragment or tolerate stress through partial mortality were more clustered. With little exception, clustering patterns were consistent with models of fragmentation and dispersal limitation. In some taxa, dispersion was linearly related to abundance, suggesting density dependence of spatial patterning. The spatial patterns of stony corals are non-random and reflect fundamental life-history characteristics of the taxa, suggesting that the reef landscape may, in many cases, have important elements of spatial predictability.

The majority of the world's coral reefs are in various stages of decline. While a suite of disturbances (overfishing, eutrophication, and global climate change) have been identified, the mechanism(s) of reef system decline remain elusive. Increased microbial and viral loading with higher percentages of opportunistic and specific microbial pathogens have been identified as potentially unifying features of coral reefs in decline. Due to their relative size and high per cell activity, a small change in microbial biomass may signal a large reallocation of available energy in an ecosystem; that is the microbialization of the coral reef. Our hypothesis was that human activities alter the energy budget of the reef system, specifically by altering the allocation of metabolic energy between microbes and macrobes. To determine if this is occurring on a regional scale, we calculated the basal metabolic rates for the fish and microbial communities at 99 sites on twenty-nine coral islands throughout the Pacific Ocean using previously established scaling relationships. From these metabolic rate predictions, we derived a new metric for assessing and comparing reef health called the microbialization score. The microbialization score represents the percentage of the combined fish and microbial predicted metabolic rate that is microbial. Our results demonstrate a strong positive correlation between reef microbialization scores and human impact. In contrast, microbialization scores did not significantly correlate with ocean net primary production, local chla concentrations, or the combined metabolic rate of the fish and microbial communities. These findings support the hypothesis that human activities are shifting energy to the microbes, at the expense of the macrobes. Regardless of oceanographic context, the microbialization score is a powerful metric for assessing the level of human impact a reef system is experiencing.

Sharks and other large predators are scarce on most coral reefs, but studies of their historical ecology provide qualitative evidence that predators were once numerous in these ecosystems. Quantifying density of sharks in the absence of humans (baseline) is, however, hindered by a paucity of pertinent time-series data. Recently researchers have used underwater visual surveys, primarily of limited spatial extent or nonstandard design, to infer negative associations between reef shark abundance and human populations. We analyzed data from 1607 towed-diver surveys (>1 ha transects surveyed by observers towed behind a boat) conducted at 46 reefs in the central-western Pacific Ocean, reefs that included some of the world's most pristine coral reefs. Estimates of shark density from towed-diver surveys were substantially lower (<10%) than published estimates from surveys along small transects (<0.02 ha), which is not consistent with inverted biomass pyramids (predator biomass greater than prey biomass) reported by other researchers for pristine reefs. We examined the relation between the density of reef sharks observed in towed-diver surveys and human population in models that accounted for the influence of oceanic primary productivity, sea surface temperature, reef area, and reef physical complexity. We used these models to estimate the density of sharks in the absence of humans. Densities of gray reef sharks (Carcharhinus amblyrhynchos), whitetip reef sharks (Triaenodon obesus), and the group \"all reef sharks\" increased substantially as human population decreased and as primary productivity and minimum sea surface temperature (or reef area, which was highly correlated with temperature) increased. Simulated baseline densities of reef sharks under the absence of humans were 1.1-2.4/ha for the main Hawaiian Islands, 1.2-2.4/ha for inhabited islands of American Samoa, and 0.9-2.1/ha for inhabited islands in the Mariana Archipelago, which suggests that density of reef sharks has declined to 3-10% of baseline levels in these areas. Los tiburones y otros depredadores mayores son escasos en la mayoría de los arrecifes de coral, pero estudios de su ecología histórica proporcionan evidencia cualitativa de que los depredadores una vez fueron numerosos en estos ecosistemas. Sin embargo, la cuantificación de la densidad de tiburones en ausencia de humanos (línea de base) es obstaculizada por la falta de datos de series de tiempo pertinentes. Recientemente, los investigadores han utilizado muestreos visuales submarinos, de extensión espacial limitada o de diseño no estándar, para inferir asociaciones negativas entre la abundancia de tiburones de arrecife y las poblaciones humanas. Analizamos datos de 1607 muestreos por remolque de buzos (transectos > muestreados por observadores remolcados por una lancha) realizados en 46 arrecifes en el Océano Pacífico centro-occidental, arrecifes que incluyeron algunos de los más prístinos del mundo. Las estimaciones de densidad de tiburones fue sustancialmente menor (<10%) que estimaciones publicadas a partir de muestreos a lo largo de transectos pequeños (<0.02 ha), lo cual no es consistente con las pirámides de biomasa invertidas (la biomasa de depredadores es mayor que la biomasa de presas) reportadas para arrecifes prístinos por otros autores. Examinamos la relación entre la densidad de tiburones de arrecife observados en los muestreos por remolque de buzos y la población humana en modelos y consideramos la influencia de la productividad oceánica primaria, la temperatura de la superficie del mar, la superficie del arrecife y su complejidad física. Utilizamos estos modelos para estimar la densidad de tiburones en ausencia de humanos. Las densidades de Carcharhinus amblyrhynchos, Triaenodon obesus y el grupo de \"tiburones estrictamente arrecifales\" incrementó sustancialmente a medida que disminuyó la población humana y que incrementó la productividad primaria y la temperatura de la superficie del mar (o superficie del arrecife, que estaba altamente correlacionada con la temperatura. Las densidades basales simuladas de tiburones arrecifales en ausencia de humanos fueron 1.1-2.4/ha para las Islas Hawaianas, 1.2-2.4/ha en islas deshabitadas de Samoa Americana y 0.9-2.1/ha e islas deshabitadas del Archipiélago Mariana, lo que sugiere que la densidad de tiburones arrecifales ha declinado entre 3 -10% en relación con los niveles basales en esas áreas.

Depth is used often as a proxy for gradients in energetic resources on coral reefs and for predicting patterns of community energy use. With increasing depth, loss of light can lead to a reduced reliance on autotrophy and an increased reliance on heterotrophy by mixotrophic corals. However, the generality of such trophic zonation varies across contexts. By combining high-resolution oceanographic measurements with isotopic analyses (δ13C, δ15N) of multiple producer and consumer levels across depths (10–30 m) at a central Pacific oceanic atoll, we show trophic zonation in mixotrophic corals can be both present and absent within the same reef system. Deep-water internal waves that deliver cool particulate-rich waters to shallow reefs occurred across all sites (2.5–5.6 events week−1 at 30 m) but the majority of events remained depth-restricted (4.3–9.7% recorded at 30 m propagated to 10 m). In the absence of other particulate delivery, mixotrophs increased their relative degree of heterotrophy with increasing depth. However, where relatively long-lasting downwelling events (1.4–3.3 times the duration of any other site) occurred simultaneously, mixotrophs displayed elevated and consistent degrees of heterotrophy regardless of depth. Importantly, these long-lasting surface pulses were of a lagoonal origin, an area of rich heterotrophic resource supply. Under such circumstances, we hypothesize heterotrophic resource abundance loses its direct linkage with depth and, with resources readily available at all depths, trophic zonation is no longer present. Our results show that fine-scale intra-island hydrographic regimes and hydrodynamic connectivity between reef habitats contribute to explaining the context specific nature of coral trophic depth zonation in shallow reef ecosystems.

Islands support unique plants, animals, and human societies found nowhere else on the Earth. Local and global stressors threaten the persistence of island ecosystems, with invasive species being among the most damaging, yet solvable, stressors. While the threat of invasive terrestrial mammals on island flora and fauna is well recognized, recent studies have begun to illustrate their extended and destructive impacts on adjacent marine environments. Eradication of invasive mammals and restoration of native biota are promising tools to address both island and ocean management goals. The magnitude of the marine benefits of island restoration, however, is unlikely to be consistent across the globe. We propose a list of six environmental characteristics most likely to affect the strength of land– sea linkages: precipitation, elevation, vegetation cover, soil hydrology, oceanographic productivity, and wave energy. Global databases allow for the calculation of comparable metrics describing each environmental character across islands. Such metrics can be used today to evaluate relative potential for coupled land–sea conservation efforts and, with sustained investment in monitoring on land and sea, can be used in the future to refine science-based planning tools for integrated land–sea management. As conservation practitioners work to address the effects of climate change, ocean stressors, and biodiversity crises, it is essential that we maximize returns from our management investments. Linking efforts on land, including eradication of island invasive mammals, with marine restoration and protection should offer multiplied benefits to achieve concurrent global conservation goals.

The Hawaiian Archipelago experienced a moderate bleaching event in 2019—the third major bleaching event over a 6-year period to impact the islands. In response, the Hawai‘i Coral Bleaching Collaborative (HCBC) conducted 2,177 coral bleaching surveys across the Hawaiian Archipelago. The HCBC was established to coordinate bleaching monitoring efforts across the state between academic institutions, non-governmental organizations, and governmental agencies to facilitate data sharing and provide management recommendations. In 2019, the goals of this unique partnership were to: 1) assess the spatial and temporal patterns of thermal stress; 2) examine taxa-level patterns in bleaching susceptibility; 3) quantify spatial variation in bleaching extent; 4) compare 2019 patterns to those of prior bleaching events; 5) identify predictors of bleaching in 2019; and 6) explore site-specific management strategies to mitigate future bleaching events. Both acute thermal stress and bleaching in 2019 were less severe overall compared to the last major marine heatwave events in 2014 and 2015. Bleaching observed was highly site- and taxon-specific, driven by the susceptibility of remaining coral assemblages whose structure was likely shaped by previous bleaching and subsequent mortality. A suite of environmental and anthropogenic predictors was significantly correlated with observed bleaching in 2019. Acute environmental stressors, such as temperature and surface light, were equally important as previous conditions (e.g. historical thermal stress and historical bleaching) in accounting for variation in bleaching during the 2019 event. We found little evidence for acclimation by reefs to thermal stress in the main Hawaiian Islands. Moreover, our findings illustrate how detrimental effects of local anthropogenic stressors, such as tourism and urban run-off, may be exacerbated under high thermal stress. In light of the forecasted increase in severity and frequency of bleaching events, future mitigation of both local and global stressors is a high priority for the future of corals in Hawai‘i.