For over three decades, scientists have conducted heat-stress experiments to predict how coral will respond to ocean warming due to global climate change. However, there are often conflicting results in the literature that are difficult to resolve, which we hypothesize are a result of unintended biases, variation in experimental design, and underreporting of critical methodological information. Here, we reviewed 255 coral heat-stress experiments to (1) document where and when they were conducted and on which species, (2) assess variability in experimental design, and (3) quantify the diversity of response variables measured. First, we found that twothirds of studies were conducted in only three countries, three coral species were more heavily studied than others, and only 4% of studies focused on earlier life stages. Second, slightly more than half of all heat-stress exposures were less than 8 d in duration, only 17% of experiments fed corals, and experimental conditions varied widely, including the level and rate of temperature increase, light intensity, number of genets used, and the length of acclimation period. In addition, 95%, 55%, and [ 35% of studies did not report tank flow conditions, light-dark cycle used, or the date of the experiment, respectively. Finally, we found that 21% of experiments did not measure any bleaching phenotype traits, 77% did not identify the Symbiodiniaceae endosymbiont, and the contribution of the coral host in the physiological response to heat-stress was often not investigated. This review highlights geographic, taxonomic, and heat-stress duration biases in our understanding of coral bleaching, and large variability in the reporting and design of heat-stress experiments that could account for some of the discrepancies in the literature. Development of some best practice recommendations for coral bleaching experiments could improve cross-studies comparisons and increase the efficiency of coral bleaching research at a time when it is needed most. Keywords Coral bleaching Á Coral heat-stress Á Temperature experiment Á Heat-stress experiment Á Bleaching experiment Á Coral bleaching review

Coral bleaching is the single largest global threat to coral reefs worldwide. Integrating the diverse body of work on coral bleaching is critical to understanding and combating this global problem. Yet investigating the drivers, patterns, and processes of coral bleaching poses a major challenge. A recent review of published experiments revealed a wide range of experimental variables used across studies. Such a wide range of approaches enhances discovery, but without full transparency in the experimental and analytical methods used, can also make comparisons among studies challenging. To increase comparability but not stifle innovation, we propose a common framework for coral bleaching experiments that includes consideration of coral provenance, experimental conditions, and husbandry. For example, reporting the number of genets used, collection site conditions, the experimental temperature offset(s) from the maximum monthly mean (MMM) of the collection site, experimental light conditions, flow, and the feeding regime will greatly facilitate comparability across studies. Similarly, quantifying common response variables of endosymbiont (Symbiodiniaceae) and holobiont phenotypes (i.e., color, chlorophyll, endosymbiont cell density, mortality, and skeletal growth) could further facilitate cross-study comparisons. While no single bleaching experiment can provide the data necessary to determine global coral responses of all corals to current and future ocean warming, linking studies through a common framework as outlined here, would help increase comparability among experiments, facilitate synthetic insights into the causes and underlying mechanisms of coral bleaching, and reveal unique bleaching responses among genets, species, and regions. Such a collaborative framework that fosters transparency in methods used would strengthen comparisons among studies that can help inform coral reef management and facilitate conservation strategies to mitigate coral bleaching worldwide.

The global impacts of climate change are evident in every marine ecosystem. On coral reefs, mass coral bleaching and mortality have emerged as ubiquitous responses to

Corals obtain nutrition from the photosynthetic products of their algal endosymbionts and the ingestion of organic material and zooplankton from the water column. Here, we use stable carbon (δ13C) and nitrogen (δ15N) isotopes to assess the proportionate contribution of photoautotrophic and heterotrophic sources to seven Hawaiian coral species collected from six locations around the island of O‘ahu, Hawaiʻi. We analyzed the δ13C and δ15N of coral tissues and their algal endosymbionts, as well as that of dissolved inorganic matter, particulate organic matter, and zooplankton from each site. Estimates of heterotrophic contribution varied among coral species and sites. Bayesian mixing models revealed that heterotrophic sources (particulate organic material and zooplankton) contributed the most to Pocillopora acuta and Montipora patula coral tissues at 49.3% and 48.0%, respectively, and the least to Porites lobata at 28.7%, on average. Estimates of heterotrophic contribution based on the difference between δ13C of the host and algal endosymbiont (δ13Ch–e) and isotopic niche overlap often differed, while estimates based on δ15Nh–e were slightly more aligned with the estimates produced using Bayesian mixing models. These findings suggest that the utility of each approach may vary with coral health status, regions, and coral species. Overall, we find that the mean heterotrophic contribution to Hawaiian coral tissues ranges from 20% to 50%, suggesting a variety of trophic strategies. However, these findings did not always match past direct measurements of heterotrophic feeding, indicating that heterotrophically acquired nutrition does not necessarily get incorporated into tissues but can be respired or exuded in mucus.