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  1. The resilience of coral reefs in oligotrophic, (sub)tropical oceans is largely due to the symbiotic relationship between scleractinian corals and Symbiodiniaceae algae, which enables efficient internal nutrient recycling. Investigating the history of this coral symbiosis can provide insights into its role in sustaining the health of both present and future coral reefs. The isotopic composition of organic nitrogen (15N/14N or δ15N) bound within coral skeletons has been utilized to trace the existence of symbiosis in fossil corals, suggesting that coral symbiosis dates back to at least 210 million years ago. The basis of this proxy is that symbiotic corals are expected to exhibit lower δ15N compared to their non-symbiotic (aposymbiotic) counterparts within the same environments, owing to internal nitrogen recycling between the coral host and algal symbiont, and reduced leakage of low-δ15N ammonium into seawater. However, this hypothesis has not been adequately tested in contemporary settings. In a laboratory experiment, we examined the δ15N differences between the symbiotic and aposymbiotic branches within the same genetic backgrounds of the facultatively symbiotic coralOculina arbusculaunder well-fed conditions. Across five different genotypes in two separate experiments, symbiotic branches consistently showed lower δ15N than their aposymbiotic counterparts. These findings corroborate the use of δ15N as a proxy for identifying coral symbiosis in the past, particularly when multiple species of corals coexisted in the same environments. 
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    Free, publicly-accessible full text available August 29, 2025
  2. Fly ash—the residuum of coal burning—contains a considerable amount of fossilized particulate organic carbon (FOC ash ) that remains after high-temperature combustion. Fly ash leaks into natural environments and participates in the contemporary carbon cycle, but its reactivity and flux remained poorly understood. We characterized FOC ash in the Chang Jiang (Yangtze River) basin, China, and quantified the riverine FOC ash fluxes. Using Raman spectral analysis, ramped pyrolysis oxidation, and chemical oxidation, we found that FOC ash is highly recalcitrant and unreactive, whereas shale-derived FOC (FOC rock ) was much more labile and easily oxidized. By combining mass balance calculations and other estimates of fly ash input to rivers, we estimated that the flux of FOC ash carried by the Chang Jiang was 0.21 to 0.42 Mt C⋅y −1 in 2007 to 2008—an amount equivalent to 37 to 72% of the total riverine FOC export. We attributed such high flux to the combination of increasing coal combustion that enhances FOC ash production and the massive construction of dams in the basin that reduces the flux of FOC rock eroded from upstream mountainous areas. Using global ash data, a first-order estimate suggests that FOC ash makes up to 16% of the present-day global riverine FOC flux to the oceans. This reflects a substantial impact of anthropogenic activities on the fluxes and burial of fossil organic carbon that has been made less reactive than the rocks from which it was derived. 
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