Climate change is transforming aquatic ecosystems. Coastal waters have experienced progressive warming, acidification, and deoxygenation that will intensify this century. At the same time, there is a scientific consensus that the public health, recreation, tourism, fishery, aquaculture, and ecosystem impacts from harmful algal blooms (HABs) have all increased over the past several decades. The extent to which climate change is intensifying these HABs is not fully clear, but there has been a wealth of research on this topic this century alone. Indeed, the United Nations' Intergovernmental Panel on Climate Change's (IPCC) Special Report on the Ocean and Cryosphere in a Changing Climate (SROCC) approved in September 2019 was the first IPCC report to directly link HABs to climate change. In the Summary for Policy Makers, the report made the following declarations with "high confidence": In addition, the report specifically outlines a series of linkages between heat waves and HABs. These statements about HABs and climate change and the high levels of confidence ascribed to them provides clear evidence that the field of HABs and climate change has matured and has, perhaps, reached a first plateau of certainty. While there are well-documented global trends in HABs being promoted by human activity, including climate change, individual events are driven by local, regional, and global drivers, making it critical to carefully evaluate the conditions and responses at appropriate scales. It is within this context that the first Special Issue on Climate Change and Harmful Algal Blooms is published in Harmful Algae.

Marine and freshwater ecosystems are warming, acidifying, and deoxygenating as a consequence of climate change. In parallel, the impacts of harmful algal blooms (HABs) on these ecosystems are intensifying. Many eutrophic habitats that host recurring HABs already experience thermal extremes, low dissolved oxygen, and low pH, making these locations potential sentinel sites for conditions that will become more common in larger-scale systems as climate change accelerates. While studies of the effects of HABs or individual climate change stressors on aquatic organisms have been relatively common, studies assessing their combined impacts have been rare. Those doing so have reported strong species- and strain-specific interactions between HAB species and climate change co-stressors yielding outcomes for aquatic organisms that could not have been predicted based on investigations of these factors individually. This review provides an ecological and physiological framework for considering HABs as a climate change co-stressor and considers the consequences of their combined occurrence for coastal ecosystems. This review also highlights critical gaps in our understanding of HABs as a climate change co-stressor that must be addressed in order to develop management plans that adequately protect fisheries, aquaculture, aquatic ecosystems, and human health. Ultimately, incorporating HAB species into experiments and monitoring programs where the effects of multiple climate change stressors are considered will provide a more ecologically relevant perspective of the structure and function of marine ecosystems in future, climate-altered systems.

Significance: This study used high-resolution (daily, quarter-degree resolution) sea-surface temperature records to model trends in growth rates and bloom-season duration for two of the most toxic and widespread harmful algal bloom species indigenous to the North Atlantic and North Pacific oceans. Alexandrium fundyense synthesizes saxitoxin and Dinophysis acuminata synthesizes okadaic acid, which cause the human health syndromes paralytic and diarrhetic shellfish poisoning, respectively. The model provided hindcasts of harmful algal bloom (HAB) events that were consistent with in situ observations from long-term monitoring programs during the same time period. This study provides evidence that increasing ocean temperatures have already facilitated the intensification of these, and likely other, HABs and thus contribute to an expanding human health threat.

There is increasing recognition that low dissolved oxygen (DO) and low pH conditions co-occur in many coastal and open ocean environments. Within temperate ecosystems, these conditions not only develop seasonally as temperatures rise and metabolic rates accelerate, but can also display strong diurnal variability, especially in shallow systems where photosynthetic rates ameliorate hypoxia and acidification by day. Despite the widespread, global co-occurrence of low pH and low DO and the likelihood that these conditions may negatively impact marine life, very few studies have actually assessed the extent to which the combination of both stressors elicits additive, synergistic or antagonistic effects in marine organisms. We review the evidence from published factorial experiments that used static and/or fluctuating pH and DO levels to examine different traits (e.g. survival, growth, metabolism), life stages and species across a broad taxonomic spectrum. Additive negative effects of combined low pH and low DO appear to be most common; however, synergistic negative effects have also been observed. Neither the occurrence nor the strength of these synergistic impacts is currently predictable, and therefore, the true threat of concurrent acidification and hypoxia to marine food webs and fisheries is still not fully understood. Addressing this knowledge gap will require an expansion of multi-stressor approaches in experimental and field studies, and the development of a predictive framework. In consideration of marine policy, we note that DO criteria in coastal waters have been developed without consideration of concurrent pH levels. Given the persistence of concurrent low pH–low DO conditions in estuaries and the increased mortality experienced by fish and bivalves under concurrent acidification and hypoxia compared with hypoxia alone, we conclude that such DO criteria may leave coastal fisheries more vulnerable to population reductions than previously anticipated.

Harmful algal blooms (HABs) cause significant economic and ecological damage worldwide. Despite considerable efforts, a comprehensive understanding of the factors that promote these blooms has been lacking, because the biochemical pathways that facilitate their dominance relative to other phytoplankton within specific environments have not been identified. Here, biogeochemical measurements showed that the harmful alga Aureococcus anophagefferens outcompeted co-occurring phytoplankton in estuaries with elevated levels of dissolved organic matter and turbidity and low levels of dissolved inorganic nitrogen. We subsequently sequenced the genome of A. anophagefferens and compared its gene complement with those of six competing phytoplankton species identified through metaproteomics. Using an ecogenomic approach, we specifically focused on gene sets that may facilitate dominance within the environmental conditions present during blooms. A. anophagefferens possesses a larger genome (56 Mbp) and has more genes involved in light harvesting, organic carbon and nitrogen use, and encoding selenium- and metal-requiring enzymes than competing phytoplankton. Genes for the synthesis of microbial deterrents likely permit the proliferation of this species, with reduced mortality losses during blooms. Collectively, these findings suggest that anthropogenic activities resulting in elevated levels of turbidity, organic matter, and metals have opened a niche within coastal ecosystems that ideally suits the unique genetic capacity of A. anophagefferens and thus, has facilitated the proliferation of this and potentially other HABs.