Extreme climatic events including marine heatwaves (MHWs) are becoming more frequent and severe in the Anthropocene. However, our understanding of how these events affect population dynamics of ecologically important species is limited, in part because extreme events are rare and difficult to predict. Here, we quantified the occurrence and severity of MHWs over 60 years in warm range edge kelp forests on both sides of the North Atlantic. The cumulative annual intensity of MHWs increased two-to four-fold during this period, coinciding with the disappearance of kelps. We experimentally demonstrated a relationship between strong and severe 2018 heatwaves and high kelp mortality in both regions. Patterns of kelp mortality were strongly linked to maximum temperature anomalies, which crossed lethal thresholds in both regions. translocation and tagging experiments revealed similar kelp mortality rates on reefs dominated by healthy kelp forests and degraded sediment-laden algal 'turfs', indicating equal vulnerability to extreme events. These results suggest a mechanistic link between MHWs and broad-scale kelp loss, and highlight how warming can make ecosystem boundaries unstable, forcing shifts to undesirable ecosystem states under episodically extreme climatic conditions. Extreme events may pose a stronger, more immediate threat to ecosystem function than shifts in average conditions, and are increasingly manifesting as key drivers of ecosystem reconfiguration as environmental conditions become more variable and extremes more frequent with climate change 1-3. Discrete periods of anomalously high ocean temperatures, known as "marine heatwaves" (MHWs) 4 , can have serious consequences for ecosystems, and are often associated with loss of ecological function and services 5,6. The increased severity and frequency of MHWs threaten biodiversity and ecosystem function on global scales 5 and the total number of MHW days per year has increased by > 50% in recent decades 1,7,8. Yet, the ecological consequences of extreme events such as MHWs have been identified as a key knowledge gap in ecology 9,10 , and our understanding of population dynamics during MHWs is limited 5. This is partly because these processes are not often studied in the field because MHWs are difficult to predict, and their impacts challenging to manipulate in situ 11,12. Compounding these problems, until recently, most ecological research on MHWs has been conducted on local scales using varying definitions of extremity, making broadscale measures and comparisons of the ecological effects of MHWs challenging 4,13. Temperature extremes appear to be particularly damaging for species located at their range edge, because they can abruptly push conditions beyond thermal tolerances, causing direct mortality 5,14. Kelps are cool-water species that are broadly distributed and respond strongly to changing abiotic conditions 15. Temperature is the most important factor controlling the range distribution of kelps 16-18 , and kelp forests in many regions have been shown to ...
Green sea urchins Strongylocentrotus droebachiensis along the coast of Nova Scotia, Canada, suffer mass mortalities from infection by the pathogenic amoeba Paramoeba invadens Jones, 1985. It has been speculated that P. invadens could be a form of Neoparamoeba pema quidensis, a species associated with disease in S. droebachiensis and lobsters in the northeast USA. During a disease outbreak in fall 2011, we isolated amoebae from moribund urchins collected from 4 locations along ~200 km of coastline. In laboratory infection trials, we found that timing and rate of morbidity corresponded to that of similar experiments conducted in the early 1980s, when P. invadens was first identified. All isolates had a similar size and morphology to the original description, including an absence of microscales. Sequences of nuclear SSU rDNA show that disease was caused by one 'species' of amoeba across the range sampled. Phylogenetic analyses prove that P. invadens is not conspecific with N. pemaquidensis, but is a distinct species most closely related to N. branchiphila, a suspected pathogen of sea urchins Diadema aff. antillarum in the Canary Islands, Spain. Morphology and closest phylogenetic affinities suggest that P. invadens would be assignable to the genus Neoparamoeba; however, nuclear SSU rDNA trees show that Neoparamoeba and Paramoeba are phylogenetically inseparable. Therefore, we treat Neoparamoeba as a junior synonym of Paramoeba, with P. invadens retaining that name, and N. pemaquidensis and N. aestuarina reverting to their original names (P. pemaquidensis and P. aestuarina), and with new combinations for N. branchiphila Dykova et al., 2005, and N. perurans Young et al., 2007, namely P. branchiphila comb. nov. and P. perurans comb. nov
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