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Intraspecific niche divergence is an important driver of species range, population abundance and impacts on ecosystem functions. Genetic changes are the primary focus when studying intraspecific divergence; however, the role of ecological interactions, particularly host‐microbiome symbioses, is receiving increased attention. The relative importance of these evolutionary and ecological mechanisms has seen only limited evaluation. To address this question, we usedMicrocystis aeruginosa, the globally distributed cyanobacterium that dominates freshwater harmful algal blooms. These blooms have been increasing in occurrence and intensity worldwide, causing major economic and ecological damages. We evaluated 46 isolates ofM. aeruginosaand their microbiomes,collected from 14 lakes in Michigan, USA, that vary over 20‐fold in phosphorus levels, the primary limiting nutrient in freshwater systems. Genomes ofM. aeruginosadiverged along this phosphorus gradient in genomic architecture and protein functions. Fitness in low‐phosphorus lakes corresponded with additional shifts withinM. aeruginosaincluding genome‐wide reductions in nitrogen use, an expansion of phosphorus assimilation genes and an alternative life history strategy of nonclonal colony formation. In addition to host shifts, despite culturing in common‐garden conditions, host‐microbiomes diverged along the gradient in taxonomy, but converged in function with evidence of metabolic interdependence between the host and its microbiome. Divergence corresponded with a physiological trade‐off between fitness in low‐phosphorus environments and growth rate in phosphorus‐rich conditions. Co‐occurrence of genotypes adapted to different nutrient environments in phosphorus‐rich lakes may have critical implications for understanding howM. aeruginosablooms persist after initial nutrient depletion. Ultimately, we demonstrate that the intertwined effects of genome evolution, host life history strategy and ecological interactions between a host and its microbiome correspond with an intraspecific niche shift with important implications for whole ecosystem function.

Ecosystems across the United States are changing in complex and unpredictable ways and analysis of these changes requires coordinated, long‐term research. This paper is a product of a synthesis effort of the U.S. National Science Foundation funded Long‐Term Ecological Research (LTER) network addressing the LTER core research area of “populations and communities.” This analysis revealed that each LTER site had at least one compelling “story” about what their site would look like in 50–100 yr. As the stories were prepared, themes emerged, and the stories were group into papers along five themes: state change, connectivity, resilience, time lags, and cascading effects. This paper addresses the cascading effects theme and includes stories from the Bonanza Creek (boreal), Kellogg Biological Station (agricultural and freshwater), Palmer (Antarctica), and Harvard Forest (temperate forest) LTER sites. We define cascading effects very broadly to include a wide array of unforeseen chains of events that result from a variety of actions or changes in a system. While climate change is having important direct effects on boreal forests, indirect effects mediated by fire activity—severity, size, and return interval—have large cascading effects over the long term. In northeastern temperate forests, legacies of human management and disturbance affect the composition of current forests, which creates a cascade of effects that interact with the climate‐facilitated invasion of an exotic pest. In Antarctica, declining sea ice creates a cascade of effects including declines in Adèlie and increases in Gentoo penguins, changes in phytoplankton, and consequent changes in zooplankton populations. An invasion of an exotic species of lady beetle is likely to have important future effects on pest control and conservation of native species in agricultural landscapes. New studies of zebra mussels, a well‐studied invader, have established links between climate, the heat tolerance of the mussels, and harmful algal blooms. Collectively, these stories highlight the need for long‐term studies to sort out the complexities of different types of ecological cascades. The diversity of sites within the LTER network facilitates the emergence of overarching concepts about trophic interactions as an important driver of ecosystem structure, function, services, and futures.