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ABSTRACT As environments worldwide change at unprecedented rates during the Anthropocene, understanding context dependency—how species interactions vary depending on environmental context—is crucial. Combining comparative genomics across 42 angiosperms with transcriptomics, genome‐wide association mapping and gene duplication origin analyses, we show for the first time that gene family expansions are important to context‐dependent regulation of species interactions. Gene families expanded in mycorrhizal fungi‐associating plants display up to 200% more context‐dependent gene expression and double the genetic variation associated with mycorrhizal benefits to plant fitness. Moreover, we discover these gene family expansions arise primarily from tandem duplications with > 2‐times more tandem duplications genome‐wide, indicating gene family expansions continuously supply genetic variation, allowing fine‐tuning of context dependency in species interactions throughout plant evolution. Taken together, our results spotlight how widespread gene duplications can provide molecular flexibility required for plant–microbial interactions to match changing environmental conditions. 

Summary Plant microbiomes have the potential to mitigate the impacts of climate change, yet both the complexity of climate change and the complexity of plant–microbe interactions make applications and future predictions challenging. Here, we embrace this complexity, reviewing how different aspects of climate change influence beneficial plant–microbe interactions and how advances in theory, tools, and applications may improve understanding and predictability of climate change effects on plants, microbiomes, and their roles within ecosystems. New advances include consideration of (1) interactions among climate stressors, such as more variable precipitation regimes combined with warmer mean temperature; (2) mechanisms that promote the stability of microbiome functions; (3) legacies of stress affecting the functionality of microbial communities under future stress; and (4) temporally repeated plant–microbe interactions or feedbacks. We also identify key gaps in each of these areas and spotlight the need for more research bridging molecular biology and ecology to develop a more mechanistic understanding of how climate change shapes beneficial microbe–plant interactions.