Species' range edges are expanding to both warmer and cooler regions. Yet, no studies directly compared the changes in range-limiting traits within the same species during both types of range expansions. To increase our mechanistic understanding of range expansions, it is crucial to disentangle the contributions of plastic and genetic changes in these traits. The aim of this study was to test for plastic and evolutionary changes in heat tolerance, life history, and behavior, and compare these during range expansions toward warmer and cooler regions. Using laboratory experiments we reconstructed the thermal performance curves (TPCurves) of larval life history (survival, growth, and development rates) and larval heat tolerance (CTmax) across two recent range expansions from the core populations in southern France toward a warmer (southeastern Spain) and a cooler (northwestern Spain) region in Europe by the damselfly Ischnura elegans. First-generation larvae from field-collected mothers were reared across a range of temperatures (16°-28°C) in incubators. The range expansion to the warmer region was associated with the evolution of a greater ability to cope with high temperatures (increased mean and thermal plasticity of CTmax), faster development, and, in part, a faster growth, indicating a higher time constraints caused by a shorter time frame available for larval development associated with a transition to a greater voltinism. Our results thereby support the emerging pattern that plasticity in heat tolerance alone is inadequate to adapt to new thermal regimes. The range expansion to the cooler region was associated with faster growth indicating countergradient variation without a change in CTmax. The evolution of a faster growth rate during both range expansions could be explained by a greater digestive efficiency rather than an increased food intake. Our results highlight that range expansions to warmer and cooler regions can result in similar evolutionary changes in the TPCurves for life history, and no opposite changes in heat tolerance.
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