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Due to global change, many species are shifting their distribution and are thereby confronted with novel thermal conditions at the moving range edges. Especially during the initial phases of exposure to a new environment, it has been hypothesized that plasticity and associated epigenetic mechanisms enable species to cope with environmental change. We tested this idea by capitalizing on the well-documented southward range expansion of the damselfly Ischnura elegans from France into Spain where the species invaded warmer regions in the 1950s in eastern Spain (old edge region) and in the 2010s in central Spain (new edge region). Using a common garden experiment at rearing temperatures matching the ancestral and invaded thermal regimes, we tested for evolutionary changes in (thermal plasticity in) larval life history and heat tolerance in these expansion zones. Through the use of de- and hypermethylating agents, we tested whether epigenetic mechanisms play a role in enabling heat tolerance during expansion. We used the phenotype of the native sister species in Spain, I. graellsii , as proxy for the locally adapted phenotype. New edge populations converged toward the phenotype of the native species through plastic thermal responses in life history and heat tolerance while old edge populations (partly) constitutively evolved a faster life history and higher heat tolerance than the core populations, thereby matching the native species. Only the heat tolerance of new edge populations increased significantly when exposed to the hypermethylating agent. This suggests that the DNA methylation machinery is more amenable to perturbation at the new edge and shows it is able to play a role in achieving a higher heat tolerance. Our results show that both (evolved) plasticity as well as associated epigenetic mechanisms are initially important when facing new thermal regimes but that their importance diminishes with time. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Stratigraphic relations, detailed petrography, microthermometry of fluid inclusions, and fine-scale isotopic analysis of diagenetic phases indicate a complex thermal history in Tithonian fluvial sandstones and lacustrine limestones of the Tera Group (North Spain). Two different thermal events have been recognized and characterized, which are likely associated with hydrothermal events that affected the Cameros Basin during the mid-Cretaceous and the Eocene. Multiple stages of quartz cementation were identified using scanning electron microscope cathodoluminescence on sandstones and fracture fills. Primary fluid inclusions reveal homogenization temperatures (Th) from 195 to 350°C in the quartz cements of extensional fracture fillings. The high variability of Th data in each particular fluid inclusion assemblage is related to natural reequilibration of the fluid inclusions, probably due to Cretaceous hydrothermal metamorphism. Some secondary fluid inclusion assemblages show very consistent data (Th = 281-305°C) and are considered not to have reequilibrated. They are likely related to an Eocene hydrothermal event or to a retrograde stage of the Cretaceous hydrothermalism. This approach shows how multiple thermal events can be discriminated. A very steep thermal gradient of 97-214°C/km can be deduced from δO values of ferroan calcites (δO −14.2/−11.8‰ V-PDB) that postdate quartz cements in fracture fillings. Furthermore, illite crystallinity data (anchizone-epizone boundary) are out of equilibrium with high fluid inclusion Th. These observations are consistent with heat-flux related to short-lived events of hydrothermal alteration focused by permeability contrasts, rather than to regional heat-flux associated with dynamo-thermal metamorphism. These results illustrate how thermal data from fracture systems can yield thermal histories markedly different from host-rock values, a finding indicative of hydrothermal fluid flow. [ABSTRACT FROM AUTHOR]