Latitude-associated evolution and drivers of thermal response curves in body stoichiometry

Trait-based studies are needed to understand the plastic and genetic responses of organisms to warming. A neglected organismal trait is elemental composition, despite its potential to cascade into effects on the ecosystem level. Warming is predicted to shape elemental composition through shifts in storage molecules associated with responses in growth, body size and metabolic rate. Our goals were to quantify thermal response patterns in body composition and to obtain insights into their underlying drivers and their evolution across latitudes. We reconstructed the thermal response curves (TRCs) for body elemental composition [C (carbon), N (nitrogen) and the C:N ratio] of damselfly larvae from high- and low-latitude populations. Additionally, we quantified the TRCs for survival, growth and development rates and body size to assess local thermal adaptation, as well as the TRCs for metabolic rate and key macromolecules (proteins, fat, sugars and cuticular melanin and chitin) as these may underlie the elemental TRCs. All larvae died at 36 degrees C. Up to 32 degrees C, low-latitude larvae increased growth and development rates and did not suffer increased mortality. Instead, growth and development rates of high-latitude larvae were lower and levelled off at 24 degrees C, and mortality increased at 32 degrees C. This latitude-associated thermal adaptation pattern matched the 'hotter-is-better' hypothesis. With increasing temperatures, low-latitude larvae decreased C:N, while high-latitude larvae increased C:N. These patterns were driven by associated changes in N contents, while C contents did not respond to temperature. Consistent with the temperature-size rule and the thermal melanism hypothesis, body size and melanin levels decreased with warming. While all traits and associated macromolecules (except for metabolic rate that showed thermal compensation) assumed to underlie thermal responses in elemental composition showed thermal plasticity, these were largely independent and none could explain the stoichiometric TRCs. Our results highlight that thermal responses in elemental composition cannot be explained by traditionally assumed drivers, asking for a broader perspective including the thermal dependence of elemental fluxes. Another key implication is that thermal evolution can reverse the plastic stoichiometric thermal responses and hence reverse how warming may shape food web dynamics through changes in body composition at different latitudes.