Microclimatic changes caused by plant invasions and warming: uncovering thermal costs and benefits to a tortoise.

Non-native plant invasions and climate warming alter the microclimatic conditions that organisms experience in their habitats, with potential implications for the fitness of native faunal species, particularly ectotherms. Predictions for species conservation increasingly use microclimate data at fine spatial scales relevant to organisms, but they typically overlook the modulating effect that vegetation changes have on the microclimates available in the habitat. Here we quantify the microclimatic changes imposed by invasive trees and simultaneous warming on native habitats and assess the resulting thermal benefits and costs to a small tortoise species (Homopus areolatus) from an organismal perspective and throughout its life cycle. We logged operative temperature above- and belowground in the field, covering the diversity of microhabitats across the four seasons of the year, and assessed the species' optimal temperature in the laboratory. Moving beyond the common use of averages, we applied a range of metrics to quantify differences between invaded and native areas in spatio-temporal temperature distributions, combined effects with warming and thermal habitat suitability for the species. We found that invaded areas became cooler and less exposed to temperatures above the species' optimal in summer. This buffering effect is expected to become more pronounced with further climate warming, turning invaded areas into potential thermal refugia. However, reduced spatial thermal heterogeneity during warm periods, more prevalent sub-optimal low temperatures in winter and colder underground incubation conditions in invaded areas could be detrimental to the species' long-term performance. Our results reveal the mixed nature of thermal effects of invasive plants on ectotherms, underscoring the importance of applying a suite of metrics to assess microclimate distribution changes. The approach used here illustrates the value of integrating thermal physiological and microclimatic information for a more mechanistic understanding of conservation problems.