The adaptive value of habitat preferences from a multi-scale spatial perspective : Insights from marsh-nesting avian species

Background. Habitat selection and its adaptive outcomes are crucial features for animal life-history strategies. Nevertheless, congruence between habitat preferences and breeding success has been rarely demonstrated, which may result from the singlescale evaluation of animal choices. As habitat selection is a complex multi-scale process in many groups of animal species, investigating adaptiveness of habitat selection in a multi-scale framework is crucial. In this study, we explore whether habitat preferences acting at different spatial scales enhance the fitness of bird species, and check the appropriateness of single vs. multi-scale models. We expected that variables found to be more important for habitat selection at individual scale(s), would coherently play a major role in affecting nest survival at the same scale(s). Methods. We considered habitat preferences of two Rallidae species, little crake (Zapornia parva) and water rail (Rallus aquaticus), at three spatial scales (landscape, territory, and nest-site) and related them to nest survival. Single-scale versus multi-scale models (GLS and glmmPQL) were compared to check which model better described adaptiveness of habitat preferences. Consistency between the effect of variables on habitat selection and on nest survival was checked to investigate their adaptive value. Results. In both species, multi-scale models for nest survival were more supported than single-scale ones. In little crake, the multi-scale model indicated vegetation density and water depth at the territory scale, as well as vegetation height at nest-site scale, as the most important variables. The first two variables were among the most important for nest survival and habitat selection, and the coherent effects suggested the adaptive value of habitat preferences. In water rail, the multi-scale model of nest survival showed vegetation density at territory scale and extent of emergent vegetation within landscape scale as the most important ones, although we found a consistent effect with the habitat selection model (and hence evidence for adaptiveness) only for the former. Discussion. Our work suggests caution when interpreting adaptiveness of habitat preferences at a single spatial scale because such an approach may under- or overestimate the importance of habitat factors. As an example, we found evidence only for a weak effect of water depth at territory scale on little crake nest survival; however, according to the multi-scale analysis, such effect turned out to be important and appeared highly adaptive. Therefore, multi-scale approaches to the study of adaptive explanations for habitat selection mechanisms should be promoted.