Landscape heterogeneity increases the stability of wild ungulate populations facing climatic variability in Mediterranean ecosystems

Mediterranean environments are characterized by strong intra- and inter-annual fluctuations in plant primary production, which are likely to regulate the carrying capacity and density-dependent responses of ungulate populations. These effects may, however, vary across spatial and temporal scales. Habitat heterogeneity, particularly when associated to differentiated phenological responses, may allow wild ungulates to mitigate temporal fluctuations in plant production by using different resources along the year. In this work, we use a 15-years dataset (including remote-sensing data on vegetation distribution, phenology and production, as well as ungulate population counts) to assess how temporal variability in plant primary production and livestock abundance influence the population dynamics of two wild ungulates: native red deer, Cervus elaphus, and introduced fallow deer, Dama dama. Results show that temporal alternation in the phenological cycles of the four different vegetation types increased plant production, thus food availability for ungulates, within each year. Furthermore, complementarity in the responses of different vegetation types to variations in the amount and timing of rainfall increased the predictability of food availability across different years. This complementarity effect was further increased by the contrasting responses of ungulate populations to variation in the production of different vegetation types. Furthermore, domestic ungulates had positive effects on wild ungulate density at low to intermediate abundances, but high livestock densities decreased ungulate density and constrained the stability of the plant-ungulate system in response to the impact of climatic variation, particularly under climate change. Our findings deepen the knowledge on vegetation-ungulate interactions in Mediterranean areas, potentially contributing to develop better management strategies of ungulate populations and adapt them to ongoing climate change.