Foraging decisions by rodents are key for the long-term maintenance of oak populations in which avian seed dispersers are absent or inefficient. Decisions are determined by the environmental setting in which acorn-rodent encounters occur. In particular, seed value, competition and predation risks have been found to modify rodent foraging decisions in forest and human-modified habitats. Nonetheless, there is little information about their joint effects on rodent behavior, and hence, local acorn dispersal (or predation). In this work, we manipulate and model the mouse-oak interaction in a Spanish dehesa, an anthropogenic savanna system in which nearby areas can show contrasting levels of ungulate densities and antipredatory cover. First, we conducted a large-scale cafeteria field experiment, where we modified ungulate presence and predation risk, and followed mouse foraging decisions under contrasting levels of moonlight and acorn availability. Then, we estimated the net effects of competition and risk by means of a transition probability model that simulated mouse foraging decisions. Our results show that mice are able to adapt their foraging decisions to the environmental context, affecting initial fates of handled acorns. Under high predation risks mice foraged opportunistically carrying away large and small seeds, whereas under safe conditions large acorns tended to be predated in situ. In addition, in the presence of ungulates lack of antipredatory cover around trees reduced mice activity outside tree canopies, and hence, large acorns had a higher probability of survival. Overall, our results point out that inter-specific interactions preventing efficient foraging by scatter-hoarders can reduce acorn predation. This suggests that the maintenance of the full set of seed consumers as well as top predators in dehesas may be key for promoting local dispersal.

Scatter-hoarding decisions by rodents are key for the long-term maintenance of scattered tree populations. Decisions are determined by seed value, competition and predation risk, so that they can be influenced by the integrity of the biological system composed by trees, rodents, ungulate competitors, and rodent predators. We manipulate and model the oak-mice interaction in a Spanish dehesa, an anthropogenic savanna system suffering chronic tree regeneration failure, and quantify the joint effect of intrinsic and extrinsic factors on acorn dispersal effectiveness. First, we conducted a large-scale cafeteria field experiment, where we modified ungulate presence and predation risk, and followed mouse scatter-hoarding decisions under contrasting levels of moonlight and acorn availability. Then, we estimated the net effects of competition and risk by means of transition probability models that simulated mouse scatter-hoarding decisions according to the environmental context. Our results show that suboptimal conditions for mice balance the interaction towards the mutualism as they force mice to forage less efficiently. Under stressful conditions (predation risks and presence of ungulates), lack of antipredatory cover around dehesa trees limited transportation of acorns, but also precluded mice activities outside tree canopies. As a result, post-dispersal predation rates were reduced and large acorns had a higher probability to survive. Our work shows that inter-specific interactions preventing efficient foraging by scatter-hoarders benefitted seed dispersal. Therefore, the maintenance of the full set of producers, consumers, dispersers and predators in ecosystems is key for promoting seed dispersal effectiveness in conditional mutualisms.

Drought-induced forest dieback can lead to a tipping point in community dominance, but the coupled response at the tree and stand-level response has not been properly addressed. New spatially and temporally integrated monitoring approaches that target different biological organization levels are needed. Here, we compared the temporal responses of dendrochronological and spectral indices from 1984 to 2020 at both tree and stand levels, respectively, of a drought-prone Mediterranean Pinus pinea forest currently suffering strong dieback. We test the influence of climate on temporal patterns of tree radial growth, greenness and wetness spectral indices; and we address the influence of major drought episodes on resilience metrics. Tree-ring data and spectral indices followed different spatio-temporal patterns over the study period (1984–2020). Combined information from tree growth and spectral trajectories suggests that a reduction in tree density during the mid-1990s could have promoted tree growth and reduced dieback risk. Additionally, over the last decade, extreme and recurrent droughts have resulted in crown defoliation greater than 40% in most plots since 2019. We found that tree growth and the greenness spectral index were positively related to annual precipitation, while the wetness index was positively related to mean annual temperature. The response to drought, however, was stronger for tree growth than for spectral indices. Our study demonstrates the value of long-term retrospective multiscale analyses including tree and stand-level scales to disentangle mechanisms triggering and driving forest dieback.