Herbivore pressure modulates soil multifunctionality in a Mediterranean landscape

Henriques, J. F.; Carvalho, J.; Torres, R. T.; Rossa, M.; Fernandes, J. M.; Malheiro, C.; Prodana, M.; Peixoto, S.; Fereira, E.; Loureiro, S.; Krummins, J.; Perea, R.; Serrano, E.; Morgado, R. G.

doi:10.64898/2025.12.12.693896

bioRxiv · 10.64898/2025.12.12.693896

Herbivore pressure modulates soil multifunctionality in a Mediterranean landscape

Henriques, J. F.·Carvalho, J.·Torres, R. T.·Rossa, M.·Fernandes, J. M.·Malheiro, C.·Prodana, M.·Peixoto, S.·Fereira, E.·Loureiro, S.·Krummins, J.·Perea, R.·Serrano, E.·Morgado, R. G.

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Abstract

Ongoing global environmental changes demand cost-effective strategies to restore ecosystem functioning. Within this context, the reintroduction of large mammalian herbivores has emerged as a promising approach to recover ecological processes and foster self-sustaining, biodiverse ecosystems. However, the impacts of such initiatives remain poorly understood, particularly regarding their effects on soil functions and ecological processes. We tested whether large herbivore densities (wild horses (Garrano breed) and cattle (Maronesa breed, an ancient lineage close to aurochs) modulates soil multifunctionality (the simultaneous provision of multiple ecosystem functions) across habitats (grasslands, shrublands, and forests) and seasons (fall and spring) in a Mediterranean landscape. Using composite multivariate indices describing soil functions related to biogeochemical cycles (i.e., enzymatic activities) and physicochemical properties (pH, conductivity, organic matter, and nutrient loads), we found that soil enzymatic activities varied seasonally, being higher in spring, and interacted with habitat and herbivore pressure gradients. Habitat and herbivore pressure explained the variation in soil multifunctionality during spring, while in fall it was mainly driven by herbivore pressure. Enzymatic stoichiometry, particularly C:P ratios, strongly predicted soil fertility and multifunctionality, showing positive relationships in forest habitats and under herbivore pressure. A mechanistic approach confirmed that herbivore impacts on soil functioning operated primarily through changes in soil properties and nutrient cycling rather than direct effects. Mediterranean landscapes are rapidly changing, and our results highlight herbivore management as a key tool to sustain soil ecological processes under the rewilding framework. Highlights- Large herbivores impacted soil functions and properties in different habitats - Large herbivores significantly impacted soil nutrient cycling and multifunctionality. - The magnitude of the effects was dependent on season and habitat - Large herbivores indirectly impact soil ecosystem multifunctionality through changes in soil properties.

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Henriques, J. F., Carvalho, J., Torres, R. T., Rossa, M., Fernandes, J. M., Malheiro, C., Prodana, M., Peixoto, S., Fereira, E., Loureiro, S., Krummins, J., Perea, R., Serrano, E., Morgado, R. G.. 2025-12-14. Herbivore pressure modulates soil multifunctionality in a Mediterranean landscape. https://doi.org/10.64898/2025.12.12.693896

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