Hierarchical spatial segregation of two Mediterranean vole species: the role of patch-network structure and matrix composition

Ricardo Pita, Xavier Lambin, António Mira, Pedro Beja

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7 Citations (Scopus)
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According to ecological theory, the coexistence of competitors in patchy environments may be facilitated by hierarchical spatial segregation along axes of environmental variation, but empirical evidence is limited. Cabrera and water voles show a metapopulation-like structure in Mediterranean farmland, where they are known to segregate along space, habitat, and time axes within habitat patches. Here, we assess whether segregation also occurs among and within landscapes, and how this is influenced by patch-network and matrix composition. We surveyed 75 landscapes, each covering 78 ha, where we mapped all habitat patches potentially suitable for Cabrera and water voles, and the area effectively occupied by each species (extent of occupancy). The relatively large water vole tended to be the sole occupant of landscapes with high habitat amount but relatively low patch density (i.e., with a few large patches), and with a predominantly agricultural matrix, whereas landscapes with high patch density (i.e., many small patches) and low agricultural cover, tended to be occupied exclusively by the small Cabrera vole. The two species tended to co-occur in landscapes with intermediate patch-network and matrix characteristics, though their extents of occurrence were negatively correlated after controlling for environmental effects. In combination with our previous studies on the Cabrera-water vole system, these findings illustrated empirically the occurrence of hierarchical spatial segregation, ranging from within-patches to among-landscapes. Overall, our study suggests that recognizing the hierarchical nature of spatial segregation patterns and their major environmental drivers should enhance our understanding of species coexistence in patchy environments.

Original languageEnglish
Pages (from-to)253-263
Number of pages11
Issue number1
Early online date11 May 2016
Publication statusPublished - Sept 2016

Bibliographical note

Acknowledgments This study was financed by FEDER funds
through the Programa Operacional Factores de Competitividade—
COMPETE, and National funds through the Portuguese Foundation
for Science and Technology—FCT, within the scope of the projects
AAG-MAA/3227/2012), and MateFrag (PTDC/BIA-BIC/6582/2014).
RP was supported by the FCT grant SFRH/BPD/73478/2010 and
SFRH/BPD/109235/2015. PB was supported by EDP Biodiversity
Chair. We thank Rita Brito and Marta Duarte for help during field
work. We thank Chris Sutherland, Douglas Morris, William Morgan,
and Richard Hassall for critical reviews of early versions of the paper.
We also thank two anonymous reviewers for helpful comments to
improve the paper.


  • Cabrera vole
  • competition
  • landscape heterogeneity
  • patchy environments
  • species coexistence
  • Southern water vole


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