Landscape Ecology

, Volume 24, Issue 4, pp 509–518 | Cite as

Boundaries and scales in shrublands of the Doñana Biological Reserve, southwest Spain

  • José Carlos Muñoz-Reinoso
Research Article


To verify that the stabilized sand dunes of Doñana, southwest Spain, are hierarchically nested, vegetation was sampled along topographic gradients at three spatial scales and Split Moving Window Boundary Analysis was applied to identify vegetation boundaries and ecotones. At small scale, only one window width was used, while for boundaries detection at upper scales the information from five windows was pooled. Environmental factors controlling plant composition were studied along topographic gradients, and diversity was estimated within the boundaries. According to several theoretical frameworks, I discuss the types of boundaries produced at different scales. Lower level boundaries are characterized by transitory gradients linked to local exchanges; intermediate boundaries are symmetric and very stable over the time; the large scale boundary is asymmetric with strong inherent abiotic constraints reinforced by strong biotic feedbacks. In spite of a similar plant composition, a plant community, the mixed shrub, works as an ecocline or an ecotone depending on the spatial scale considered. A certain parallelism exists between shrub composition along dune slopes and dune generations; however, processes at upper scale constraint plant composition at lower scale resulting in different mature formations.


Boundaries Doñana Ecocline Ecotone Nested hierarchy Split Moving Window Shifting transition Stationary transition 



Doñana Biological Reserve


Split Moving Window Boundary Analysis


Multi-response permutation procedures



I greatly thank Dr de Castro and Dra Fernández-Alés for their suggestions during the preparation of the paper. I also thank the two anonymous reviewers for their helpful comments on the manuscript. The Doñana Biological Station provided permission to carry out the field survey in the Reserve.


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Copyright information

© Springer Science+Business Media B.V. 2009

Authors and Affiliations

  1. 1.Departamento de Biología Vegetal y EcologíaUniversidad de SevillaSevillaSpain

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