, Volume 186, Issue 3, pp 805–816 | Cite as

Shifts in the importance of the species pool and environmental controls of epiphytic bryophyte richness across multiple scales

  • Nagore G. MedinaEmail author
  • Matthew A. Bowker
  • Joaquín Hortal
  • Vicente Mazimpaka
  • Francisco Lara
Community ecology – original research


Species richness is influenced by a nested set of environmental factors, but how do these factors interact across several scales? Our main aim is to disentangle the relative importance of environmental filters and the species pool on the richness of epiphytic bryophytes across spatial scales. To do so, we sampled epiphytic bryophytes in 43 oak forests across the northwest of the Iberian Peninsula. As predictors we used climate, descriptors of forest structure and micro-environment. We applied structural equation modeling to relate these variables with richness and cover at three scales: locality (forest), stand (three stands per forest), and sample (a quadrate in a tree). We assumed top–down relationships, so that large-scale variables influenced lower scale variables, and in which cover directly influenced richness. Richness at the next larger scale (locality to stand and stand to sample) is considered a surrogate of the species pool and included as a predictor of richness at the next smaller scale. Environmental variables explain locality richness, but as we decrease the spatial scale, its importance decreases and the dependence on species pool increases. In addition, we found unexpected bottom–up relationships (between micro-scale environment to locality richness). Our results point to the scale dependence of niche vs. neutral processes: niche processes are important at the locality (forest) scale, while neutral processes are significant at the small (sample) scale. We propose a modified conceptualization of the factors influencing biodiversity at different spatial scales by adding links across different scales (between micro-environment and locality-scale richness in our study).


Plant Diversity Neutral processes Regional species pool Scale 



This research has been funded by the Spanish Ministry of Science and Innovation (Grants CGL2007-61389, CGL2010-15693 and CGL2011-28857). JH is supported by a Spanish Dirección General de Ciencia y Tecnología (DGCyT) Ramón y Cajal Grant. NGM is supported by the Grant AP2007-00905 of the Spanish Ministry of Science. We would like to thank two anonymous referees for their thorough review and constructive comments.

Author contribution statement

NGM, FL, JH, and VM originally formulated the idea; NGM and VM conducted fieldwork; NGM and MAB analyzed the data; NGM lead the writing process; FL, JH, MAB, and VM actively contributed to discuss the results and write the manuscript.

Supplementary material

442_2018_4066_MOESM1_ESM.doc (1.6 mb)
Supplementary material 1 (DOC 1635 kb)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  • Nagore G. Medina
    • 1
    • 2
    Email author
  • Matthew A. Bowker
    • 3
  • Joaquín Hortal
    • 4
  • Vicente Mazimpaka
    • 1
  • Francisco Lara
    • 1
  1. 1.Departamento de Biología (Botánica), Facultad de CienciasUniversidad Autónoma de MadridMadridSpain
  2. 2.Department of Botany, Faculty of SciencesSouth Bohemia UniversityCeske BudejoviceCzech Republic
  3. 3.School of ForestryNorthern Arizona UniversityFlagstaffUSA
  4. 4.Departamento de Biogeografía y Cambio GlobalMuseo Nacional de Ciencias Naturales (MNCN-CSIC)MadridSpain

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