Scale-dependent species-area and species-isolation relationships: A review and a test study from a fragmented semi-arid agro-ecosystem

Itamar Giladi, Felix May, Michael Ristow, Florian Jeltsch, Yaron Ziv

Research output: Contribution to journalArticlepeer-review

28 Scopus citations


Aim: Patterns that relate species richness with fragment area (the species-area relationship, SAR) and with isolation (the species-isolation relationship, SIR) are well documented. However, those that relate species density - the number of species within a standardized area - with fragment area (D-SAR) or isolation (D-SIR) have not been sufficiently explored, despite the potential for such an analysis to disentangle the underlying mechanisms of SARs and SIRs. Previous spatial theory predicts that a significant D-SAR or D-SIR is unlikely to emerge in taxa with high dispersal limitation, such as plants. Furthermore, a recent model predicts that the detection and the significance of D-SARs or D-SIRs may decrease with grain size. We combined a literature review with grain size-dependent sampling in a fragmented landscape to evaluate the prevalence and grain size-dependent nature of D-SARs and D-SIRs in plants. Location: Worldwide (review) and a semi-arid agro-ecosystem in Israel (case study). Methods: We combined an extensive literature review of 31 D-SAR studies of plants in fragmented landscapes with an empirical study in which we analysed grain size-dependent D-SARs and D-SIRs using a grain size-dependent hierarchical sampling of species density and species richness in a fragmented, semi-arid agro-ecosystem. Results: We found that significantly increasing D-SARs are rare in plant studies. Furthermore, we found that the detection of a significant D-SAR is often possible only after the data have been stratified by species, habitat or landscape characteristics. The results from our case study indicated that the significance and the slopes of both D-SARs and D-SIRs increase as grain size decreases. Main conclusions: These results call for a careful consideration of scale while analysing and interpreting the responses of species richness and species density to fragmentation. Our results suggest that grain size-dependent analyses of D-SARs and D-SIRs may help to disentangle the mechanisms that generate SARs and SIRs and may enable early detection of the effects of fragmentation on plant biodiversity.

Original languageEnglish
Pages (from-to)1055-1069
Number of pages15
JournalJournal of Biogeography
Issue number6
StatePublished - 1 Jan 2014


  • Conservation biogeography
  • Extinction debt
  • Habitat fragmentation
  • Habitat islands
  • Island biogeography theory
  • Island ecology
  • Isolation
  • Scale-dependence
  • Species density
  • Species-area relationship

ASJC Scopus subject areas

  • Ecology, Evolution, Behavior and Systematics
  • Ecology


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