Measurements of area and the (island) species–area relationship: new directions for an old pattern


  • Kostas A. Triantis,

  • David Nogués-Bravo,

  • Joaquín Hortal,

  • Paulo A. V. Borges,

  • Henning Adsersen,

  • José María Fernández-Palacios,

  • Miguel B. Araújo,

  • Robert J. Whittaker

K. A. Triantis ( and R. J. Whittaker, Biodiversity Research Group, Oxford Univ. Centre for the Environment, South Parks Road, Oxford, OX1 3QY, UK. – D. Nogués-Bravo and M. B. Araújo, Dept of Biodiversity and Evolutionary Biol., National Museum of Natural Sciences, CSIC, C/Gutiérrez Abascal 2, ES–28006, Madrid, Spain. – J. Hortal, NERC Centre for Population Biology, Imperial College London, Silwood Park Campus, Ascot, SL5 7PY, UK. – P. A. V. Borges, Azorean Biodiversity Group, Depto de Ciências Agrárias – CITA-A, Univ. dos Açores, Angra do Heroísmo, Terceira, Açores, Terra-Chã, 9700-851. – H. Adsersen, Centre for Macroecology, Dept of Biology, Univ. of Copenhagen, Universitetsparken 15, DK–2100 Copenhagen Ø, Denmark. – J. M. Fernández-Palacios, Depto de Ecología, Facultad de Biología, Univ. de La Laguna, ES–38206 La Laguna, Tenerife, Islas Canarias, Spain.


The species–area relationship is one of the strongest empirical generalizations in geographical ecology, yet controversy persists about some important questions concerning its causality and application. Here, using more accurate measures of island surface size for five different island systems, we show that increasing the accuracy of the estimation of area has negligible impact on the fit and form of the species–area relationship, even though our analyses included some of the most topographically diverse island groups in the world. In addition, we show that the inclusion of general measurements of environmental heterogeneity (in the form of the so-called choros model), can substantially improve the descriptive power of models of island species number. We suggest that quantification of other variables, apart from area, that are also critical for the establishment of biodiversity and at the same time have high explanatory power (such as island age, distance, productivity, energy, and environmental heterogeneity), is necessary if we are to build up a more predictive science of species richness variation across island systems.