Origins and basic form
The nineteenth-century British botanist Hewett Watson (1859), compiling lists of British plants, recorded that larger areas contain more species. Olof Arrhenius (1921) expressed this as the power function S = cA^z, while Gleason (1922) proposed an alternative model in which species number is proportional to the logarithm of area. Taking logarithms of both sides of the power form gives a straight line, so the slope z describes how quickly species number rises with area. A commonly used rule of thumb is that a tenfold increase in area roughly doubles the number of species. The relationship is a classic biological example of a power law.
Theoretical explanations and island biogeography
Preston (1962) proposed the canonical distribution theory, showing that if species abundances follow a lognormal distribution, the z-value of the species–area curve is about 0.26. MacArthur and Wilson's (1967) theory of island biogeography held that the number of species on an island is set by a balance between immigration of new species and extinction of existing ones, and that larger islands have lower extinction rates and so sustain more species. Connor and McCoy (1979) distinguished habitat diversity, sampling effects and immigration–extinction balance as causes of the relationship and set out the statistical issues involved. Rosenzweig (1995) synthesized evidence that the shape and slope of the curve change with spatial scale.
Conservation applications and limits
Drawing on the species–area relationship, Diamond (1975) proposed the design principle that a single large reserve is better than several small ones, and Simberloff and Abele (1976) countered that this need not be so, starting what became known as the SLOSS debate. Pimm et al. (1995) and Thomas et al. (2004) used the relationship to estimate future extinctions from habitat loss and climate change. He and Hubbell (2011), however, argued that applying the species–area curve in reverse overestimates extinction rates from habitat loss. Triantis, Guilhaumon and Whittaker (2012) compared which mathematical models best fit island data, addressing both biological interpretation and statistical model choice.
FAQ
- What is the species–area relationship in simple terms?
- The larger the area you survey, the more species you find, and the increase follows a regular mathematical rule. Roughly, ten times the area gives about twice the species.
- Why does it matter for conservation?
- It is used to think about the size and design of reserves and to estimate how many species habitat loss might eliminate, though it can overestimate extinction rates.
Sources reviewed: 2026-10-26
References
- Arrhenius, O. (1921). Species and area. Journal of Ecology, 9(1), 95–99.
- Gleason, H. A. (1922). On the relation between species and area. Ecology, 3(2), 158–162.
- Watson, H. C. (1859). Cybele Britannica (Vol. 4). London: Longman.
- Preston, F. W. (1962). The canonical distribution of commonness and rarity: Part I. Ecology, 43(2), 185–215.
- MacArthur, R. H., & Wilson, E. O. (1967). The Theory of Island Biogeography. Princeton: Princeton University Press.
- Connor, E. F., & McCoy, E. D. (1979). The statistics and biology of the species–area relationship. American Naturalist, 113(6), 791–833.
- Diamond, J. M. (1975). The island dilemma: Lessons of modern biogeographic studies for the design of natural reserves. Biological Conservation, 7(2), 129–146.
- Simberloff, D. S., & Abele, L. G. (1976). Island biogeography theory and conservation practice. Science, 191(4224), 285–286.
- Rosenzweig, M. L. (1995). Species Diversity in Space and Time. Cambridge: Cambridge University Press.
- Pimm, S. L., Russell, G. J., Gittleman, J. L., & Brooks, T. M. (1995). The future of biodiversity. Science, 269(5222), 347–350.
- Thomas, C. D., et al. (2004). Extinction risk from climate change. Nature, 427(6970), 145–148.
- He, F., & Hubbell, S. P. (2011). Species–area relationships always overestimate extinction rates from habitat loss. Nature, 473(7347), 368–371.
- Triantis, K. A., Guilhaumon, F., & Whittaker, R. J. (2012). The island species–area relationship: Biology and statistics. Journal of Biogeography, 39(2), 215–231.
