Biogeochemistry and Geographical Ecology: Embracing All Twenty-Five Elements Required to Build Organisms.

Biogeochemistry is a key but relatively neglected part of the abiotic template that underlies ecology. The template has a geography, one that is increasingly being rearranged in this era of global change. Justus von Liebig's law of the minimum has played a useful role in focusing attention on biogeochemical regulation of populations, but given that ∼25+ elements are required to build organisms and that these organisms use and deplete nutrients in aggregates of communities and ecosystems, we make the case that it is time to move on. We review available models that suggest the many different mechanisms that give rise to multiple elements, or colimitation. We then review recent empirical data that show that rates of decomposition and primary productivity may be limited by multiple elements. In that light, given the tropics' high species diversity and generally more weathered soils, we predict that colimitation at community and ecosystem scales is more prevalent closer to the equator. We conclude with suggestions for how to move forward with experimental studies of colimitation.