The nature of serpentine endemism.

Serpentine soils are a model system for the study of plant adaptation, speciation, and species interactions. Serpentine soil is an edaphically stressful, low productivity soil type that hosts stunted vegetation and a spectacular level of plant endemism. One of the first papers on serpentine plant endemism was by Arthur Kruckeberg, titled "Intraspecific variability in the response of certain native plant species to serpentine soil." Published in the American Journal of Botany in 1951, it has been cited over 100 times. Here, I review the context and content of the paper, as well as its impact. On the basis of the results of reciprocal transplant experiments in the greenhouse, Kruckeberg made three important conclusions on the nature of serpentine plant endemism: (1) Plants are locally adapted to serpentine soils, forming distinct soil ecotypes; (2) soil ecotypes are the first stage in the evolutionary progression toward serpentine endemism; and (3) serpentine endemics are restricted from more fertile nonserpentine soils by competition. Kruckeberg's paper inspired a substantial amount of research, especially in the three areas reviewed here: local adaptation and plant traits, speciation, and the interaction of climate and soil in plant endemism. In documenting soil ecotypes, Kruckeberg identified serpentine soils as a potent selective factor in plant evolution and helped establish serpentine soils as a model system in evolution and ecology.