Where do seedlings go? A spatio-temporal analysis of seedling mortality in a semi-arid gypsophyte

Studies of seedling population dynamics often focus on survival because it provides an integrated measure of seedling performance. However, this approach involves a substantial loss of information because survival is the net result of a wide range of mechanisms. The present study overcomes these shortcomings by investigating spatial and temporal patterns in the causes of plant mortality in a population of Helianthemum squamatum seedlings. We use new point pattern analyses based on K functions combined with a new null model ("independent labeling"). A total of 871 seedlings of H. squamatum were mapped and regularly monitored over an 18-month period. More than 60% of seedlings died during this period. Causes of mortality were spatially structured, and these structures shifted through time. Small differences in either the time of emergence or the environment surrounding H. squamatum seedlings had profound influences on their fate. Seedlings emerging late in the season under the canopy of adult plants died from drought more often than expected, whereas those emerging earlier in the same microsite survived more than expected. The identity of neighbors also affected the spatio-temporal dynamics of mortality causes. Our results show that seedling-adult interactions cannot be easily predicted from simple models, and that the time of seedling emergence, its age and the identity of its neighbors determine the sign and the spatial scale of these interactions. The new methods introduced in this article open an avenue for the detailed analyses of the spatio-temporal dynamics of plant mortality and can help to disentangle the complexity of biotic interactions along environmental severity gradients.