Seedling dynamics of Acer mono and Fagus crenata: an environmental filter limiting their adult distributions

Spatial pattern of a biotic population may change over time as its component individuals grow or die out, but whether this is the case for desert annual plants is largely unknown. Here we examined seasonal changes in spatial patterns of two annuals, Eriogonum abertianum and Haplopappus gracilis, in initial (winter) and final (summer) densities. The density was measured as the number of individuals from 384 permanent quadrats (each 0.5 m × 0.5 m) in the Chihuahuan Desert near Portal, Arizona, USA. We used three probability distributions (binomial, Poisson, and negative binomial or NB) that represent three basic spatial patterns (regular, random, and clumped) to fit the observed frequency distributions of densities of the two annuals. Both species showed clear clumped patterns as characterized by the NB and had similar inverse J-shaped frequency distribution curves in two density categories. Also, both species displayed a reduced degree of aggregation from winter to summer after the spring drought (massive die-off), as indicated by the increased k-parameter of the NB and decreased values of another NB parameter p, variance/mean ratio, Lloyd’s Index of Patchiness, and David and Moore’s Index of Clumping. Further, we hypothesized that while the NB (i.e., Poisson-logarithmic) well fits the distribution of individuals per quadrat, its components, the Poisson and logarithmic, may describe the distributions of clumps per quadrat and of individuals per clump, respectively. We thus obtained the means and variances for (1) individuals per quadrat, (2) clumps per quadrat, and (3) individuals per clump. The results showed that the decrease of the density from winter to summer for each plant resulted from the decrease of individuals per clump, rather than from the decrease of clumps per quadrat. The great similarities between the two annuals indicate that our observed temporal changes in spatial patterns may be common among desert annual plants.