DEMOGRAPHY OF A SAVANNA PALM TREE: PREDICTIONS FROM COMPREHENSIVE SPATIAL PATTERN ANALYSES

Existing statistical methods for spatial pattern analysis now permit precise analyses of patterns and, given the appropriate interpretation keys, the formulation of testable hypotheses about the underlying processes. From a comprehensive analysis of the spatial pattern of a plant population and its environment, we infer scenarios of links between demographic processes, plant spatial patterns, and environmental heterogeneity. The palm tree Borassus aethiopum has easily identifiable demographic stages, a root foraging strategy enabling adults to reach distant nutrient sources, and a marked senescence starting with the onset of reproduction. We analyzed map data for palm individuals (in three different life history stages plus two sexes for adults) and for nutrient-rich patches (clumps of other tree species and termite mounds) in three vegetation types (facies) of a humid savanna of West Africa (Lamto, Ivory Coast). Spatial analyses were based on Diggle's nearest neighbor functions F and G and on Ripley's K function. The main results were as follows: (1) juveniles and seedlings are aggregated, while adults have a random pattern or are more loosely aggregated; (2) all stages except female adults are spatially associated with nutrient-rich patches, but association distances increase with stage in the life cycle; and (3) seedlings are associated with female adults, whereas the association of juveniles at longer distances is not clear-cut. We propose from these results a parsimonious scenario linking spatial pattern and mortality pattern during the life cycle. The initial pattern of seedlings (close to maternal trees) results from low dispersal distance. Later stages (older seedlings and juveniles) are mostly restricted to nutrient-rich patches through nutrient shortage away from these patches (environment-induced mortality) and form dense clumps of immature palms. Competition on nutrient-rich patches then favors the few juveniles that manage to survive farther from these patches (density-dependent mortality). Finally, the last surviving juvenile of a clump suddenly experiences almost no competition with conspecifics, due to the long distance between clumps of juveniles, and owing to its root-foraging ability, it can now recruit to the adult stage, subject only to senescence. Pattern variations among savanna facies are consistent with this scenario.