Notes: Abstract A Monte Carlo method based on Ripley's K function-a cumulative function related to the number of plants encountered at different distances from other plants-is used to test the null hypothesis of random distribution of shrub clumps in a desert dwarf shrub community in Namaqualand, South Africa, where Psilocaulon arenosum is the dominant shrub. The method takes into account the apparent regularity of pattern caused by the finite size (up to 2 m diameter) of the clumps. It is shown that the clump centres are significantly aggregated (compared to random expectation) at distances on the order of 1 m. Such aggregation is expected, as a simple result of regeneration near to seed sources, if the time between catastrophic droughts is short in relation to the time required for development of a non-aggregated or regular pattern determined by moisture competition. No significant regulatiry was detected at distances of 3 m or less. One subplot showed regularity above 3 m, but this pattern was not shown by the other subplot and may not be a competition effect. These results support a hypothesis of aggregation caused by regeneration pattern decaying slowly toward randomness as larger individuals compete.

Notes: Abstract Range limits and broad-scale geographic variations in the productivity of boreal and northern deciduous tree species in Europe are simulated with a bioclimatic model (STASH). STASH is based on a small number of distinct mechanisms by which climate is thought to affect the survival, regeneration and growth of trees. Survival is limited by summer warmth requirements and winter cold tolerance; regeneration, additionally by winter chilling requirements; and growth rate by net assimilation, which in turn is related to photosynthetically active radiation, growing-season length, temperature (relative to species-specific optima) and soil moisture supply (relative to evaporative demand). These mechanisms are quantified either as thresholds (for survival and regeneration) or multipliers (for growth), based on bioclimatic variables computed from monthly climate normals interpolated three-dimensionally to a 10-minute grid. Growing-season and growing degree day calculations take into account the effects of chilling. The drought calculations also take into account the effects of soil-moisture storage by means of a physically based evapotranspiration calculation coupled to a one-layer soil hydrology model. STASH is used to examine changes in potential range limits under a 2xCO2 climate-change scenario. Associated transient responses at selected sites in the boreal and boreo-nemoral zones of Sweden are also simulated, using the forest gap model FORSKA2. The species-specific survival and regeneration constraints and growth responses of STASH modify the growth, establishment and mortality of trees in FORSKA2. The results obtained in this way differ sharply from the results of conventional forest gap models, where growth rates are assumed to decline to zero at minimum and maximum growing degree day limits. For example, towards the southern limit ofPicea abies (Norway spruce), STASH correctly shows no decline in productivity, but rather an abrupt cut-off corresponding to a chilling requirement during regeneration that is not met further south or west. In transient warming scenarios, this mechanism has the effect that natural regeneration can be blocked due to the warm winters even as yield is increasing due to the longer and warmer summers. STASH predicts drastic changes in species distributions in response to the large climate changes (especially winter warming) expected for northern Europe. Some of the common boreal species (e.g.,Picea abies; Pinus sylvestris; Alnus incana) are unlikely to survive in much of their present range, withdrawing to the far north. Other species already widespread may be able to occupy some of the few sites that are today unavailable to them (e.g.,Betula spp.;Corylus avellana). Other temperate deciduous species such asFagus sylvatica could have dramatic range expansions, potentially occupying large tracts of the present boreal zone. FORSKA2 transient simulations illustrate some of the possible routes towards different types of forest in a changed climate. Some sites in the north show little change in species composition, but sites towards the southern boundary of the boreal zone could develop a new suite of dominants. The degree of sensitivity of a particular site depends both on the climate change prediction and on the transient dynamics of the forest community. Many types of transient behaviour are shown to be possible. Coupled with uncertainties about the future role of dispersal and changes in disturbance rate, the complexity and variety of these transient responses imply a highly uncertain future for the north European boreal forests.