Scaling patterns of biomass and soil properties: an empirical analysis

We argue that studies at multiple scales must necessarilychange the extent of measurements, not just the spacing, in order toeffectivelycapture information regarding processes at multiple scales. We have implementeda multi-scale sampling scheme using transects of 10 cm, 1m, 10 m, 100 m, and 1 km ateach of four sites along an elevational gradient from dry foothills forest toalpine tundra in the Front Range of Colorado; these four sites form anadditional transect of 22 km. Along each of these transects wetookten equally spaced soil cores and measured variables important in determiningboth microbial and plant community structure: soil water content, organicmattercontent, pH, and total soil biomass. With this sampling scheme we are able totreat scale as an independent variable in our analyses, and our data show thatboth particular sites and particular variables can determine whether estimatesof mean values are scale-dependent or not. A geostatistical analysis using allof our data shows common relationships between scales across ecologicallydiverse sites; biomass shows the most complex pattern of distribution acrossscales, as measured by fractal dimension. Our analyses also reveal theinadequacy of several standard geostatistical models when applied to data frommultiple scales of measurement – we recommend the use of the boundedpowerlaw model in such cases. We hypothesize that because biological communitiesmustrespond simultaneously to multiple variables with differing patterns of spatialvariation, the spatial variation of biological communities will be at least ascomplex as the most complex environmental variable at any given site.This revised version was published online in May 2005 with corrections to the Cover Date.