Wardhaugh, C.W., Blakely, T.J., Greenberg, R., Morris, P.D., Barnden, A., Rickard, S., Atkinson, B., Fagan, L.L., Ewers, R.M., & Didham, R.K. 2006 Vertical stratification in the spatial distribution of the beech scale insect (Ultracoelostoma assimile) in Nothofagus tree canopies in New Zealand.. Ecological Entomology 31(2): 185-195.

Notes: The degree of infestation by New Zealand sooty beech scale insects (Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees (Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full-census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy-dominant red beech (Nothofagus fusca) trees (size range 38.7-107.6 cm diameter at breast height) to determine the degree of within-tree heterogeneity in herbivore density. The within-tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time. Because of the high degree of within-tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population.