Wardhaugh, C.W., & Didham, R.K. 2005 Density-dependent effects on the reproductive fitness of the New Zealand beech scale insect (Ultracoelostoma assimile) across multiple spatial scales.. Ecological Entomology 30(6): 733-738.

Notes: Intraspecific competition can be an important factor influencing the individual fitness of organisms. This study reports density-dependent effects on the fecundity of the beech scale insect (Ultracoelostoma assimile, Hemiptera: Margarodidae) on naturally occurring, canopy-dominant red beech (Nothofagus fusca, Fagaceae) trees in New Zealand. For the first time an increasing intensity of intraspecific competition at increasing spatial scales within individual host trees is demonstrated. Beech scale insect 'tests' containing adult female scale insects and eggs were collected from the trunks of 10 red beech trees [17.7-48.5 cm in diameter at breast height (1.4 m), d.b.h.] with varying densities of scale insect infestation. The relationship between individual female fecundity and scale insect density at three spatial scales: local (within 5 cm), lower trunk (below 2 m above ground level), and whole tree, was tested. Beech scale insect fecundity was density dependent, with total female egg load decreasing with increasing scale insect density. The strength of the density-dependent effect increased with increasing spatial scale, suggesting that scale insects are creating a tree-wide drain on the quality of phloem sap, rather than depleting nutrients from localized, high-density areas within trees. These results indicate that at high densities, the New Zealand beech scale insect can have a negative effect on the nutritional quality of the phloem of red beech, thus negatively affecting conspecifics elsewhere on individual host trees. The increasing intensity of the effect with increasing spatial scale within individual trees emphasizes the importance of measuring density-dependent effects at the appropriate spatial scale.