Distribution of Insect Attacks in Biological Control of Weeds: Infestation of Centaurea virgata Flowerheads by a Gall Fly

The success of biological control efforts to reduce weed density through release of insects may depend as much on the distribution of insect attacks among individual plants or plant parts as on the mean level of infestation. We used an index of dispersion to describe the distribution of Urophora quadrifasciata (Diptera: Tephritidae) galls among squarrose knapweed (Centaurea virgata) flowerheads at 18 west central Utah sites in the first 5 years following introduction of the biological control agent. Two thirds of the samples showed a significantly aggregated distribution of galls among flowerheads. Statistical analysis showed that site and year accounted for relatively small proportions of the variance in the index of dispersion. The degree of gall aggregation among flowerheads was positively correlated with the mean flowerhead quality (mean number of seeds per flowerhead; P = 0.013) and tended to be negatively correlated with the mean fly density per flowerhead at a site in a given year (P = 0.097). Our data suggest that higher quality flowerheads, and possibly higher quality plants, are preferentially attacked by U. quadrifasciata and therefore are more heavily subject to reduced reproductive potential through biological control. However, an aggregated distribution of fly attacks may undercut the potential of the fly to reduce seed production by the weed population as a whole. Understanding both the distribution of insect attacks among individual plants and the behavioral mechanisms producing such distribution patterns is important to the biological control of weeds.