Ecology of Drosophila aggregation pheromone: a multitrophic approach

<font size="3"><p>Many insect species use an aggregation pheromone to form groups with conspecifics in certain localities of the environment. This type of behaviour has a variety of implications for ecological interactions, both directly through the effect of the pheromone on the behaviour of con- and heterospecifics, and indirectly through the consequential aggregative distributions that may affect species interactions. The evolutionary ecology of the use of aggregation pheromone has received only little attention. Yet, these pheromones may play an intricate role in food web interactions by providing an accompanying information web.</p><p>The aim of this thesis is to further our understanding on the ecological and evolutionary aspects of the use of aggregation pheromone in insects. By unravelling costs and benefits that arise from the use of aggregation pheromone in our ecological model organism, <em>Drosophila melanogaster</em> , we strive to answer <em>why</em> they use an aggregation pheromone and elucidate the ecological consequences of an aggregation pheromone in a food web context.</p><p>In laboratory and field studies, we identified behaviours and interactions of the fruit fly <em>D. melanogaster</em> that were affected by its aggregation pheromone. The pheromone affected the distribution of adults, their eggs, competitor species and parasitoids. Moreover, a number of costs and benefits to the use of aggregation pheromone were indicated. In subsequent studies, the major hypotheses on costs and benefits were examined.</p><p>A major benefit of using aggregation pheromone was shown to be aggregated oviposition. Aggregated oviposition enhanced the quality of the larval resource, as indicated by a higher survival of the larvae and larger size of the emerging flies. This Allee effect was characterised by a positive effect of adult density on larval fitness components, and may have arisen from the interaction between adult flies and micro-organisms (yeasts and fungi). Fungi antagonise yeast and larval development, while adults can inoculate yeast on a substrate and temper fungal growth. Larvae also tempered fungal growth, but an increased larval density did not result in an Allee effect but in competition instead.</p><p>A major cost of using aggregation pheromone arose from an increased risk of parasitism. The parasitoid <em>Leptopilina heterotoma</em> uses the aggregation pheromone of adult fruit flies to localise the larval hosts, and based on this information this parasitoid can differentiate quantitatively at long range between substrates that differ in profitability. After arrival on a substrate, the pheromones no longer play a role in the host searching behaviour. A behaviour-based model was developed to predict the individual risk of parasitism for hosts in differently sized host aggregations. The functional and numerical responses of the parasitoids were combined with a flexible patch leaving decision rule for the parasitoid, to assess whether aggregation could also comprise a benefit to the hosts in terms of a diluted risk ( <em>sensu</em> Hamilton 1971). The model prediction reads that aggregation is not beneficial in the context of the <em>Drosophila</em> - <em>Leptopilina</em> interaction, and these predictions were supported by field data.</p><p>In a simple spatio-temporal simulation model, the population dynamics arising from several modes of dispersal, food competition and an Allee effect were explored. The model is a first step towards a more extensive model that incorporates the responses of insects to spatially heterogeneous resources and chemical information (e.g., aggregation pheromone).</p><p>The main conclusion from this thesis is that the aggregation pheromone of <em>D. melanogaster</em> plays an intricate role within a foodweb context, and that a variety of costs and benefits arise from multitrophic interactions. To understand the dynamic interactions in this and many other ecological systems, it is essential to gain more insight into the effect of aggregation pheromone on the behaviour of individuals.</p></font&gt