Figure 1: Tinbergen’s four quadrat framework applied to evolutionary questions about Escovopsis parasitism of fungus-farming ant crops. Ontogeny refers to the description of development, from DNA to progressive phenotype, mechanism refers to the physiological and cellular processes that organisms have available, phylogeny refers to the idiosyncratic evolutionary history of a lineage, and adaptation refers to traits that acquired their extant function because of specific selective advantages, modified from [6668].