The forcibly launched spores of ascomycete fungi must eject through several millimeters of nearly still air surrounding fruiting bodies to reach dispersive air flows. Because of their microscopic size, spores experience great fluid drag, and although this drag can aid transport by slowing sedimentation out of dispersive air flows, it also causes spores to decelerate rapidly after launch. We hypothesize that spores are shaped to maximize their range in the nearly still air surrounding fruiting bodies. To test this hypothesis we numerically calculate optimal spore shapes-shapes of minimum drag for prescribed volumes-and compare these shapes with real spore shapes taken from a phylogeny of >100 species. Our analysis shows that spores are constrained to remain within 1% of the minimum possible drag for their size. From the spore shapes we predict the speed of spore launch, and confirm this prediction through high-speed imaging of ejection in Neurospora tetrasperma. By reconstructing the evolutionary history of spore shapes within a single ascomycete family we measure the relative contributions of drag minimization and other shape determinants to spore shape evolution. Our study uses biomechanical optimization as an organizing principle for explaining shape in a mega-diverse group of species and provides a framework for future measurements of the forces of selection toward physical optima.hydrodynamics | biological optimization | fungal spores M any organisms have visible adaptations for minimizing drag, including the streamlined shapes of fast-swimming fish and of Mayfly nymphs that cleave to rocks in rapidly flowing streams, or the precisely coordinated furling of tulip tree leaves in strong winds (1). However, although drag minimization may improve some aspects of individual fitness, it is also clear that physiological and ecological trade-offs will constrain the evolution of body shape. Similarly, although signatures of optimization can be seen in diverse features of organism morphology (2), behavior (3), and resource allocation (4, 5), the strength of the force of selection for one physical optimum over another can not be readily quantified (6).Most fungi grow on highly heterogeneous landscapes and must move between disjoint patches of suitable habitat. Sexual spores can be carried by air flows and allow dispersal between patches (7, 8). To reach dispersive air flows, spores of many species of ascomycete fungi are ejected from asci, fluid-filled sacs containing the spores (9). At maturity the turgor pressure within each sac climbs until a critical pressure is reached, whereupon a hole opens at the apex and spores are ejected (8). Spores must travel far enough from the originating fruiting body to enter dispersive air flows. In particular, it is necessary that they pass through a boundary layer of still air of thickness ∼1 mm that clings to the fungal fruiting body (10).Multiple independently evolved adaptations enhance spore range in disparate fungal species. For example, appendages and mucilaginous sheathes pro...