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Microstructures on insect wings can promote directional drop shedding, and the local orientation of these structures is expected to facilitate drop removal. However, microstructures may exhibit very different orientations at different locations on the wing. Using the march fly Penthetria heteroptera, we propose that local orientation of small hairs (microtrichia) reflects a balance of three nonexclusive strategies: (1) preventing water from becoming stuck in intervenous grooves (microtrichia point upslope), (2) shedding water off the wing as readily as possible (microtrichia point towards the nearest edge), and, (3) shedding water away from the body (microtrichia point distally). We present evidence for all three and show that local microtrichial orientation is seldom determined by any one factor. We develop a mathematical model that employs factor-specific weighting values determined via optimization. Our predictions are tested against the orientation of microtrichia randomly sampled from a P. heteroptera specimen. Using the best-fit weighting parameters, the model displays a median residual of 20°; no residual is greater than 46°. The model also reproduces qualitative aspects of microtrichial orientation, such as bifurcation midway between veins and convergence toward peaks. This strong correspondence between modelled and observed orientation supports the role of microtrichia as directional antiwetting devices and highlights the importance of considering both function and wing geometry to explain the organization of natural microstructure arrays.

Hybridization between distinct populations or species is increasingly recognized as an important process for generating biodiversity. However, the interaction between hybridization and speciation is complex, and the diverse evolutionary outcomes of hybridization are difficult to differentiate. Here we characterize potential hybridization in a species group of swallowtail butterflies using microsatellites, DNA sequences, and morphology, and assess whether adaptive introgression or homoploid hybrid speciation was the primary process leading to each putative hybrid lineage. Four geographically separated hybrid populations were identified in the Papilio machaon species group. One distinct mitochondrial DNA clade from P. machaon was fixed in three hybrid taxa (P. brevicauda, P. joanae, and P. m. kahli), while one hybrid swarm (P. zelicaon x machaon) exhibited this hybrid mtDNA clade as well as widespread parental mtDNA haplotypes from both parental species. Microsatellite markers and morphology showed variable admixture and intermediacy, ranging from signatures of prolonged differential introgression from the paternal species (P. polyxenes/P. zelicaon) to current gene flow with both parental species. Divergences of the hybrid lineages dated to early- to mid-Pleistocene, suggesting that repeated glaciations and subsequent range shifts of parental species, particularly P. machaon hudsonianus, facilitated initial hybridization. Although each lineage is distinct, P. joanae is the only taxon with sufficient evidence (ecological separation from parental species) to define it as a homoploid hybrid species. The repetition of hybridization in this group provides a valuable foundation for future research on hybridization, and these results emphasize the potential for hybridization to drive speciation in diverse ways.