Lepidoptera demonstrate the relevance of Murray’s Law to circulatory systems with tidal flow

Abstract

Background: Murray’s Law, which describes the branching architecture of bifurcating tubes, predicts the morphology of vessels in many amniotes and plants. Here, we use insects to explore the universality of Murray’s Law and to evaluate its predictive power for the wing venation of Lepidoptera, one of the most diverse insect orders. Lepidoptera are particularly relevant to the universality of Murray’s Law because their wing veins have tidal, or oscillatory, flow of air and hemolymph. We examined over one thousand wings representing 667 species of Lepidoptera. Results: We found that veins with a diameter above approximately 50 microns conform to Murray’s Law, with veins below 50 microns in diameter becoming less and less likely to conform to Murray’s Law as they narrow. The minute veins that are most likely to deviate from Murray’s Law are also the most likely to have atrophied, which prevents efficient fluid transport regardless of branching architecture. However, the veins of many taxa continue to branch distally to the areas where they atrophied, and these too conform to Murray’s Law at larger diameters (e.g., Sesiidae). Conclusions: This finding suggests that conformity to Murray’s Law in larger taxa may reflect requirements for structural support as much as fluid transport, or may indicate that selective pressures for fluid transport are stronger during the pupal stage—during wing development prior to vein atrophy—than the adult stage. Our results increase the taxonomic scope of Murray’s Law and provide greater clarity about the relevance of body size.