Universal foliage-stem scaling across environments and species in dicot trees: plasticity, biomechanics and Corner's Rules

Trees range from small-leaved, intricately branched species with slender stems to large-leaved, coarsely branched ones with thick stems. We suggest a mechanism for this pattern, known as Corner's Rules, based on universal scaling. We show similar crown area–stem diameter scaling between trunks and branches, environments, and species spanning a wide range of leaf size and stem biomechanics. If crown and stem maintain metabolically driven proportionality, but similar amounts of photosynthates are produced per unit crown area, then the greater leaf spacing in large-leaved species requires lower density stem tissue and, meeting mechanical needs, thicker stems. Congruent with this scenario, we show a negative relationship between leaf size and stem Young's modulus. Corner's Rules emerge from these mutual adjustments, which suggest that adaptive studies cannot consider any of these features independently. The constancy of scaling despite environmental challenges identifies this trait constellation as a crucial axis of plant diversification.