Climate and symbioses with ants modulate leaf/stem scaling in epiphytes

In most seed plants, leaf size is isometrically related to stem cross-sectional area, a relationship referred to as Corner's rule. When stems or leaves acquire a new function, for instance in ant-plant species with hollow stems occupied by ants, their scaling is expected to change. Here we use a lineage of epiphytic ant-plants to test how the evolution of ant-nesting structures in species with different levels of symbiotic dependence has impacted leaf/stem scaling. We expected that leaf size would correlate mostly with climate, while stem diameter would change with domatium evolution. Using a trait dataset from 286 herbarium specimens, field and greenhouse observations, climatic data, and a range of phylogenetic-comparative analyses, we detected significant shifts in leaf/stem scaling, mirroring the evolution of specialized symbioses. Our analyses support both predictions, namely that stem diameter change is tied to symbiosis evolution (ant-nesting structures), while leaf size is independently correlated with rainfall variables. Our study highlights how independent and divergent selective pressures can alter allometry. Because shifts in scaling relationships can impact the costs and benefits of mutualisms, studying allometry in mutualistic interactions may shed unexpected light on the stability of cooperation among species.