Ontogenetic correlates, not direct adaptation, explain the evolution of stelar morphology

The primary vascular system of plants (the stele) has attracted interest from paleobotanists, developmental biologists, systematists, and physiologists for nearly two centuries. Ferns, with their diverse stelar morphology, deep evolutionary history, and prominent fossil record, have been a major focus in studies of the stele. To explain the diversity of stelar morphology, past adaptive hypotheses have invoked biomechanics, hydraulics, and drought tolerance as key selection pressures in the evolution of stelar complexity; but, these hypotheses often isolate the stele from a whole-plant developmental context, ignoring potential covariation between vascular patterning and shoot morphology. Furthermore, incongruence between expected patterns and observed data challenge adaptive hypotheses, precluding a comprehensive explanation of stelar evolution. While ontogeny has been previously recognized as a factor in stelar diversification, it has not been fully integrated into a comprehensive framework. Here we synthesize 150-years of research on stelar morphology, incorporating developmental, physiological, and phylogenetic data to present the ontogenetic hypothesis of stelar evolution. This hypothesis posits that stelar morphology is an integrated feature of whole-plant ontogeny, not a trait shaped by direct selection for adaptive patterns. This shift in perspective provides an updated framework for understanding the determinants of stelar morphology and focusses future efforts to ask more incisive questions about the evolution and function of primary vascular architecture.