Contrasting trait variations and trait correlation networks across heteromorphic leaves of <i>Populus euphratica</i>

Abstract Traits and their correlation networks can reflect plant adaptive strategies. However, variations in traits and trait correlation networks across heteromorphic leaves within species remain largely unexplored. In this study, we systematically quantified a diverse array of leaf traits—spanning morphology, anatomy, physiology and biochemistry—among the striped, lanceolate, ovate, and broadly ovate leaves of Populus euphratica, aiming to elucidate the adaptive differences across these various leaf types. We found that the four heteromorphic leaves showed significant differences in leaf traits. From striped leaves to broadly ovate leaves, leaf size, leaf thickness, water use efficiency and catalase content significantly increased, while specific leaf area showed the opposite pattern. Principal component analysis and cluster analysis revealed distinct aggregation and clear demarcation of the four leaf types, indicating substantial variations in trait compositions and their distinct ecological adaptations. Plant trait networks varied significantly across the four leaf types, with the broadly ovate leaves exhibiting a fragmented network structure that enhances their modularity. This suggests strong resilience to disturbances and is consistent with the characteristic foliage on mature trees. Regardless of leaf type, nitrogen and phosphorus consistently emerged as hub traits within plant trait networks, underscoring their fundamental role in driving physiological processes and influencing phenotypic expression. This study meticulously delineates the variations in both individual leaf traits and trait correlation networks across the heteromorphic leaves of P. euphratica, significantly deepening our understanding of plant adaptive strategies.