Macrostructure of Malus Leaves and Its Taxonomic Significance

Leaves are the most ubiquitous plant organs, whose macrostructures exhibit close correlations with environmental factors while simultaneously reflecting inherent genetic and evolutionary patterns. These characteristics render them highly significant for plant taxonomy, ecology, and related disciplines. Therefore, this study presents the first comprehensive evaluation of Malus leaf macrostructures for infraspecific classification. By establishing a trait-screening system, we conducted a numerical taxonomic analysis of leaf phenotypic variation across 73 Malus germplasm (34 species and 39 cultivars). Through ancestor-inclined distribution characteristic analysis, we investigated phylogenetic relationships at both the genus level and infraspecific ranks within Malus. A total of 21 leaf phenotypic traits were selected from 50 candidate traits based on the following criteria: high diversity, abundance, and evenness (D ≥ 0.50, H ≥ 0.80, and E ≥ 0.60); significant intraspecific uniformity and interspecific distinctness (CV¯ ≤ 10% and CV ≥ 15%). Notably, the selected traits with low intraspecific variability (CV¯ ≤ 10%) exhibit environmental robustness, likely reflecting low phenotypic plasticity of these specific traits under varying conditions. This stability enhances their taxonomic utility. It was found that the highest ancestor-inclined distribution probability reached 90% for 10 traceable cultivars, demonstrating reliable breeding lines. Based on morphological evidence, there was a highly significant correlation between the evolutionary orders of (Sect. Docyniopsis → Sect. Sorbomalus → Sect. Malus) and group/sub-groups (B₁ → B₂ → A). This study demonstrates that phenotypic variation in leaf macrostructures can effectively explore the affinities among Malus germplasm, exhibiting taxonomic significance at the infraspecific level, thereby providing references for variety selection. However, hybrid offspring may exhibit mixed parental characteristics, leading to blurred species boundaries. And convergent evolution may create false homologies, potentially misleading morphology-based taxonomic inferences. The inferred taxonomic relationships present certain limitations that warrant further investigation.