Abstract
Nutlet micromorphological traits represent highly stable genetic markers with significant diagnostic value for resolving taxonomic complexities in phenotypically plastic Lamiaceae genera. This study provides a comprehensive macro- and ultrastructural evaluation of mature nutlets across seven localized Stachys taxa (S. mardinensis, S. baytopiorum, S. megalodonta subsp. mardinensis, S. menthoides, S. burgsdorffioides subsp. burgsdorffioides, S. melampyroides, and S. cretica subsp. garana) from South-Eastern Anatolia. The evaluated group includes endemic taxa (S. menthoides, S. baytopiorum, and S. megalodonta subsp. mardinensis) and the rare species S. mardinensis. Biometric measurements revealed notable morphometric extensions; most prominently in S. cretica subsp. garana, whose nutlet dimensions (mean: 3.20 × 2.27 mm) significantly exceed current literature limits. Scanning electron microscopy (SEM) identified five distinct exine sculpturing patterns (reticulate, reticulate-foveolate, areolate, tuberculate, and reticulate-ruminate) that provide unequivocal species-level diagnostic criteria. Multivariate statistical analysis integrated via Principal Component Analysis (PCA) accounted for 73.93% of the total cumulative variance (PC1: 49.57%, PC2: 24.36%), effectively segregating taxa based on dimensional traits and surface ornamentation. Hierarchical UPGMA clustering based on Gower’s distance delimited the taxa into distinct lineages (~0.13 to 0.66 distance thresholds), strongly supporting their taxonomic and sectional boundaries. Based on these stable carpological attributes, a diagnostic key is provided to facilitate rapid identification. These findings expand the current carpological circumscription of Stachys within the flora of Türkiye.
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