Taxonomic Reassessment and Rediscovery of Tulipa scardica Bornm. in Greece: Insights from Integrated Analyses Compared to T. undulatifolia Boiss.

Tulipa scardica (Balkan endemic) was last recorded in Greece in 1896, possibly attributed to longstanding taxonomic ambiguity, as it has frequently been considered as conspecific with T. gesneriana or T. undulatifolia. In the present study we aimed to investigate the historical Greek locality of T. scardica and to reassess its taxonomic status in comparison with the closely related T. undulatifolia (also native to Greece and member of T. scardica complex). Targeted field surveys were conducted to verify the presence of T. scardica in Greece. The newly identified tulip population was subjected to an integrated analytic approach, including qualitative and quantitative morphological assessment, seed micromorphology, DNA barcoding, karyological investigation, and habitat/soil properties analyses. These datasets were comparatively evaluated against four reference populations of T. undulatifolia. Molecular data did not provide consistent species-level resolution, whereas morphological and karyological evidence statistically supported their distinction. Mitotic metaphase chromosomes of T. scardica were documented herein for the first time, while first-time scanning electron microscopy (SEM) analysis revealed the presence of different types of stomatal complexes in seed coats of both taxa. In addition, soil parameters differed between the examined taxa, and those of the rediscovered population were consistent with habitat preferences of T. scardica. Although both taxa exhibited considerable variability, the combined evidence derived from the present study strongly supported the rediscovery of T. scardica in Greece after approximately 130 years, unless proven otherwise in a wider context across its Balkan range.