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This strongly supported monophylum, exemplified here for the first time, is early diverging and sister to the respective Cam08–Cam12 assemblages. Campanula aizoides, C. aizoon, and C. columnaris are three narrow-distributed, Greek endemic species, morphologically similar and caracterized by their robust taproot, dense rosette of leaves, from which arises a thyrsoid inflorescence with large, tubular-campanulate flowers (Contandriopolous & al. 1973). Campanula aizoides presents a striking bi-regional and disjunct distribution in western Crete (Lefka Ori) and northern Peloponnese (Mt Chelmos), whereas C. aizoon (Mts Parnassos and Giona) and C. columnaris (Mt Vardhousia) are found in some places of the mountain ranges of Central Greece (Sterea Ellas). The divergence age estimate at the lineage stem node is 30,86 Ma [18,58–35,81]), indicating an ancient separation of this Greek lineage from the Cam08–Cam12 sister clade. Interestingly, the whole lineage seem to have diversified very recently (c. 1.5 Ma), forming two mainland lineages and an insular one, contradicting a paleo-subendemic status postulated for the Cretan C. aizoides (Cellinese & al. 2009). Alternatively, the three species could represent a single entity of an older lineage whose remnant populations in both mainland Greece and Crete may have escaped from extinction by taking refuge in and/or adapting to mountain habitats. Overall, the low genetic distances estimated for the respective taxa, the identical chromosome numbers (n = 8), weak morphological differences, and different ecological preferences (Contandriopolous & al. 1973) would better favor the second hypothesis.

From: Mansion & al. (2012: 15)


Cellinese N., Smith S.A., Edwards E.J, Kim S.T., Haberle R.C., & al. 2009: Historical biogeography of the endemic Campanulaceae of Crete. – J. Biogeogr. 36: 1253–1269

Contandriopolous J., Quezel P. & Zaffran J. 1973: A propos des Campanules du groupe aizoon en Grèce méridionale et en Crête. – B. Soc. Bot. Fr-Lett. 120: 331–340

Mansion G., Parolly G., Crowl A.A., Mavrodiev E., Cellinese N., Oanesian M., Fraunhofer K., Kamari G., Phitos D., Haberle R., Akaydin G., Ikinci N., Raus T. & Borsch T. 2012: How to Handle Speciose Clades? Mass Taxon-Sampling as a Strategy towards Illuminating the Natural History of Campanula (Campanuloideae). – PLoS ONE 7 (11).