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This well-supported clade nearly entirely encompasses the subgenus Roucela Dumort., a group of 12 small dichotomously branched annual species lacking calyx appendages, and showing disc-like capsules opening by three valves (Carlström 1986). However, the inferred clade does not contain Campanula scutellata, a Balkan native species differing from all the remaining taxa by its large habit size and broad corolla. The placement of C. scutellata into Roucela has been questioned (Carlström 1986), but potential affinities with annuals of the subgenus Megalocalyx (see Cam16) have never been suggested. Other than C. scutellata, most Roucela species are endemic to narrow areas of Greece, the Aegean, and W Turkey, except the widespread, self-compatible C. erinus distributed throughout the Mediterranean Basin, from Macaronesia to Iran.

Clade Cam14 can be further divided into three lineages, with an early diverging Campanula simulans sister to two subclades, a general pattern congruent with a previous study by Roquet (unpublished thesis). Campanula simulans (2 n = 28) has been proposed by Carlström (1986) to describe a Turkish species morphologically and cytologically related to C. drabifolia (2 n = 28) from southern Greece. Nonetheless, molecular data do not support sister relationships between these two species, C. drabifolia belonging to a well-supported subclade otherwise encompassing the Cretan endemic C. creutzburgii and the widespread C. erinus. The timing of diversification for this subclade (0.87 Ma [0.31–2.85]) is congruent with the previous study by Cellinese & al. (2009), who also inferred a recent split of 2.5±2 Ma between C. erinus and C. creutzburgii, suggesting a recent dispersal event from the mainland to Crete during the Pleistocene, after the isolation of Crete.

A second subclade comprises five species with very narrow distributions, namely Campanula delicatula (SE Aegean, SW Turkey), C. rhodensis (endemic to Rhodos), C. pinatzii (endemic to Kasos, Karpathos, and Saria), C. veneris (endemic to Cyprus), and C. podocarpa (Aegean Islands and SW Turkey and Cyprus). The last two species are poorly resolved as sister lineages, C. podocarpa differing from other species of the subclade by its non-stellate calyx, and some particular edaphic affinities (serpentine tolerant). Interestingly, populations from Cyprus have been recently rediscovered (R. Hand, personal communication), and are genetically close to the Turkish accessions included here (G. Mansion, unpublished data). Species delimitation in this group is not easy (Carlström 1986), and some morphs cannot be identified properly (G. Parolly & G. Mansion, pers. obs.), further suggesting reticulate evolution in the group. A more detailed and collaborative study is currently on the way (A. Crowl & al., unpublished data).

From: Mansion & al. (2012: 18)


Carlström A. 1986: A revision of the Campanula drabifolia complex (Campanulaceae). – Willdenowia 15: 375–387

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.

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).