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This clade shows high support for branches and contains 8 species (11 subspecies) with similar chromosomal valence (most derived from x = 10). Close relationships between C. patula (2 n = 20, 40), a species widespread in European woodlands and meadows, and the East-Mediterranean perennial geophyte C. spatulata (2 n = 20) were first revealed by Borsch & al. (2009), within their Campanula rotundifolia-clade. The current increased sampling of Mediterranean species, such as the annual C. lusitanica (2 n = 18, 20), C. phrygia (2 n = 16), and C. sparsa (2 n = 20), and the biennial- perennial C. olympica (2 n = 20), C. pontica (2 n = n/a), and C. rapunculus (2 n = 20), reveals sister relationships between C. lusitanica and the rest of the species, a pattern supported by a more detailed ITS-based phylogenetic study (Cano-Maqueda & al. 2008). Cano-Maqueda & al. (2008) further included five annual, Iberian native species, which formed a well-supported clade including C. lusitanica, and sister to a C. rapunculus–C. sparsa–C. patula lineage. Surprisingly, C. lusitanica was inferred as sister to a C. elatines–C. elatinoides clade by the ITS study of Park & al. (2006), a relationship not supported here. Discrepancies between the respective cp- and nrDNA based signals in this clade would deserve further studies.

Within the C. lusitanica sister clade, ML reconstruction moderately support sister relationships between C. phrygia (2 n = 16) and the rest of the species (2 n = 20), overall suggesting some episodes of descending dysploidy in the lineage. Morphologically, C. phrygia shows some affinities with C. sparsa, both species sharing characteristic ribbed capsule opening by three apical to median pores (Damboldt 1978). Phylogenetic inference also moderately supports affinities between the northern Anatolian species C. pontica and C. olympica. The relationships between C. patula (3 subspp.) and C. spatulata (3 subspp.) remain unresolved. The origin of the Cretan endemic C. spatulata subsp. filicaulis was recently estimated to 17 (±8) Ma for a reduced C. lusitanica–C. spatulata subsp. filicaulis clade [19], The current study would support similar age for the divergence between C. lusitanica and its sister clade (13.10 Ma [4,60–17,55]), but a much younger origin for the C. spatulata–C. filicaulis lineage (stem node 8.60 Ma [1,14–12,90]), overall suggesting a more recent dispersal event in C. spatulata from the mainland to Crete, after the isolation of Crete, such as the very recent split between C. erinus and C. creutzburgii discussed under clade Cam14 below.

From: Mansion & al. (2012: 15)


Borsch T., Korotkova N,, Raus T., Lobin W. & Löhne C. 2009: The petD group II intron as a species level marker: utility for tree inference and species identification in the diverse genus Campanula (Campanulaceae). – Willdenowia 39: 7–33

Cano-Maqueda J., Talavera S., Arista M. & Catalan P. 2008: Speciation and biogeographical history of the Campanula lusitanica complex (Campanulaceae) in the Western Mediterranean region. – Taxon 57:

Damboldt J. 1978: Campanulaceae – Pp. 2–65 in: Davis P.H., (ed.), Flora of Turkey and the East Aegean Islands. – Edinburgh: University Press.

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).

Park J.M., Kovacic S., Liber Z., Eddie W.M.M. & Schneeweiss G.M. 2006: Phylogeny and biogeography of isophyllous species of Campanula (Campanulaceae) in the Mediterranean area. – Syst. Bot. 31: 862–880.