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This well-supported monophyletic group contains five “isophyllous” species of Campanula, namely C. garganica, C. elatines, C. fenestrellata, C. portenschlagiana, and C. poscharskyana. The Isophylla group is morphologically (isophylly, both the basal and cauline leaves having cordate to ovate blade; erect capsules opening with basal pores) and karyologically (2 n = 34) well defined, and encompasses some 12 species disjunctly distributed in the sub-Mediterranean Adriatic Mountains (Park & al. 2006, Damboldt 1965, Greuter & al. 1984). Isophylla has been further divided into three morphological groups (Lovašen-Eberhardt & Trinajstic 1978), and corresponding three well-supported, albeit non-sister ITS clades (Park & al. 2006). Our study also inferred the polyphyly of the isophyllous assemblage with Cam08 corresponding to the tentative “garganica” clade of Parks & al. (2006), their “fragilis” and part of the “elatines” clades being imbedded in our Cam12 lineage (see below).

Despite great similarities between the respective petD (this study) and ITS (Park & al. 2006) inference, some taxa show strongly incongruent topological position. Indeed, our current petD analysis does not support the sister relationships between C. elatines and C. elatinoides, the former being sister to C. fenestrellata and the latter included in clade Cam12, a result congruent with Borsch & al. (2009). The "elatines” group, treated under “garganica” by Damboldt (1965), was described to encompass two narrowly-distributed alpine species (C. elatines and C. elatinoides), characterized by intermediate morphological characters between the “fragilis” and “garganica” clades (Park & al. 2006). Interestingly, isozyme evidence (Frizzi & Tammaro 1991) support closer relationships between C. elatinoides and C. isophylla (fragilis clade), a result in line with our current inference (C. elatinoides and C. isophylla in clade Cam12). Furthermore, some ecological differences, including the strong affinity of C. elatines (Piemont) for gneiss or granite versus calcareous rocks for C. elatinoides (Insubrian Alps), would add further support for their phylogenetic divergence (Park & al. 2006).

On the whole, Cam08, as currently circumscribed, is a genetically well-supported clade with strong morphological, karyological, and geographical structure. Indeed, most species are similar in habit and floral shape, share a diploid to hexaploid chromosome number based on x = 17, and mainly occur in the Transadriatic Mediterranean area.

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

Damboldt J. 1965: Zytotaxonomische Revision der isophyllen Campanulae in Europa. – Bot. Jahrb. Syst.. 84: 302–358.

Frizzi G., Tammaro F. 1991: Electrophoretic study and genetic affinity in the Campanula elatines and C. fragilis (Campanulaceae) rock-plants group from Italy and W Jugoslavia. – Plant Syst. Evol. 174: 67–73.

Greuter W., Burdet H.M. & Long G. 1984: Campanula L. – Pp. 123–145 in: Med-Checklist. – Genève: Conservatoire et Jardin botaniques.

Lovašen-Eberhardt Z. & Trinajstić I. 1978: O geografskoj distribuciji morfoloških karakteristika vrsta serije Garganicae roda Campanula L. u flori Jugoslavije - On geographic distribution of morphological characteristics of Campanula L. species of Garganicae series in Yugoslavian flora (in Croatian). – Biosistematika 4: 273–280.

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.