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This clade shows weak sister relationships between a lineage of two perennial species (Campanula
rumeliana and C. jacquinii), and an assemblage containing both annuals (11) and perennials (3). The strong affinitiy between C. rumeliana and C. jacquinii has already been suggested (Tan & Iatrou 2001), but the absence of genetic relationships with the otherwise morphologically similar species (e.g. C. asperuloides, C. buseri, or C. myrtifolia) here included in Cam13, refutes their taxonomic inclusion in either Diosphaera or Tracheliopsis.

The second lineage shows further affinities between annual species of the respective subgenera Sicyocodon (C. macrostyla), Megalocalyx (C. propinqua, C. strigosa, C. hierosolymitana, C. camptoclada, C. cecilii, and C. reuteriana), Roucela (C. scutellata), and the perennials C. damascena, C. mardinensis, and C. lourica. Although most species of the subgenus Megalocalyx are very polymorphic and difficult to separate morphologically (Sáez & Aldasoro 2003), they appear to have evolved in two lineages that originated in the early Miocene (24,67 Ma (Boissier, 1875, Turner & al. 2009, Eddie & Ingrouille 1999,Roquet & al. 2008, Federov 1957, Damboldt 1976, Oganesian 1995, Quézel 1953, Borsch & al. 2009, Cellinese & al. 2009, Eddie & al 2003, Haberle & al. 2009, Park & al. 2006, Heath & al. 2008, Hillis & al. 2003, Pollock & al. 2002, Hedtke & al. 2006, Nabhan & Sarkar 2012, Muller & al. 2006, Drummond 2008). On the one hand, most species of Megalocalyx are sister to C. macrostyla, a singular species with a combination of characters not found in any other extant species of Campanula, subsequently classified in the monotypic subgenus Sicyocodon (Damboldt 1976, Sáez & Aldasoro 2003). Albeit partially unresolved, this clade depicts relationships between annuals currently occurring in the Near-East region, from Turkey to Egypt. On the other hand, an annual C. scutellata–C. stellaris lineage is sister to the Iranian perennial C. lourica. Both C. scutellata and C. stellaris differ by the presence (C. scutellata) vs. absence (C. stellaris) of calyx appendages, but exhibit particular stellate and accrescent calyces after fructification. Campanula scutellata has long been considered a particular species within subgenus Roucela, and must be clearly excluded from it. As mentioned for the annual species-rich clade Cam14, the possibility of reticulate evolution exists in the current clade, whose natural history inference would necessitate increasing taxonomic and geographic sampling, and more sensitive molecular markers.

From: Mansion & al. (2012: 19)


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