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Evolution and biogeography of heterostyly in Linum (Linaceae)

José Ruiz Martín, University of Seville. Supervisors: Rocío Pérez-Barrales & Juan Arroyo. December 2017.
Summary:
The genus Linum L. includes about 250 species with sub-cosmopolitan distribution. Linum species are highly diverse in life forms, floral colours, pollination systems and arrangement of sex organs. Particularly interesting are this last aspect, and Darwin (The Different Forms of Flowers, 1877) already noticed this variation and used it to formulate his ideas about the mechanisms that triggered heterostyly. Linum species include monomorphism, approach herkogamy, style dimorphism, distyly, and 3-D distyly, a putatively derived form of style polymorphism (Armbruster et al. 2006. New Phytol 171: 581–590). To test if evolutionary transitions between stylar conditions agree with existing hypotheses, it is crucial to have a solid phylogenetic hypothesis for species and populations. Results of previous research (McDill et al., 2009 Syst. Bot. 34: 386–405) concluded that heterostyly in Linum seems a condition mostly restricted to the Mediterranean Basin, but there have been reported cases of heterostyly in other regions, such as the South African Cape Floristic Region, under similar Mediterranean climate conditions.

Below are outlined the objectives of my thesis, which aim at testing Darwin´s ideas on Linum species at different levels:
1. (i) To resolve the phylogenetic and phylogeographic relationships among Mediterranean and South African species and populations ;(ii) to characterize the style polymorphism of each species by measuring 30 flowers of each species per morph and thus determining the level of herkogamy present in each population and species; (iii) to reconstruct the ancestral character state of the flower polymorphism, and finally (iv) to infer evolutionary transitions between the states of this character in the phylogeny.

2. In a further step, I wish to examine the transitions of style conditions among populations of the Linum tenuifolium aggr. in the Mediterranean Basin. This group is represented by two distinct species: L. tenuifolium L. (stylar monomorphic, herkogamous species) and L. suffruticosum L. (distylous species, with a special 3-D reciprocal herkogamy, supposedly derived from the former), although the latter harbours much variation. All populations collected (125 pops: L. tenuifolium (39 pops) & L. suffruticosum (86 pops)) will be included in phylogenetic and phylogeographic analysis to disentangle the taxonomic status and evolutionary relationships of the group. Once the phylogenetic results are obtained, a chronology of the evolutionary events in resolved clades will be performed using internal and external calibration points.

3. Reproductive biology and stylar polymorphism in Linum tenuifolium aggr. in natural conditions. This study aims at determining the breeding systems of the species, including the possible heteromorphic incompatibility system of stylar morphs. This study is being undertaken in three different populations of the Iberian Peninsula (Málaga, Guadalajara and Pyrenees).

4. After the preliminary field work around Mediterranean Basin some collateral works are emerging. The recognition of a new heterostylous species from Morocco, and the description of a new polymorphism in the genus in a species from the Eastern Mediterranean, will be noteworthy results. These results illustrate the value of conducting research in biodiversity hotspots such as the Mediterranean Basin, as usually it reveals new processes beyond taxonomical patterns.