Paleoendemic species of the monophyletic genus Ramonda (R. myconi, R. serbica and R.~nathaliae) are the remnants of the Tertiary tropical and subtropical flora in Europe. They are the rare resurrection plants of Northern Hemisphere temperate zone. Ramonda serbica and R. nathaliae are chorologically differentiated in the Balkan Peninsula and occupy similar habitats in calcareous, northward slopes in canyons and mountainsides. They remain well-hydrated during spring, late autumn and even in winter. In summer and early autumn when plants are subjected to drought and thermal stress, their desiccation tolerance comes into operation and they fall into anabiosis. Investigations revealed the permanent presence of ubiquitine and its conjugates, high amounts of oxalic acid and proline. Both species are homoiochlorophyllous. It enables them to rapidly resume photosynthesis upon rehydration, but also makes them susceptible to reactive oxygen species formation. Dehydration induces activation of antioxidative enzymes (ascorbate peroxidase, glutathione reductase, polyphenol oxidase), increase in amounts of AsA and GSH, phenolic acids, dehydrins, sucrose, and inorganic ions. Plasma membranes, characterized by high amount of cholesterol, are subjected to decrease in membrane fluidity mostly on account of increased level of lipid saturation. Cytogenetic analysis revealed that R. nathaliae is a diploid (2n = 48) and probably evolutionary older species, while R. serbica is a hexaploid (2n = 144). Two species live together in only two localities forming hybrid individuals (2n = 96). Polyploidization is the major evolutionary mechanism in the genus Ramonda that together with hybridization ability indicates that these relict species which have preserved an ancient survival strategy are not the evolutionary “dead end.”The species of the genus Ramonda are promising sources of data important for understanding the complex strategy of resurrection plants’ survival, appraised through a prism of their evolutionary and adaptive potential for multiple environmental stresses.
The pollen and seeds of 3 paleoendemic resurrection species, Ramonda myconi (L.) Rchb. (Iberian Peninsula), R. nathaliae Pančić & Petrović, and R. serbica Pančić (Balkan Peninsula), as well as of natural hybrids between the 2 last species, have been analyzed using light and scanning electron microscopy. Their general structural differences, taxonomic and phylogenetic significance, pollen viability, and seed germination capacity, as well as the correlation of pollen and seed characteristics and plant ploidy levels, have been studied. The pollen grains are small (R. myconi, R. nathaliae) to medium (R. serbica) in size, ranging from 10 to 28 µm, and 3-colporate, isopolar monads with microreticulate-perforate exine. Seeds are small, 309 to 1000 µm long and 80 to 425 µm wide, elongated, with a reticulate surface and auriculate ornamentation. Micromorphologies of the pollen exine ornamentation and seed surface revealed significant differences among the 3 species. Pollen from hybrid individuals was heterogeneous in size and morphology, and the germination of their tiny seeds was very low. A strong correlation was found between pollen size, DNA content, and chromosome number. The environmental influence on pollen and seeds of 3 species and especially of the R. nathaliae population growing on serpentine is also discussed.
Analyses of mitotic chromosome numbers and nuclear DNA content were performed for 39 populations of 17 perennial Cerastium taxa from south-eastern Europe. The DNA content ranged from 2C = 2.43 to 8.78 pg, revealing four ploidy levels corresponding to 4x (2n = 36), 8x, 12x and 16x. High-polyploid cytotypes with a greater range of ploidy (up to 2n = 144) occur mostly in the central mountainous parts of the Balkan Peninsula. The chromosome number was determined for the first time for C. dinaricum (2n = 36 + 1B), C. decalvans subsp. orbelicum (2n = 36), C. decalvans subsp. glutinosum (2n = 36), C. neoscardicum (2n = 144), C. malyi subsp. serpentini (2n = 144) and C. moesiacum s.s. (2n = 144). New chromosome counts were recorded for C. arvense subsp. strictum (2n = 108), C. banaticum subsp. kosaninii (2n = 36) and C. grandiflorum (2n = 36). For the first time, flow cytometry was used to estimate C values for six species (15 taxonomic entities). The intraspecific variation quotient of C values is high, ranging from 1.003 in C. malyi to 1.306 in C. decalvans subsp. decalvans. The variation in chromosome size among both tetra-and octoploid members of Cerastium is much more prominent than in most other angiosperm polyploid series. Significant genome downsizing after polyploidization was observed in some investigated taxa. Differences in ploidy levels and monoploid genome size values confirm the taxonomic status of C. decalvans subsp. glutinosum and C. decalvans subsp. leontopodium. The results obtained indicate a possible close relationship between C. banaticum and C. grandiflorum, but not C. arvense.
Polyploidization has played an important role in the diversification of the genus Sesleria (Poaceae), which comprises c. 48 species and subspecies mostly distributed in Europe. The genus' centre of diversity clearly is the Balkan Peninsula, harbouring about 80% of the species, half of which are endemic to this area. We employed chromosome counts, measurements of absolute genome size and determination of relative DNA-content for 460 populations belonging to 43 species of Sesleria. Our main aim was to provide essential baseline data for future molecular genetic reconstructions of the genus' evolutionary history. Relative genome size allowed for a mostly clear separation of four ploidy levels. The most frequent and widespread cytotypes are tetraploids followed by octoploids, while di-and dodecaploids were only found in a few species. We present first chromosome numbers for the tetraploid species S. doerfleri, S. phleoides, S. skipetarum and S. tuzsonii as well as for diploid S. ovata. Based on relative and partly also on absolute genome size measurements, ploidy level was determined in tetraploid S. rhodopaea and S. voronovii for the first time, and new cytotypes were identified in S. interrupta, S. kalnikensis and S. wettsteinii (tetraploids), S. caerulea, S. klasterskyi, S. latifolia, S. tenerrima, S. ujhelyii and S. vaginalis (octoploids), and S. albanica and S. vaginalis (dodecaploids). While most Sesleria species are ploidy-uniform, several comprise two or even, in the case of S. vaginalis, three ploidy levels. Genome downsizing after polyploidization was confirmed by significant negative correlation between ploidy level and monoploid genome size. Finally, we found a significant increase in monoploid relative genome size towards the margin of the genus' distribution area, which may be triggered by increased activity of transposable element in populations exposed to environmental or genomic stress.
Reconstruction of relationships among populations of the morphologically polymorphic and taxonomically intricate Sesleria rigida sensu Fl. Eur. based on Amplified Fragment Length Polymorphisms (AFLPs) revealed four clearly differentiated genetic groups that did only partly follow recent taxonomic concepts, but were strictly allopatric. While some of the previously described taxa constitute distinct genetic entities, others have no taxonomic value. Synthesizing our AFLP data with ploidy‐level information obtained from all genetically investigated individuals as well as with chromosome counts revealed that tetraploid individuals prevail, while octoploids occur only within S. filifolia. Lack of AFLP divergence between tetra‐ and octoploids suggests an autopolyploid origin of the latter. The genetic differentiation pattern was reflected by morphological differentiation, allowing for a taxonomic revision of the constituents of S. rigida sensu Fl. Eur. resulting in recognition of the four species S. achtarovii, S. filifolia, S. rigida, and S. serbica.
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