Many plant species can reproduce by both sexual and vegetative means. Clonal diversity and degree of intermingling of clones in the vegetative reproductive mode can influence the mating and fertility of individuals. The aim of the study was to assess the clonal structure and its potential influence on gene flow and generative reproduction efficiency in six endangered Betula humilis populations from the southwestern margin of the species range. Analyses of seven microsatellite loci revealed 86 genets among 522 samples. In general, the phalanx strategy dominated in the populations considered, as 76% of ramets shared the same genotype with their closest neighbour. Nevertheless, substantial clonal and genetic diversities and high contribution of unrelated individuals in all B. humilis stands suggest that panmictic pollination prevails. On the other hand, positive and significant relationships between genetic and geographic distances in the two populations could be a consequence of biparental inbreeding resulting from the pollen and seed flow limitations. The seed germination capacity was very low (2.70%); however, the populations characterised by the lowest and highest values of clonal diversity parameters did not differ significantly in the number of germinated seeds, which indicates that clonality is not responsible for seed production failure.
Juniperus excelsa s. str. (Greek juniper) in Crimea is a relic species on the limits of its range, and represents the Mediterranean flora in the Sub-Mediterranean part of the peninsula. Its origin and history in this area remains unresolved. We aimed to analyze phylogeny and potential demographic expansion of the juniper in the Crimea as well as to study its morphological differentiation. We analyzed plant material from 59 trees inhabiting eight populations. Genetic variation assessments were based on the four non-coding chloroplast DNA (cpDNA) fragments and the nuclear internal transcribed spacer region ITS1-5,8S-ITS2 (ITS). To examine the morphological differentiation, eight measured/counted traits of cones, seeds, and shoots were chosen and eight ratios were calculated. Morphological parameters were compared using ANOVA, Student’s t test, discrimination analysis and Kruskal-Wallis and U Mann-Whitney tests. Two cpDNA fragments were polymorphic and, in total, 10 cpDNA haplotypes were found. Haplotype diversity (Hd) ranged from 0.0 to 0.9. Based on both cpDNA and ITS sequences variation, phylogenetic analyses revealed a close relationship of the Crimean junipers to the individuals from other parts of the species range. In general, our molecular results confirmed the low level of genetic differentiation of J. excelsa individuals inhabiting different parts of the species range, likely resulting from a common ancestry. Only slight morphological differences were found between populations with different geographic location or habitat. The analyzes showed the distinctness of the populations from the southern part of the coast. Some unique morphological and molecular features of southern coastal populations imply that they are remnants of Late Pleistocene abundant forests. We suggest that the recent fragmentation of the Juniperus populations in the Crimean Peninsula could have arisen during the Atlantic period of the Holocene.
Key message
Lowered pollen and seed quality of B. humilis compared to its widespread tree relatives may threaten its populations at the southwestern edge of the species range.
Abstract
Inability to produce vital propagules is the main threat to the persistence of rare plants at the edges of species distribution. In the present investigation, factors responsible for the low germinability of shrub birch (Betula humilis) seeds in the peripheral populations were analysed. As shrub birch populations are found in phosphorus (P)-limited peatlands with high groundwater levels, the impact of groundwater PO43− ion contents and groundwater table for B. humilis flowering was also studied. Qualities of the B. humilis seeds and pollen were compared to those of tree congeners silver birch (B. pendula) and downy birch (B. pubescens), which usually represent a high reproductive performance. We revealed that the poor germination ability of B. humilis seeds was caused by the high proportions of empty seeds (44.8%) and seeds with partially developed embryos (47.3%). The insufficient numbers of viable seeds may be a consequence of low-quality shrub birch pollen, as only 1.6% of pollen grains sprouted. The germinability of shrub birch pollen was significantly lower than that of Betula tree pollen; thus, there is a risk of pollination of B. humilis female flowers by pollen grains of tree birches. Positive relationships between the P concentrations in seeds and seed germinability were revealed in tree birches, but not in B. humilis. The availability of PO43− ions in the groundwater did not affect B. humilis flowering, but male inflorescences were more numerous in stands with higher groundwater levels. Consequently, the maintenance of peatlands is crucial for the efficient sexual reproduction of shrub birches.
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