The female advantage in natural populations of gynodioecious Plantago coronopus: seed quantity vs. offspring quality.

In gynodioecious plant species, females can only persist when they have a reproductive advantage in comparison with hermaphrodites. However, several studies have shown that females do not necessarily produce more seeds than hermaphrodites, since seed production can be affected by population characteristics, such as female frequency or population size. The aim of this study was to quantify the female advantage across a large number of natural populations, examine its relationship with population sex ratio and size, and to assess the role of competition on the magnitude of the female advantage. We sampled 27 populations of Plantago coronopus (nuclear-cytoplasmic gynodioecy) along the Belgian and Dutch coast. In each population, we estimated population sex ratio and size, and assessed seed production per flower and seed production per plant. Subsequently, germination, growth, and competition experiments were performed in the greenhouse to determine the female advantage regarding offspring quality. Females produced fewer seeds per plant than hermaphrodites (FA = 0.90), and seed production was negatively related to female frequency. Since both sex morphs were equally affected by pollen availability, the female advantage was not related to population sex ratio. On the other hand, offspring of females showed higher germination and growth rates, resulting in higher competitive abilities when seeds of a female and a hermaphrodite were grown together. Overall, these results indicate that differences in competitive abilities between the offspring of females and hermaphrodites may have contributed to the maintenance of females in relatively high frequencies in populations of this short-lived gynodioecious plant species.

The impact of hybridization on long-term persistence of polyploid Dactylorhiza species.

PREMISE OF THE STUDY: Hybridization may pose severe threats to the long-term survival of the parental taxa through introgression and the formation of hybrid swarms. However, when the resulting hybrids show strong male and female sterility, backcrossing and introgression are unlikely to occur, but the parental species may suffer from reduced male and female fitness. METHODS: We assessed the impact of hybridization on the long-term persistence of two food-deceptive orchids in the genus Dactylorhiza (the common Dactylorhiza maculata and the rare D. sphagnicola). The extent of hybridization was investigated using both molecular markers and morphometric measurements. To determine the strength of postmating reproductive isolation, hand pollinations were conducted between pure and hybrid individuals. Finally, fruit set and seed viability of open-pollinated plants were determined in sympatric and allopatric populations to investigate the impact of hybridization on the reproductive output of the pure parental species. KEY RESULTS: Our results showed that postmating reproductive isolation was weak and that hybridization occurred frequently within the studied sympatric population. Although hybrids were characterized by very low female fitness, mainly because of strongly reduced seed viability, backcrossing appeared to occur and was asymmetric toward the rare D. sphagnicola. Fruit set and seed viability of open-pollinated plants were also significantly lower in the sympatric population than in the allopatric populations, indicating that hybridization and ongoing introgression incurred fitness costs in the pure parental species. CONCLUSIONS: Overall, our results suggest that extensive hybridization can affect the long-term viability of the parental species through the combined effect of introgression following interspecific hybrid fertilization and reduced fitness of the parental species.

Recent range expansion of a terrestrial orchid corresponds with climate-driven variation in its population dynamics.

The population dynamics and distribution limits of plant species are predicted to change as the climate changes. However, it remains unclear to what extent climate variables affect population dynamics, which vital rates are most sensitive to climate change, and whether the same vital rates drive population dynamics in different populations. In this study, we used long-term demographic data from two populations of the terrestrial orchid Himantoglossum hircinum growing at the northern edge of their geographic range to quantify the influence of climate change on demographic vital rates. Integral projection models were constructed to study how climate conditions between 1991 and 2006 affected population dynamics and to assess how projected future climate change will affect the long-term viability of this species. Based on the parameterised vital rate functions and the observed climatic conditions, one of the studied populations had an average population growth rate above 1 (λ = 1.04), while the other was declining at ca. 3 % year(-1) (λ = 0.97). Variation in temperature and precipitation mainly affected population growth through their effect on survival and fecundity. Based on UK Climate Projection 2009 estimates of future climate conditions for three greenhouse gas emission scenarios, population growth rates are expected to increase in one of the studied populations. Overall, our results indicate that the observed changes in climatic conditions appeared to be beneficial to the long-term survival of the species in the UK and suggest that they may have been the driving force behind the current range expansion of H. hircinum in England.

The effect of phenological variation in sex expression on female reproductive success in Saxifraga granulata.

PREMISE OF THE STUDY: Differences in timing of flowering within and among protandrous plants shift the floral sex ratio from male to female dominance during the flowering season. Hence, the number of seeds produced by a single flower depends on traits of the flower itself (e.g., allocation to male and female function, position within an inflorescence, and flower size), as well as plant traits (e.g., timing of flowering, number of flowers, and plant height). Although it is clear that characteristics of individual flowers and whole plants can affect the number of seeds produced per flower, their relative importance for plant fitness remains largely unknown. METHODS: We examined how phenological sex expression affected seed number per flower in two populations of the protandrous grassland herb Saxifraga granulata. Seed number was assessed for >1200 flowers and related to their position within an inflorescence, male and female phase duration, timing of flowering, number of flowers per plant, and plant height. KEY RESULTS: Seed number within and among plants decreased significantly over time. Early lateral flowers were larger and produced more seeds in comparison to late lateral flowers, indicating that flower position significantly affected seed number through its combined effect on sex allocation, timing of flowering, and attractiveness. CONCLUSIONS: Our results showed that female reproductive success of a single flower was best explained by its position within an inflorescence and that plant traits such as first flowering date and number of flowers per plant had a smaller impact on seed number per flower.

Genetic Diversity and Spatial Genetic Structure of the Grassland Perennial Saxifraga granulata along Two River Systems.

Due to changes in land use, the natural habitats of an increasing number of plant species have become more and more fragmented. In landscapes that consist of patches of suitable habitat, the frequency and extent of long-distance seed dispersal can be expected to be an important factor determining local genetic diversity and regional population structure of the remaining populations. In plant species that are restricted to riparian habitats, rivers can be expected to have a strong impact on the dynamics and spatial genetic structure of populations as they may enable long-distance seed dispersal and thus maintain gene flow between fragmented populations. In this study, we used polymorphic microsatellite markers to investigate the genetic diversity and the spatial genetic structure of 28 populations of Saxifraga granulata along two rivers in central Belgium. We hypothesized that rivers might be essential for gene flow among increasingly isolated populations of this species. Genetic diversity was high (HS = 0.68), which to a certain extent can be explained by the octoploid nature of S. granulata in the study area. Populations along the Dijle and Demer rivers were also highly differentiated (G"ST = 0.269 and 0.164 and DEST = 0.190 and 0.124, respectively) and showed significant isolation-by-distance, indicating moderate levels of gene flow primarily between populations that are geographically close to each other. Along the river Demer population genetic diversity was higher upstream than downstream, suggesting that seed dispersal via the water was not the primary mode of dispersal. Overall, these results indicate that despite increasing fragmentation populations along both rivers were highly genetically diverse. The high ploidy level and longevity of S. granulata have most likely buffered negative effects of fragmentation on genetic diversity and the spatial genetic structure of populations in riparian grasslands.

Microsatellite primers for the gynodioecious grassland perennial Saxifraga granulata (Saxifragaceae).

PREMISE OF THE STUDY: Nine polymorphic and 12 monomorphic microsatellite loci (simple sequence repeats [SSRs]) were isolated and characterized for the gynodioecious grassland perennial Saxifraga granulata. • METHODS AND RESULTS: Based on genomic screening of leaf material of four individuals from four populations, a total of 21 microsatellite primer pairs were designed for S. granulata. Nine loci were polymorphic and were optimized into two PCR multiplex reactions and tested on 100 individuals from five riparian populations from central Belgium. The number of alleles of the polymorphic loci ranged from three to 18, and gametic heterozygosity ranged from 0.26 to 0.94. • CONCLUSIONS: The markers that are presented here are the first microsatellite markers reported for S. granulata and will be used to assess how river systems shape the spatial genetic structure and diversity of riparian populations of this species.