Strong differences in genetic structure across disjunct, edge, and core populations of the distylous forest herb Pulmonaria officinalis (Boraginaceae).

PREMISE OF THE STUDY: Populations at the edge of a species' distribution area are often small and have low levels of gene flow resulting in lower genetic variation and higher differentiation compared to core populations. This study examined genetic variation among populations of the distylous temperate forest herb Pulmonaria officinalis located in the core, the edge, and outside the species' main distribution range. METHODS: We compared patterns of genetic variation for eight microsatellite loci between disjunct (Belgium), edge (western Germany), and core (eastern Germany) populations of P. officinalis. KEY RESULTS: Disjunct populations contained only a subset of alleles found in edge and core populations and had significantly lower within-population genetic variation. No significant differences, however, in within-population genetic variation were found between edge and core populations, except for allelic and genotypic richness. Genetic differentiation was highest among disjunct (F(ST) = 0.11) and lowest among core populations (F(ST) = 0.03). Significant (P < 0.01) isolation by distance was found for disjunct and edge populations (r(M) = 0.29 and 0.50, respectively), but not for core populations (r(M) = 0.18). CONCLUSIONS: The results are best interpreted from a "dynamic range" point of view in which the observed low levels of genetic diversity and high genetic differentiation in disjunct populations are best explained through historical processes, most likely the introduction of the species in medieval times. Lower levels of gene flow caused by the pronounced fragmentation of forests in Belgium may further have contributed to the genetic structure of P. officinalis in these disjunct populations.

Biased morph ratios and skewed mating success contribute to loss of genetic diversity in the distylous Pulmonaria officinalis.

BACKGROUND AND AIMS: In heterostylous plant species, skewed morph ratios are not uncommon and may arise from a range of factors. Despite the recognized importance of skewed morph ratios on overall reproductive success within populations, little is known about the impact of skewed morph ratios on population genetic diversity and differentiation in heterostylous species. This study specifically aimed to clarify the effect of population size and morph bias on population genetic diversity and differentiation in the temperate forest herb Pulmonaria officinalis. This species is characterized by a distylous breeding system and shows morph-specific differences in reproductive success. METHODS: Genetic diversity was determined for 27 P. officinalis populations in northern Belgium by using eight recently developed microsatellite markers. Multiple regressions were used to assess the relationship between genetic diversity, morph bias and population size, and F(ST)-values were calculated for short- and long-styled morphs separately to study genetic differentiation as a function of morph type. KEY RESULTS: For all genetic measures used, morph bias was more important in explaining patterns of genetic diversity than population size, and in all cases patterns of population genetic diversity followed a quadratic function, which showed a symmetrical decrease in genetic diversity with increasing morph bias. However, probably due to the reproductive advantage of L-morphs relative to S-morphs, maximum genetic diversity was found in populations showing an excess of L-morphs (60·7 % L-morph). On the other hand, no significant difference in pairwise genetic distances between populations was observed between L- (0·107) and S-morphs (0·106). CONCLUSIONS: Our results indicate that significant deviations from equal morph ratios not only affect plant reproductive success but also population genetic diversity of heterostylous plant species. Hence, when defining conservation measures for populations of heterostylous plant species, morph ratios should be considered as an important trait affecting their long-term population viability.

Floral display size and spatial distribution of potential mates affect pollen deposition and female reproductive success in distylous Pulmonaria officinalis (Boraginaceae).

In animal-pollinated plants, both the spatial distribution of flowering individuals and the number of flowers that an individual displays affect pollen deposition rates and female reproductive success. Heterostylous species are likely to be particularly sensitive to the contingencies of spatial distribution, as they are reproductively subdivided into distinct mating groups, which usually exhibit self- and intra-morph incompatibility and differ in floral morphology. In this paper, we explore the joint effects of both spatial distribution of potential mates and floral display size on morph-specific pollen deposition rates and seed set patterns in two natural populations of Pulmonaria officinalis, a distylous species with a weak self-incompatibility system. Both total stigmatic pollen load and the proportion of legitimate pollen decreased with increasing spatial isolation. Legitimate (intermorph) pollen transfer was, however, asymmetric and decreased more rapidly with decreasing proximity to a compatible legitimate mating partner in the S-morph than in the L-morph. Total stigmatic pollen loads per flower increased with increasing floral display size, indicating that large plants are disproportionately more visited than smaller individuals. However, because legitimate pollen deposition decreased with increasing floral display size, these results also suggest that larger numbers of flowers increase the degree of geitonogamous pollination. In both the L- and S-morph, seed set significantly decreased with increasing isolation from a legitimate mating partner, but in the L-morph seed set was less dependent on the spatial distribution of the S-morph. In addition, seed set significantly increased with floral display size in the L-morph, but not in the S-morph. These findings indicate that the spatial distribution of potential mates and variation in floral display size may cause morph-specific differences in pollen deposition rates and female reproductive success.