Selection and Floral Evolution in Platanthera bifolia and P. chlorantha (Orchidaceae)

Natural selection mediated by pollinators has influenced the evolution of floral diversity of the flowering plants (angiosperms). The scope of this thesis was to study: 1) phenotypic selection, 2) mating systems, and 3) floral shifts involved in plant speciation. Model plant species were Platanthera bifolia and P. chlorantha (Orchidaceae). These orchids are moth-pollinated, strictly co-sexual (bisexual flowers), and produce a spike that displays 10-20 white flowers. I explored the influence of characters on plant fitness by using multiple linear regressions. Pollen removal (male fitness) and fruit set (female fitness) increased with more flowers per plant in three P. bifolia populations. There was selection towards longer spurs in a dry year when average spur length was shorter than in normal-wet years. Female function was sensitive to drought, which enabled an application of the male function hypothesis of floral evolution (Bateman's principle). The results show that selection may vary between populations, years, and sex-functions. I examined inbreeding by estimating levels of geitonogamy (self-pollination between flowers of an individual) with an emasculation method in two P. bifolia populations. Geitonogamy did not vary with inflorescence size. Levels of geitonogamy was 20-40% in the smaller, but non-significant in the larger population. This may relate to lower number of possible mates and pollinator activity in the smaller population. Platanthera bifolia exhibits the ancestral character state of tongue-attachment of pollinia on the pollinator. Its close relative P. chlorantha attaches its pollinia onto the pollinator's eyes. To explore the mechanism of a floral shift, pollination efficiency and speed was compared between the two species. The results showed no differences in pollination efficiency, but P. chlorantha had faster pollen export and import. Efficiency of pollination in terms of speed may cause floral shifts, and thus speciation.

Reindeer habitat selection under the risk of brown bear predation during calving season

The depredation of semi-domesticated reindeer by large carnivores reflects an important human-wildlife conflict in Fennoscandia. Recent studies have revealed that brown bears (Ursus arctos) may kill substantial numbers of reindeer calves (Rangifer tarandus tarandus) in forest areas in Sweden. Several authors have suggested that predation risk is an important driver of habitat selection in wild Rangifer populations where predation is a limiting factor, but little is known about these mechanisms in semi-domesticated populations. We examined the habitat selection of female reindeer in relation to spatial and temporal variations in brown bear predation risk on the reindeer calving grounds and evaluated the simultaneous responses of brown bears and reindeer to landscape characteristics. We used GPS data from 110 reindeer years (97 individuals) and 29 brown bear years (19 individuals), from two reindeer herding districts in the forest area of northern Sweden. Our results did not indicate that reindeer alter their behavior in response to spatiotemporal variation in brown bear predation risk, on the scale of the calving range. Instead, we suggest that spatiotemporal behavioral adjustments by brown bears were the main driver of prey-predator interactions in our study system. Contrasting responses by brown bears and reindeer to clear-cuts and young forest indicate that forestry can influence species interactions and possibly yield negative consequences for the reindeer herd. Even if clear-cuts may be beneficial in terms of calf survival, logging activity will eventually cause greater abundance of young regenerating forest, reducing available reindeer habitats and increasing habitat preferred by brown bears. Domestication may have made semi-domesticated reindeer in Fennoscandia less adapted to cope with predators. Areal restrictions, limiting the opportunity for dispersion and escape, possibly make the calves more susceptible to predation. Also, a generally higher population density in semi-domesticated herds compared to wild populations can make dispersion a less efficient strategy and the reindeer calves easier prey. Overall, the lack of ability of the reindeer females to reduce brown bear encounter risk on the scale of the calving range is probably an important reason for the high brown bear predation rates on reindeer calves documented in our study areas.