Gender expression of individual cushions of Silene acaulis was shown to vary between years. Fifteen of thirty-nine (= 38%) cushions monitored changed gender expression between 1993 and 1995. Cushions dominated by female flowers were shown to be more stable in gender expression than cushions dominated by male or hermaphrodite flowers.
Pollen viability among genders and limitation of female seed production in a natural trioecious population of the circumpolar cushion plant Silene acaulis was examined. Pollen viability was estimated by an in vitro pollen germination experiment. Both male and hermaphrodite flowers displayed large variation in pollen viability (0-53% in hermaphrodite and 0-54% in male flowers). There was a significant difference between genders in pollen viability: male plants had on average higher pollen viability than hermaphrodite plants. Resource and pollen limitation of seed production was studied by an experiment consisting of three treatments; (I) hand-pollination and removal of all other flowers on the cushion, (II) harid-pollination without removal of other flowers, and (III) open pollination without removal of flowers, Hand-pollination increased seed production, whereas removal of flowers had no effect on seed production. Abortion of pollinated ovules during seed development and seed mass did not differ among treatments. To control for effect of fruit number on seed production, data from naturally pollinated individuals was used. There was a positive correlation between both total number of seeds and fruit number, mean seed number per fruit and fruit number, respectively. These results indicate that seed production of S. acaulis is mainly limited by pollen availability whereas resource competition between fruits is not important as a limiting factor. The possible role of male quality differences between genders and pollen limitation of seed production for maintenance of trioecious reproductive systems is discussed.
The aim of this study was to test if early no overing species respond with increased seed production to climate warming as is predicted for late-flowering seed-risk strategists. Experimental climate warming of about 3 degrees C was applied to two populations of the cushion-forming plant Silene acaulis (L.) Jacq. The experiment was run at one subarctic site and one alpine site for 2 years and 1 year, respectively, using open-top chambers (OTC). The 2-year temperature enhancement at the subarctic site had a marked effect on the flowering phenology. Cushions inside the OTC started flowering substantially earlier than control cushions. Both the male and female phases developed faster in the OTCs, and maturation of capsules occurred earlier. The cushions also responded positively in reproductive terms and produced more mature seeds and had a higher seed/ovule ratio. After 1 year temperature enhancement at the alpine site there was a weak trend for earlier flowering, but there was no significant difference in seed production or seed/ovule ratio.
Predicted increase of female frequencies in gynodioecious Silene acaulis was tested along an altitude gradient in northern Sweden. Average female frequencies for the four sites increased with altitude from 42% to 59% within a short geographical distance. This follows the outcrossing hypothesis, that female frequencies should be positively correlated with selfing rates of hermaphrodites in populations. More adverse environmental conditions should favour gynodioecy in areas where reproduction to a greater part relies on vegetative reproduction or selfing. Further, a significant difference in corolla width was found between females and the larger hermaphrodites, but not between sites. Cushion size and the number of flowers per cushion decreased with altitude.
The International Tundra Experiment (ITEX) is a collaborative, multisite experiment using a common temperature manipulation to examine variability in species response across climatic and geographic gradients of tundra ecosystems. ITEX was designed specifically to examine variability in arctic and alpine species response to increased temperature. We compiled from one to four years of experimental data from 13 different ITEX sites and used meta-analysis to analyze responses of plant phenology, growth, and reproduction to experimental warming. Results indicate that key phenological events such as leaf bud burst and flowering occurred earlier in warmed plots throughout the study period; however, there was little impact on growth cessation at the end of the season. Quantitative measures of vegetative growth were greatest in warmed plots in the early years of the experiment, whereas reproductive effort and success increased in later years. A shift away from vegetative growth and toward reproductive effort and success in the fourth treatment year suggests a shift from the initial response to a secondary response. The change in vegetative response may be due to depletion of stored plant reserves, whereas the lag in reproductive response may be due to the formation of flower buds one to several seasons prior to flowering. Both vegetative and reproductive responses varied among life-forms; herbaceous forms had stronger and more consistent vegetative growth responses than did woody forms. The greater responsiveness of the herbaceous forms may be attributed to their more flexible morphology and to their relatively greater proportion of stored plant reserves. Finally, warmer, low arctic sites produced the strongest growth responses, but colder sites produced a greater reproductive response. Greater resource investment in vegetative growth may be a conservative strategy in the Low Arctic, where there is more competition for light, nutrients, or water, and there may be little opportunity for successful germination or seedling development. In contrast, in the High Arctic, heavy investment in producing seed under a higher temperature scenario may provide an opportunity for species to colonize patches of unvegetated ground. The observed differential response to warming suggests that the primary forces driving the response vary across climatic zones, functional groups, and through time.
1 Macrolichens are important for the functioning and biodiversity of cold northern ecosystems and their reindeer-based cultures and economics. 2 We hypothesized that, in climatically milder parts of the Arctic, where ecosystems have relatively dense plant canopies, climate warming and/or increased nutrient availability leads to decline in macrolichen abundance as a function of increased abundance of vascular plants. In more open high-arctic or arctic-alpine plant communities such a relationship should be absent. To test this, we synthesized cross-continental arctic vegetation data from ecosystem manipulation experiments simulating mostly warming and increased nutrient availability, and compared these with similar data from natural environmental gradients. 3 Regressions between abundance or biomass of macrolichens and vascular plants were consistently negative across the subarctic and mid-arctic experimental studies. Such a pattern did not emerge in the coldest high-arctic or arctic-alpine sites. The slopes of the negative regressions increased across 10 sites as the climate became milder (as indicated by a simple climatic index) or the vegetation denser (greater site above-ground biomass). 4 Seven natural vegetation gradients in the lower-altitude sub- and mid-arctic zone confirmed the patterns seen in the experimental studies, showing consistent negative relationships between abundance of macrolichens and vascular plants. 5 We conclude that the data supported the hypothesis. Macrolichens in climatically milder arctic ecosystems may decline if and where global changes cause vascular plants to increase in abundance. 6 However, a refining of our findings is needed, for instance by integrating other abiotic and biotic effects such as reindeer grazing feedback on the balance between vascular plants and lichens.
Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.
Climate change was simulated by increasing temperature and nutrient availability in an alpine landscape. We conducted a field experiment of BACI-design (before/after control/impact) running for five seasons in two alpine communities (heath and meadow) with the factors temperature (increase of ca. 1.5-3.0A degrees C) and nutrients (5 g N, 5 g P per m(2)) in a fully factorial design in northern Swedish Lapland. The response variables were abundances of plant species and functional types. Plant community responses to the experimental perturbations were investigated, and the responses of plant functional types were examined in comparison to responses at the species level. Nutrient addition, exclusively and in combination with enhanced temperature increase, exerted the most pronounced responses at the species-specific and community levels. The main responses to nutrient addition were increases in graminoids and forbs, whereas deciduous shrubs, evergreen shrubs, bryophytes, and lichens decreased. The two plant communities of heath or meadow showed different vegetation responses to the environmental treatments despite the fact that both communities were located on the same subarctic-alpine site. Furthermore, we showed that the abundance of forbs increased in response to the combined treatment of temperature and nutrient addition in the meadow plant community. Within a single-plant functional type, most species responded similarly to the enhanced treatments although there were exceptions, particularly in the moss and lichen functional types. Plant community structure showed BACI responses in that vegetation dominance relationships in the existing plant functional types changed to varying degrees in all plots, including control plots. Betula nana and lichens increased in the temperature-increased enhancements and in control plots in the heath plant community during the treatment period. The increases in control plots were probably a response to the observed warming during the treatment period in the region.
Effects of simulated environmental change on bryophyte and lichen species richness and diversity in alpine tundra were investigated in a 5-year experiment at Latnjajaure, northern Sweden. The experiment had a factorial design including fertilisation and temperature enhancement in one meadow and one heath plant community. Responses in species richness, biodiversity, and species composition of bryophytes and lichens to experimental treatments were compared to the observed variation in six naturally occurring plant communities. The combination of fertilisation and enhanced temperature resulted in a species impoverishment, for bryophytes in the bryophyte-dominated community, and for lichens in the lichen-dominated communities, but the species composition stayed within the observed natural variation. During the course of the study, no species new to the investigated mid-alpine landscape were recorded, but that scenario is realistic within a decade when comparing with the processes seen in vascular plants.
The predicted changes in climate over the next 50 years are expected to be most pronounced in arctic and subarctic regions. In the present study, we examine the responses of a subarctic-alpine rich meadow and poor heath community to factorial manipulations of temperature and nutrient treatments. Specifically, we address response to the treatments in terms of biodiversity and relative cover of the bryophyte, lichen and vascular plant communities. We point out that the responses differ among mosses, lichens, vascular plants, and communities, and this will probably cause shifts in the dominance of both bottom layer and canopy layer species. It is important to note that the decrease in cover and species number of the bottom layer mainly occurred due to a decline in mosses; in contrast, lichen cover increased in all treatments in both communities. Climate change may thus cause a shift in the bottom layer from being dominated by mosses, to become dominated by lichens.
We simulated a predicted environmental change in a subarctic-alpine plant community to study short-term growth in Hylocomium splendens, and three-year effects in abundance changes of the five most common bryophytes at Latnjajaure, northern Sweden. The experiment had a factorial design with increased temperature and nutrients, alone and in combination. Moss growth was measured during the 1995 growing season, and we measured species abundance before and after three years of environmental perturbation. The combined treatments of fertilizer and temperature enhancement caused a decreased growth in length and dry weight in H. splendens. There was a significant decrease in abundance of Rhytidium rugosum in the combined temperature and fertilizer treatment. The other four common bryophyte species (Aulacomnium turgidum, Dicranum groenlandicum, Hylocomium splendens, and Ptilidium ciliare) showed weaker, but not significant trends in the same direction. As the bryophytes are an important component of arctic and subarctic vegetation, we expect that they will play a key role in the impact of anticipated Global Change on the ecosystems.
The present work investigates C and N allocation patterns in two forage plants: a forb, Oxyria digyna, and a sedge, Eriophorum angustifolium, in subarctic Sweden and high arctic Canada. Short-term changes in concentrations after simulated or natural herbivory (caused by Gynaephora groenlandica on Oxyria in the high arctic habitat) were also investigated. There were no clear differences in concentrations of C and N between the high arctic and subarctic sites in either species. In Oxyria of the subarctic habitat, the minimum N concentrations occurred at earlier phenological stages compared with plants in the high arctic habitat. Simulated herbivory increased the concentration of C in belowground tissues relative to those in control plants in Oxyria at the subarctic site, which may indicate increased allocation of non-C compounds to the growing shoots or daughter ramets. Herbivory by Gynaephora groenlandica caterpillars increased the N concentrations of Oxyria both in aboveground and belowground tissues, possible indicating increased uptake of N in the high arctic habitat. Eriophorum did not show clear trends in concentrations relating to habitat, phenology or simulated herbivory. The difference between Oxyria and Eriophorum in their response to herbivory apparently resulted from contrasting growth habits between the species. Tiller death after reproduction and long leaf life span may be the main reasons for the lack of clear patterns in concentrations in Eriophorum. Compensation after herbivory may be attained by the early production of daughter tillers in Eriophorum instead of the regrowth of the damaged ramets, as in Oxyria. Monitoring the responses for only one season is apparently a too short time period in these long-lived plants.
Changes in growing season temperature and duration may have profound effects on the population dynamics of arctic and alpine plant species in snow-bed and fell-field habitats. We examined how a typical herbaceous pioneer species, Ranunculus glacialis, responded to experimental climate change in open-top chambers for three seasons at an alpine site in southern Norway. Warming had no significant effect on any reproductive, growth or phenological variables, except for seed weight, which increased significantly during the first 2 ears. Despite large differences in average date of snow-melt among years, average reproductive output and ramet size differed little among years. Within-year variation in date of snowmelt had no impact on seed number or weight in either control or warmed plots. Leaf width and ramet leaf number decreased significantly with later snowmelt within a year. Experimental warming reduced the negative effect on ramet size of late snowmelt within a year to some extent. In general, R. glacialis reacts contrary to most other arctic/alpine species to experimental warming. Species with such low responsiveness to environmental conditions may be particularly vulnerable to climatic change, especially if their habitat is invaded by other species with higher phenotypic plasticity and a better competitive ability.
Recent observations of changes in some tundra ecosystems appear to be responses to a warming climate. Several experimental studies have shown that tundra plants and ecosystems can respond strongly to environmental change, including warming; however, most studies were limited to a single location and were of short duration and based on a variety of experimental designs. In addition, comparisons among studies are difficult because a variety of techniques have been used to achieve experimental warming and different measurements have been used to assess responses. We used metaanalysis on plant community measurements from standardized warming experiments at 11 locations across the tundra biome involved in the International Tundra Experiment. The passive warming treatment increased plant-level air temperature by 1-3 degrees C, which is in the range of predicted and observed warming for tundra regions. Responses were rapid and detected in whole plant communities after only two growing seasons. Overall, warming increased height and cover of deciduous shrubs and graminoids, decreased cover of mosses and lichens, and decreased species diversity and evenness. These results predict that warming will cause a decline in biodiversity across a wide variety of tundra, at least in the short term. They also provide rigorous experimental evidence that recently observed increases in shrub cover in many tundra regions are in response to climate warming. These changes have important implications for processes and interactions within tundra ecosystems and between tundra and the atmosphere.