(Table 1) Species richness and diversity of tundra plants in herbivore exclosure and control plots near Barrow, Alaska

To determine the role lemmings play in structuring plant communities and their contribution to the 'greening of the Arctic', we measured plant cover and biomass in 50 + year old lemming exclosures and control plots in the coastal tundra near Barrow, Alaska. The response of plant functional types to herbivore exclusion varied among land cover types. In general, the abundance of lichens and bryophytes increased with the exclusion of lemmings, whereas graminoids decreased, although the magnitude of these responses varied among land cover types. These results suggest that sustained lemming activity promotes a higher biomass of vascular plant functional types than would be expected without their presence and highlights the importance of considering herbivory when interpreting patterns of greening in the Arctic. In light of the rapid environmental change ongoing in the Arctic and the potential regional to global implications of this change, further exploration regarding the long-term influence of arvicoline rodents on ecosystem function (e.g. carbon and energy balance) should be considered a research priority.

Flower and seed biomass, and germination rate of arctic tundra plants in response to long term experimental warming

We provide new information on changes in tundra plant sexual reproduction in response to long-term (12 years) experimental warming in the High Arctic. Open-top chambers (OTCs) were used to increase growing season temperatures by 1-2 °C across a range of vascular plant communities. The warming enhanced reproductive effort and success in most species; shrubs and graminoids appeared to be more responsive than forbs. We found that the measured effects of warming on sexual reproduction were more consistently positive and to a greater degree in polar oasis compared with polar semidesert vascular plant communities. Our findings support predictions that long-term warming in the High Arctic will likely enhance sexual reproduction in tundra plants, which could lead to an increase in plant cover. Greater abundance of vegetation has implications for primary consumers - via increased forage availability, and the global carbon budget - as a function of changes in permafrost and vegetation acting as a carbon sink. Enhanced sexual reproduction in Arctic vascular plants may lead to increased genetic variability of offspring, and consequently improved chances of survival in a changing environment. Our findings also indicate that with future warming, polar oases may play an important role as a seed source to the surrounding polar desert landscape.

Initial biomass and biomass change of tundra and forest plants in three herbivore treatments in Norway and Sweden

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana, has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (> 0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (< 0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.

Environmental characteristics, and growth traits and leaf chemistry of tundra plants in a warming experiment at Alexandra Fiord

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate-vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open-top chambers (OTCs) to passively warm vegetation by 1-2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf d15N. Long-term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect d15N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.

Sampling sites, sampled birds and plants of young larch plantations in Tokachi district, eastern Hokkaido, northern Japan, 2011

Models and data used to describe species-area relationships confound sampling with ecological process as they fail to acknowledge that estimates of species richness arise due to sampling. This compromises our ability to make ecological inferences from and about species-area relationships. We develop and illustrate hierarchical community models of abundance and frequency to estimate species richness. The models we propose separate sampling from ecological processes by explicitly accounting for the fact that sampled patches are seldom completely covered by sampling plots and that individuals present in the sampling plots are imperfectly detected. We propose a multispecies abundance model in which community assembly is treated as the summation of an ensemble of species-level Poisson processes and estimate patch-level species richness as a derived parameter. We use sampling process models appropriate for specific survey methods. We propose a multispecies frequency model that treats the number of plots in which a species occurs as a binomial process. We illustrate these models using data collected in surveys of early-successional bird species and plants in young forest plantation patches. Results indicate that only mature forest plant species deviated from the constant density hypothesis, but the null model suggested that the deviations were too small to alter the form of species-area relationships. Nevertheless, results from simulations clearly show that the aggregate pattern of individual species density-area relationships and occurrence probability-area relationships can alter the form of species-area relationships. The plant community model estimated that only half of the species present in the regional species pool were encountered during the survey. The modeling framework we propose explicitly accounts for sampling processes so that ecological processes can be examined free of sampling artefacts. Our modeling approach is extensible and could be applied to a variety of study designs and allows the inclusion of additional environmental covariates.

Effects of endophyte colonization of Vicia faba (Fabaceae) plants on the life

The effects of fungal endophytes Beauveria bassiana isolates ICIPE 279, G1LU3, S4SU1 and Hypocrea lixii isolate F3ST1 on the life-history of Phaedrotoma scabriventris and Diglyphus isaea, parasitoids of the pea leafminer Liriomyza huidobrensis, were studied in the laboratory. Parasitoids were allowed to parasitize 2nd-3rd L. huidobrensis larvae reared on endophytically-inoculated Vicia faba. In the control, parasitoids were reared on non-inoculated host plants. Parasitism, pupation, adult emergence and survival were recorded. No significant difference was observed between the control and the endophyte-inoculated plants in terms of parasitism rates of P. scabriventris (p = 0.68) and D. isaea (p = 0.45) and adult' survival times (p = 0.06). The survival period of the F1 progeny of P. scabriventris was reduced (p < 0.0001) in B. bassiana S4SU1 to 28 days as compared to more than 40 days for B. bassiana G1LU3, ICIPE 279 and H. lixii F3ST1. However, no significant difference (p = 0.54) was observed in the survival times of the F1 progeny of D. isaea. This study has therefore demonstrated the beneficial effects of both endophytes and parasitoids in L. huidobrensis population suppression.

(Table 1) Chlorophyll concentration and pigment composition of carotenoids in three heathland bryophytes, northern Sweden

Background and Aims: Anthropogenic depletion of stratospheric ozone in Arctic latitudes has resulted in an increase of ultraviolet-B radiation (UV-B) reaching the biosphere. UV-B exposure is known to reduce aboveground biomass and plant height, to increase DNA damage and cause accumulation of UV-absorbing compounds in polar plants. However, many studies on Arctic mosses tended to be inconclusive. The importance of different water availability in influencing UV-B impacts on lower plants in the Arctic has been poorly explored and might partially explain the observed wide variation of responses, given the importance of water in controlling bryophyte physiology. This study aimed to assess the long-term responses of three common sub-Arctic bryophytes to enhanced UV-B radiation (+UV-B) and to elucidate the influence of water supply on those responses. Results: Responses were species specific: H. splendens responded most to +UV-B, with reduction in both annual growth (-22%) and sporophyte production (-44%), together with increased b-carotene, violaxanthin, total chlorophyll and NPQ, and decreased zeaxanthin and de-epoxidation of the xanthophyll cycle pool (DES). Barbilophozia lycopodioides responded less to +UV-B, showing increased b-carotene and sclerophylly and decreased UV-absorbing compounds. Polytrichum commune only showed small morphogenetic changes. No effect of UV-B on bryophyte cover was observed. Water availability had profound effects on bryophyte ecophysiology, and plants showed, in general, lower growth and ETR, together with a higher photoprotection in the drier site. Water availability also influenced bryophyte responses to +UV-B and, in particular, responses were less detectable in the drier site. Conclusions: Impacts of UV-B exposure on Arctic bryophytes were significant, in contrast to modest or absent UV-B effects measured in previous studies. The impacts were more easily detectable in species with high plasticity such as H. splendens and less obvious, or more subtle, under drier conditions. Species biology and water supply greatly influences the impact of UV-B on at least some Arctic bryophytes and could contribute to the wide variation of responses observed previously.