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.

(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.

Species occurrence and richness of lichen, bryophytes and vascular plants near Abisko, Sweden and Toolik lake, Alaska

Little is known about the impact of changing temperature regimes on composition and diversity of cryptogam communities in the Arctic and Subarctic, despite the well-known importance of lichens and bryophytes to the functioning and climate feedbacks of northern ecosystems. We investigated changes in diversity and abundance of lichens and bryophytes within long-term (9-16 years) warming experiments and along natural climatic gradients, ranging from Swedish subarctic birch forest and subarctic/subalpine tundra to Alaskan arctic tussock tundra. In both Sweden and Alaska, lichen diversity responded negatively to experimental warming (with the exception of a birch forest) and to higher temperatures along climatic gradients. Bryophytes were less sensitive to experimental warming than lichens, but depending on the length of the gradient, bryophyte diversity decreased both with increasing temperatures and at extremely low temperatures. Among bryophytes, Sphagnum mosses were particularly resistant to experimental warming in terms of both abundance and diversity. Temperature, on both continents, was the main driver of species composition within experiments and along gradients, with the exception of the Swedish subarctic birch forest where amount of litter constituted the best explanatory variable. In a warming experiment in moist acidic tussock tundra in Alaska, temperature together with soil ammonium availability were the most important factors influencing species composition. Overall, dwarf shrub abundance (deciduous and evergreen) was positively related to warming but so were the bryophytes Sphagnum girgensohnii, Hylocomium splendens and Pleurozium schreberi; the majority of other cryptogams showed a negative relationship to warming. This unique combination of intercontinental comparison, natural gradient studies and experimental studies shows that cryptogam diversity and abundance, especially within lichens, is likely to decrease under arctic climate warming. Given the many ecosystem processes affected by cryptogams in high latitudes (e.g. carbon sequestration, N2-fixation, trophic interactions), these changes will have important feedback consequences for ecosystem functions and climate.

Sampling data, experimental data and fecal pellet characteristics of water pump samples in the strait of Øresund between Denmark and Sweden during 2004/2005

Copepod fecal pellets are often degraded at high rates within the upper part of the water column. However, the identity of the degraders and the processes governing the degradation remain unresolved. To identify the pellet degraders we collected water from Øresund (Denmark) approximately every second month from July 2004 to July 2005. These water samples were divided into 5 fractions (<0.2, <2, <20, <100, <200 µm) and total (unfractionated). We determined fecal pellet degradation rate and species composition of the plankton from triplicate incubations of each fraction and a known, added amount of fecal pellets. The total degradation rate of pellets by the natural plankton community of Øresund followed the phytoplankton biomass, with maximum degradation rate during the spring bloom (2.5 ± 0.49 d**-1) and minimum (0.52 ± 0.14 d**-1) during late winter. Total pellet removal rate ranged from 22% d**-1 (July 2005) to 87% d**-1 (May). Protozooplankton (dinoflagellates and ciliates) in the size range of 20 to 100 µm were the key degraders of the fecal pellets, contributing from 15 to 53% of the total degradation rate. Free-living in situ bacteria did not affect pellet degradation rate significantly; however, culture-originating bacteria introduced in association with the pellets contributed up to 59% of the total degradation rate. An effect of late-stage copepod nauplii (>200 µm) was indicated, but this was not a dominating degradation process. Mesozooplankton did not contribute significantly to the degradation. However, grazing of mesozooplankton on the pellet degraders impacts pellet degradation rate indirectly. In conclusion, protozooplankton seems to include the key organisms for the recycling of copepod fecal pellets in the water column, both through the microbial loop and, especially, by functioning as an effective 'protozoan filter' for fecal pellets.

Establishment, expansion and regeneration of aspen (Populus tremula) in the Torneträsk area, subarctic Sweden

In subarctic Sweden, recent decadal colonization and expansion of aspen (Populus tremula L.) were recorded. Over the past 100 years, aspen became c. 16 times more abundant, mainly as a result of increased sexual regeneration. Moreover, aspen now reach tree-size (>2 m) at the alpine treeline, an ecotone that has been dominated by mountain birch (Betula pubescens ssp. czerepanovii) for at least the past 4000 years. We found that sexual regeneration in aspen probably occurred seven times or more within the last century. Whereas sexual regeneration occurred during moist years following a year with an exceptionally high June-July temperature, asexual regeneration was favored by warm and dry summers. Disturbance to the birch forest by cyclic moth population outbreaks was critical in aspen establishment in the subalpine area. At the treeline, aspen colonization was less determined by these moth outbreaks, and was mainly restricted by summer temperature. If summer warming persists, aspen spread may continue in subarctic Sweden, particularly at the treeline. However, changing disturbance regimes, future herbivore population dynamics and the responses of aspen's competitors birch and pine to a changing climate may result in different outcomes.

Characteristics and changes of thermokarst lakes in Canada, Russia and Sweden

Peat plateaus are widespread at high northern latitudes and are important soil organic carbon reservoirs. A warming climate can cause either increased ground subsidence (thermokarst) resulting in lake formation or increased drainage as the permafrost thaws. A better understanding of spatiotemporal variations in these landforms in relation to climate change is important for predicting the future thawing permafrost carbon feedback. In this study, dynamics in thermokarst lake extent during the last 35-50 years has been quantified through time series analysis of aerial photographs and high-resolution satellite images (IKONOS/QuickBird) in three peat plateau complexes, spread out across the northern circumpolar region along a climatic and permafrost gradient. From the mid-1970s until the mid-2000s there has been an increase in mean annual air temperature, winter precipitation, and ground temperature in all three study areas. The two peat plateaus located in the continuous and discontinuous permafrost zones, respectively, where mean annual air temperatures are below -5°C and ground temperatures are -2°C or colder, have experienced small changes in thermokarst lake extent. In the peat plateau located in the sporadic permafrost zone where the mean annual air temperature is around -3°C, and the ground temperature is close to 0°C, lake drainage and infilling with fen vegetation has been extensive and many new thermokarst lakes have formed. In a future progressively warmer and wetter climate permafrost degradation can cause significant impacts on landscape composition and greenhouse gas exchange also in areas with extensive peat plateaus, which presently still experience stable permafrost conditions.

Molecular dissolved organic matter composition in lakes across Sweden as relative intensities of FT-ICR-MS peaks and PARAFAC components and optical indices

Whether intrinsic molecular properties or extrinsic factors such as environmental conditions control the decomposition of natural organic matter across soil, marine and freshwater systems has been subject to debate. Comprehensive evaluations of the controls that molecular structure exerts on organic matter's persistence in the environment have been precluded by organic matter's extreme complexity. Here we examine dissolved organic matter from 109 Swedish lakes using ultrahigh-resolution mass spectrometry and optical spectroscopy to investigate the constraints on its persistence in the environment. We find that degradation processes preferentially remove oxidized, aromatic compounds, whereas reduced, aliphatic and N-containing compounds are either resistant to degradation or tightly cycled and thus persist in aquatic systems. The patterns we observe for individual molecules are consistent with our measurements of emergent bulk characteristics of organic matter at wide geographic and temporal scales, as reflected by optical properties. We conclude that intrinsic molecular properties are an important control of overall organic matter reactivity.