Tab. 1: Mean water loss by mosses and lichens at 40%

The loss of water in a desiccating atmosphere (c.40% r.h. at 10°C) and uptake of water from a saturated atmosphere (100% r.h. at 10°C) was recorded at intervals over periods of many hours or days in the dominant mosses and macroiichens occurring near the Australian Casey Station. Wilkes Land, continental Antarctica. While major differences exist in the water holding capacity and rates of water loss between mosses and lichens, the minimum levels attained after prolonged exposure to desiccating conditions are remarkably similar. By contrast, the volume of water absorbed from a saturated atmosphere is very similar in both groups of cryptogams. Morphological and anatomical characters are responsible for many of the differences, both between species, and within species exhibiting different growth features. Thus, significantly larger amounts of water are held by colonies of Bryum algens with a dense tomentum of rhizoids than those with sparse rhizoids; similarly, the rhizinate Umbilicaria aprina held a greater volume of water than the erhizinate U. decussata. The filamentous mat form of Alectoria mimiscula permits a much higher water content to be attained than in the coarser fruticose forms of Usnea sphacelata and U. antarctica. The dense shoot arrangement in Schistidium antarcticum accounts for the high water holding capacity in the hydric turf form whereas the less densely packed shoots and thicker cell walls of the xeric cushion form maintain a lower water content. The rate of water loss (as percentage dry weight) was much faster in the turf form of Schistidium and tomenlose form of Bryum, although this trend was reversed when expressed as percentage of the initial water content. Minimal water contents arc achieved by the lichens in desiccating conditions within 6-12 hours, whereas the mosses take several times longer. The water relations characteristics of these cryptogams are considered in the light of their distribution in the field and of their metabolic activity under prevailing Antarctic conditions.

14C activity of Antarctic lichens

By the nuclear bomb tests during the 1950s and early 1960s, the radiocarbon content of the atmospheric CO, on the Southern Hemisphere rose within a few years from 98 to 162% of the standard recent value and then dropped to 122% (at the end of 1984). This rapid fluctuation was used to determine the lifetime of five species of lichens collected in the beginning of 1985 in the maritime Antarctic. Under the assumption that Lichens assimilate each year carbon at the same rate and that carbon once fixed at least in main branches never will be exchanged later on. The age of mature thalli of Caioplaco regalis, Ramalino tetebrata and Ustiea antarctica was determined to 32 years, while U, aurantiaco-atra and Himantormia lugubris gave an age of ca. 38 years and ca. 60 years, respectively.

(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.

Gene sequences of the lichen Cetraria aculeata

Lichens are symbioses between fungi (mycobionts) and photoautotrophic green algae or cyanobacteria (photobionts). Many lichens occupy large distributional ranges covering several climatic zones. So far, little is known about the large-scale phylogeography of lichen photobionts and their role in shaping the distributional ranges of lichens. We studied south polar, temperate and north polar populations of the widely distributed fruticose lichen Cetraria aculeata. Based on the DNA sequences from three loci for each symbiont, we compared the genetic structure of mycobionts and photobionts. Phylogenetic reconstructions and Bayesian clustering methods divided the mycobiont and photobiont data sets into three groups. An AMOVA shows that the genetic variance of the photobiont is best explained by differentiation between temperate and polar regions and that of the mycobiont by an interaction of climatic and geographical factors. By partialling out the relative contribution of climate, geography and codispersal, we found that the most relevant factors shaping the genetic structure of the photobiont are climate and a history of codispersal. Mycobionts in the temperate region are consistently associated with a specific photobiont lineage. We therefore conclude that a photobiont switch in the past enabled C. aculeata to colonize temperate as well as polar habitats. Rare photobiont switches may increase the geographical range and ecological niche of lichen mycobionts by associating them with locally adapted photobionts in climatically different regions and, together with isolation by distance, may lead to genetic isolation between populations and thus drive the evolution of lichens.

Chlorophyll content and temperature-dependent net photosynthesis and respiration rate in the lichen Cetraria aculeata

We studied polar and temperate samples of the lichen Cetraria aculeata to investigate whether genetical differences between photobionts are correlated with physiological properties of the lichen holobiont. Net photosynthesis and dark respiration (DR) at different temperatures (from 0 to 30 °C) and photon flux densities (from 0 to 1,200 ?mol/m**2/s) were studied for four populations of Cetraria aculeata. Samples were collected from maritime Antarctica, Svalbard, Germany and Spain, representing different climatic situations. Sequencing of the photobiont showed that the investigated samples fall in the polar and temperate clade described in Fernández-Mendoza et al. (2011, doi:10.1111/j.1365-294X.2010.04993.x). Lichens with photobionts from these clades differ in their temperature optimum for photosynthesis, maximal net photosynthesis, maximal DR and chlorophyll content. Maximal net photosynthesis was much lower in Antarctica and Svalbard than in Germany and Spain. The difference was smaller when rates were expressed by chlorophyll content. The same is true for the temperature optima of polar (11 °C) and temperate (15 and 17 °C) lichens. Our results indicate that lichen mycobionts may adapt or acclimate to local environmental conditions either by selecting algae from regional pools or by regulating algal cell numbers (chlorophyll content) within the thallus.

Carbon/Nitrogen ratio of plant tissues and biomass of belowground consumers after 11 years of experimental treatment, Toolik lake

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The Exploitation Ecosystem Hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.

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.

Plant Functional Type (PFT) map over Siberia derived from GlobCover 2005 dataset, link to dataset in NetCDF format

High-latitude ecosystems play an important role in the global carbon cycle and in regulating the climate system and are presently undergoing rapid environmental change. Accurate land cover data sets are required to both document these changes as well as to provide land-surface information for benchmarking and initializing Earth system models. Earth system models also require specific land cover classification systems based on plant functional types (PFTs), rather than species or ecosystems, and so post-processing of existing land cover data is often required. This study compares over Siberia, multiple land cover data sets against one another and with auxiliary data to identify key uncertainties that contribute to variability in PFT classifications that would introduce errors in Earth system modeling. Land cover classification systems from GLC 2000, GlobCover 2005 and 2009, and MODIS collections 5 and 5.1 are first aggregated to a common legend, and then compared to high-resolution land cover classification systems, vegetation continuous fields (MODIS VCFs) and satellite-derived tree heights (to discriminate against sparse, shrub, and forest vegetation). The GlobCover data set, with a lower threshold for tree cover and taller tree heights and a better spatial resolution, tends to have better distributions of tree cover compared to high-resolution data. It has therefore been chosen to build new PFT maps for the ORCHIDEE land surface model at 1 km scale. Compared to the original PFT data set, the new PFT maps based on GlobCover 2005 and an updated cross-walking approach mainly differ in the characterization of forests and degree of tree cover. The partition of grasslands and bare soils now appears more realistic compared with ground truth data. This new vegetation map provides a framework for further development of new PFTs in the ORCHIDEE model like shrubs, lichens and mosses, to represent the water and carbon cycles in northern latitudes better. Updated land cover data sets are critical for improving and maintaining the relevance of Earth system models for assessing climate and human impacts on biogeochemistry and biophysics.

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.

Haplotype and nucleotide characteristics of photobiont and mycobiont gene sequences in the lichen Cetraria aculeata

Lichens, symbiotic associations of fungi (mycobionts) and green algae or cyanobacteria (photobionts), are poikilohydric organisms that are particularly well adapted to withstand adverse environmental conditions. Terrestrial ecosystems of the Antarctic are therefore largely dominated by lichens. The effects of global climate change are especially pronounced in the maritime Antarctic and it may be assumed that the lichen vegetation will profoundly change in the future. The genetic diversity of populations is closely correlated to their ability to adapt to changing environmental conditions and to their future evolutionary potential. In this study, we present evidence for low genetic diversity in Antarctic mycobiont and photobiont populations of the widespread lichen Cetraria aculeata. We compared between 110 and 219 DNA sequences from each of three gene loci for each symbiont. A total of 222 individuals from three Antarctic and nine antiboreal, temperate and Arctic populations were investigated. The mycobiont diversity is highest in Arctic populations, while the photobionts are most diverse in temperate regions. Photobiont diversity decreases significantly towards the Antarctic but less markedly towards the Arctic, indicating that ecological factors play a minor role in determining the diversity of Antarctic photobiont populations. Richness estimators calculated for the four geographical regions suggest that the low genetic diversity of Antarctic populations is not a sampling artefact. Cetraria aculeata appears to have diversified in the Arctic and subsequently expanded its range into the Southern Hemisphere. The reduced genetic diversity in the Antarctic is most likely due to founder effects during long-distance colonization.

(Table 2) Collection sites and number of specimens of Usnea subgenus Neuropogon species

Usnea species of the Neuropogon group are amongst the most widespread and abundant macrolichens in Antarctic regions. Four principal species, U. antarctica, U. aurantiaco-atra, U. sphacelata and U. subantarctica, have been described on morphological grounds. However, identification to species level is often difficult and atypical morphologies frequently arise. Over 400 specimens were collected on the Antarctic Peninsula and Falkland Islands. Both morphological and molecular characters (ITS and RPB1) were used to compare samples to clarify taxonomic relationships. Morphological characteristics used included presence of apothecia, apothecial rays, soredia, papillae, fibrils, pigmentation and the diameter of the central axis as a proportion of branch diameter. Results revealed a very close relationship between U. antarctica and U. aurantiaco-atra, suggesting that they might constitute a species pair or be conspecific. Usnea sphacelata was comprised of at least two genetically distinct groups with no clear differences in morphology. One group included the first reported fertile specimen of this species. Usnea subantarctica was phylogenetically distinct from the other main Antarctic Usnea species, but clustered with U. trachycarpa. Genetic variation was evident within all species although there was no clear correlation between geographic origin and genetic relatedness. Phylogenetic analyses indicated that species circumscription in the Neuropogon group needs revision, with the principal species being non-monophyletic. None of the morphological characters, or groups of characters, used in this study proved to be completely unambiguous markers for a single species. However, axis thickness was supported as being informative for the identification of monophyletic lineages within the group.

Vegetation community and geochemistry in the Uummannaq District, West Greenland

Beach and salt marsh vegetation of the Uummannaq District, northern West Greenland (c. 70°15' N - 72° N, 49° W - 54° W) was studied 1998 according to the Braun-Blanquet phytosociological approach. Habitat analyses included soil chemistry. Such vegetation locally occurs and is not developed over extensive areas. On gravely stony beaches a Mertensia maritima ssp. maritima community occurs, while a Honckenya peploides var. diffusa community is confined to sandy beaches. The association Honckenyo diffusae-Elymetum mollis Thannh. 1975 is confined to sandy shore walls and low dunes. All vegetation types are assigned to the alliance Honckenyo- Elymion arenariae Tx. 1966, which again is a unit of the order Honckenyo- Elymetalia arenariae Tx. 1966, which is sub ordered to the class Honckenyo-Elymetea arenariae Tx. 1966. On fine sediments along sheltered coasts salt marsh vegetation is locally developed mainly on fiord deltas and outwash plains of small rivers and streams. A distinct zonation pattern in vegetation can be observed from the lower to upper salt marsh: Puccinellietum phryganodis Hadac 1946 association, Caricetum subspathaceae Hadac 1946 association, Caricetum ursinae Hadac 1946 association (all assigned to the alliance Puccinellion phryganodis Hadac 1946) and Festuco-Caricetum glareosae Nordh. 1954 association (assigned to the alliance Armerion maritimae Br.-Bl. et de Leeuw 1936). Both alliances are units of the order Glauco- Puccinellietalia Beeftink et Westhoff in Beeftink 1965, which is assigned to the class Asteretea tripolii Westhoff et Beeftink in Beeftink 1962. TWINSPAN and CCA support the vegetation classification and the CCA with soil chemistry parameters shows that salinity (related to position above MHW) and Ncontent are strongly correlated with the floristical differentiation of the vegetation of the Honckenyo-Elymetea class. In the Asteretea tripolii class, position above MHW (negatively correlated with pH, conductivity and Clcontent) and fresh water supply are likely the main factors, which affect vegetation differentiation. A synoptic survey of vegetation types from Greenland based on published phytosociological tables is presented and distribution of the vegetation types is addressed, just as their position in a circumpolar context. Moreover a Cochlearia groenlandica-Melandrium triflorum community is described as a new vegetation type, occurring on shallow soil on cliffs influenced by salt spray.