Greening of Svalbard.

Svalbard, located between 76°30'N and 80°50'N, is among the regions in the world with the most rapid temperature increase. We processed a cloud-free time-series of MODIS-NDVI for Svalbard. The dataset is interpolated to daily data during the 2000-2022 period with 232 m pixel resolution. The onset of growth, with a clear phenological definition, has been mapped each year. Then the integrated NDVI from the onset (O) of growth each year to the time of average (2000-2022) peak (P) of growth (OP NDVI) have been calculated. OP NDVI has previously shown high correlation with field-based tundra productivity. Daily mean temperature data from 11 meteorological stations are compared with the NDVI data. The OP NDVI values show very high and significant correlation with growing degree days computed from onset to time of peak of growth for all the meteorological stations used. On average for the entire Svalbard, the year 2016 first had the highest greening (OP NDVI values) recorded since the year 2000, then the greening in 2018 surpassed 2016, then 2020 surpassed 2018, and finally 2022 was the year with the overall highest greening by far for the whole 2000-2022 period. This shows a rapid recent greening of Svalbard very strongly linked to temperature increase, although there are regional differences: the eastern parts of Svalbard show the largest variability between years, most likely due to variability in the timing of sea-ice break-up in adjacent areas. Finally, we find that areas dominated by manured moss-tundra in the polar desert zone require new methodologies, as moss does not share the seasonal NDVI dynamics of tundra communities.

Changes in Onset of Vegetation Growth on Svalbard, 2000-2020.

The global temperature is increasing, and this is affecting the vegetation phenology in many parts of the world. The most prominent changes occur at northern latitudes such as our study area, which is Svalbard, located between 76°30'N and 80°50'N. A cloud-free time series of MODIS-NDVI data was processed. The dataset was interpolated to daily data during the 2000-2020 period with a 231.65 m pixel resolution. The onset of vegetation growth was mapped with a NDVI threshold method which corresponds well with a recent Sentinel-2 NDVI-based mapping of the onset of vegetation growth, which was in turn validated by a network of in-situ phenological data from time lapse cameras. The results show that the years 2000 and 2008 were extreme in terms of the late onset of vegetation growth. The year 2020 had the earliest onset of vegetation growth on Svalbard during the 21-year study. Each year since 2013 had an earlier or equally early timing in terms of the onset of the growth season compared with the 2000-2020 average. A linear trend of 0.57 days per year resulted in an earlier onset of growth of 12 days on average for the entire archipelago of Svalbard in 2020 compared to 2000.

The genus Massalongia (lichenised ascomycetae) in the Southern Hemisphere.

The species of Massalongia recorded and described from the Southern Hemisphere are revised and it is shown that only one is present; M. patagonica which is widespread, with populations in Australia and New Zealand that differ from the South American populations, but at present best regarded as part of the variation of that species. Records from this hemisphere of all other species placed in the genus are incorrect. The type species, M. carnosa, is restricted to the Northern Hemisphere. Two species, M. antarctica and M. novozelandica cannot be identified precisely due to lack of sufficient type material and with the types as the only collections known of these, but none belongs in Massalongia according to available data. Massalongia griseolobata (from Gough Isl.) is shown here to belong in the Pannariaceae and is part of the parmelielloid clade. M. intricata (from South Georgia) and M. olechiana (from South Shetland) have both recently been correctly transferred to the genus Steinera in the Arctomiaceae.

Algal and fungal diversity in Antarctic lichens.

The composition of lichen ecosystems except mycobiont and photobiont has not been evaluated intensively. In addition, recent studies to identify algal genotypes have raised questions about the specific relationship between mycobiont and photobiont. In the current study, we analyzed algal and fungal community structures in lichen species from King George Island, Antarctica, by pyrosequencing of eukaryotic large subunit (LSU) and algal internal transcribed spacer (ITS) domains of the nuclear rRNA gene. The sequencing results of LSU and ITS regions indicated that each lichen thallus contained diverse algal species. The major algal operational taxonomic unit (OTU) defined at a 99% similarity cutoff of LSU sequences accounted for 78.7-100% of the total algal community in each sample. In several cases, the major OTUs defined by LSU sequences were represented by two closely related OTUs defined by 98% sequence similarity of ITS domain. The results of LSU sequences indicated that lichen-associated fungi belonged to the Arthoniomycetes, Eurotiomycetes, Lecanoromycetes, Leotiomycetes, and Sordariomycetes of the Ascomycota, and Tremellomycetes and Cystobasidiomycetes of the Basidiomycota. The composition of major photobiont species and lichen-associated fungal community were mostly related to the mycobiont species. The contribution of growth forms or substrates on composition of photobiont and lichen-associated fungi was not evident.

Outbreaks by canopy-feeding geometrid moth cause state-dependent shifts in understorey plant communities.

The increased spread of insect outbreaks is among the most severe impacts of climate warming predicted for northern boreal forest ecosystems. Compound disturbances by insect herbivores can cause sharp transitions between vegetation states with implications for ecosystem productivity and climate feedbacks. By analysing vegetation plots prior to and immediately after a severe and widespread outbreak by geometrid moths in the birch forest-tundra ecotone, we document a shift in forest understorey community composition in response to the moth outbreak. Prior to the moth outbreak, the plots divided into two oligotrophic and one eutrophic plant community. The moth outbreak caused a vegetation state shift in the two oligotrophic communities, but only minor changes in the eutrophic community. In the spatially most widespread communities, oligotrophic dwarf shrub birch forest, dominance by the allelopathic dwarf shrub Empetrum nigrum ssp. hermaphroditum, was effectively broken and replaced by a community dominated by the graminoid Avenella flexuosa, in a manner qualitatively similar to the effect of wild fires in E. nigrum communities in coniferous boreal forest further south. As dominance by E. nigrum is associated with retrogressive succession the observed vegetation state shift has widespread implications for ecosystem productivity on a regional scale. Our findings reveal that the impact of moth outbreaks on the northern boreal birch forest system is highly initial-state dependent, and that the widespread oligotrophic communities have a low resistance to such disturbances. This provides a case for the notion that climate impacts on arctic and northern boreal vegetation may take place most abruptly when conveyed by changed dynamics of irruptive herbivores.

PHYLOGENETIC PATTERNS AMONG NOSTOC CYANOBIONTS WITHIN BI- AND TRIPARTITE LICHENS OF THE GENUS PANNARIA(1).

Phylogenetic relationships between Nostoc cyanobionts in the lichen genus Pannaria were studied to evaluate their correlation to geography, habitat ecology, and other patterns previously reported. The 16S rRNA gene sequences of a total of 37 samples of 21 Pannaria species from seven countries from the Northern and Southern hemispheres were analyzed and compared with 69 free-living and symbiotic cyanobacterial strains. The sequences from Pannaria were distributed throughout a branch of Nostoc sequences previously called "the Nephroma guild," and within two subgroups from another branch, referred to as the "Peltigera guild," although there was a gradual transition between the two major groups. There is a more diverse pattern of relationships between Nostoc sequences from bipartite versus tripartite lichen species in Pannaria, compared with other well-studied genera, such as Nephroma and Peltigera. Cyanobionts from several tripartite Pannaria species from the Southern Hemisphere and corticolous bipartite species from both hemispheres were grouped together. Four sequences of Pannaria and Pseudocyphellaria cyanobionts from rocks in the Chilean Juan Fernández Islands were nested within corticolous cyanobionts, whereas the terricolous "Pannaria sphinctrina clade" was placed with other terricolous strains. The cluster patterns derived from phylogenetic analysis were partly reflecting lichen taxonomy, in two groups of lichen species, possibly indicating coevolution. The phylogram partly also reflected lichen ecology. Three Pannaria species have very different cyanobiont strains when they grow in different habitats.

Seasonal trends in usnic acid concentrations of Arctic, alpine and Patagonian populations of the lichen Flavocetraria nivalis.

The widespread secondary metabolite usnic acid, a dibenzofuran derivative, is the principal acetone-soluble compound in the lichen Flavocetraria nivalis. Seasonal variation in concentrations were studied in four populations of this lichen, three from Arctic-alpine habitats in the Northern Hemisphere, and one from Patagonian heathland in the Southern Hemisphere. Usnic acid is produced in large amounts, making up between 4% and 8% of thallus dry weight. Large seasonal variation is seen, with a trend towards peak levels in late spring and early summer, and generally low levels during autumn and winter. However, at an Arctic steppe in Central West Greenland, remarkably high levels were also detected during late autumn and early winter. Comparisons with environmental data using model selection procedures show that usnic acid levels of three of the populations are positively correlated with time of season, as measured by the proximity in time to nearest summer solstice, solar radiation levels, and temperature conditions. All these three variables are intercorrelated, thus indicating the same overall trend. For the three driest sites, precipitation rates are included in the models that best explain the variation in usnic acid. However, the explanatory powers of the models are generally low, partly due to high variation between thalli growing together and sampled at the same time. This is the first attempt to compare statistically seasonal variation in usnic acid concentrations and environmental variables, and thus also the first time it is shown that the concentration in various populations of the same lichen species shows different types of correlation with seasonal climatic changes.

Effects of ultraviolet radiation and PAR on the content of usnic and divaricatic acids in two arctic-alpine lichens.

The fruticose lichen Flavocetraria nivalis and the crustose lichen Ophioparma ventosa, both common in light-exposed arctic-alpine environments, were exposed to ultraviolet radiation (UVR) in growth chambers for 30 days. Treatment with visible light (PAR) served as control. Both species accumulate the UV-absorbing phenolic compound usnic acid in the upper cortex. The latter species also synthesises several UV-absorbing medullary compounds, among them divaricatic acid. The effects of treatment with UVR on the synthesis of these two compounds were investigated by analysing the compounds quantitatively by RP-HPLC. UV-exposed thallus tips of F. nivalis contained higher concentrations of usnic acid than those not grown under UVR. Both treatments had a positive effect on the synthesis of usnic acid in O. ventosa. An additional experiment with O. ventosa was performed by first storing samples in a low-light habitat for 1 year to obtain near-zero levels of phenolics, and thereby exposing the samples to UVR and PAR for 90 days. A rapid resynthesis of usnic acid was observed for both treatments. The amounts of divaricatic acid were highly variable in all groups, and were not correlated with usnic acid concentrations or treatments. A comparison of O. ventosa from three different habitat types showed that the highest usnic acid amounts were found in the habitat with the highest levels of solar radiation. Results indicate that the induction of usnic acid production by UVR depends on the species studied, and on how well acclimatised the lichen samples are to solar radiation before they are exposed to supplementary UVR. In lichens with an already well-developed internal screening capacity, like the population of F. nivalis, enhanced UVR need not induce further accumulation of usnic acid, but removal of UVR may induce a biodegradation of usnic acid. Results also indicate that PAR is just as important as UVR for triggering the resynthesis of usnic acid in shade-adapted lichens. Divaricatic acid seems to be of low importance for the UV-screening properties of O. ventosa.