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Arctic ecosystems are increasingly exposed to extreme climatic events throughout the year, which can affect species performance. Cryptogams (bryophytes and lichens) provide important ecosystem services in polar ecosystems but may be physiologically affected or killed by extreme events. Through field and laboratory manipulations, we compared physiological responses of seven dominant sub-Arctic cryptogams (three bryophytes, four lichens) to single events and factorial combinations of mid-winter heatwave (6°C for 7 days), re-freezing, snow removal and summer nitrogen addition. We aimed to identify which mosses and lichens are vulnerable to these abiotic extremes and if combinations would exacerbate physiological responses. Combinations of extremes resulted in stronger species responses but included idiosyncratic species-specific responses. Species that remained dormant during winter (March), irrespective of extremes, showed little physiological response during summer (August). However, winter physiological activity, and response to winter extremes, was not consistently associated with summer physiological impacts. Winter extremes affect cryptogam physiology, but summer responses appear mild, and lichens affect the photobiont more than the mycobiont. Accounting for Arctic cryptogam response to multiple climatic extremes in ecosystem functioning and modelling will require a better understanding of their winter eco-physiology and repair capabilities.

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Climate change is one of many ongoing human-induced environmental changes, but few studies consider interactive effects between multiple anthropogenic disturbances. In coastal sub-arctic heathland, we quantified the impact of a factorial design simulating extreme winter warming (WW) events (7 days at 6–7∘C) combined with episodic summer nitrogen (+N) depositions (5 kg N ha-1) on plant winter physiology, plant community composition and ecosystem CO2 fluxes of an Empetrum nigrum dominated heathland during 3 consecutive years in northern Norway. We expected that the +N would exacerbate any stress effects caused by the WW treatment. During WW events, ecosystem respiration doubled, leaf respiration declined (-58%), efficiency of Photosystem II (Fv/Fm) increased (between 26 and 88%), while cell membrane fatty acids showed strong compositional changes as a result of the warming and freezing. In particular, longer fatty acid chains increased as a result of WW events, and eicosadienoic acid (C20:2) was lower when plants were exposed to the combination of WW and +N. A larval outbreak of geometrid moths (Epirrita autumnata and Operophtera brumata) following the first WW led to a near-complete leaf defoliation of the dominant dwarf shrubs E. nigrum (-87%) and Vaccinium myrtillus (-81%) across all experimental plots. Leaf emergence timing, plant biomass or composition, NDVI and growing season ecosystem CO2 fluxes were unresponsive to the WW and +N treatments. The limited plant community response reflected the relative mild winter freezing temperatures (-6.6∘C to -11.8∘C) recorded after the WW events, and that the grazing pressure probably overshadowed any potential treatment effects. The grazing pressure and WW both induce damage to the evergreen shrubs and their combination should therefore be even stronger. In addition, +N could have exacerbated the impact of both extreme events, but the ecosystem responses did not support this. Therefore, our results indicate that these sub-arctic Empetrum-dominated ecosystems are highly resilient and that their responses may be limited to the event with the strongest impact.