1. Climate change in northern high latitudes is predicted to be greater in winter rather than summer, yet little is known about the effects of winter climate change on northern ecosystems. Among the ...unknowns are the effects of an increasing frequency of acute, short-lasting winter warming events. Such events can damage higher plants exposed to warm, then returning cold, temperatures after snow melt, and it is not known how bryophytes and lichens, which are of considerable ecological importance in high-latitude ecosystems, are affected by such warming events. However, even physiological adaptations of these cryptogams to winter environments in general are poorly understood. 2. Here we describe findings from a novel field experiment that uses heating from infrared lamps and soil warming cables to simulate acute mid-winter warming events in a sub-Arctic heath. In particular, we report the growing season responses of the dominant lichen, Peltigera aphthosa, and bryophyte, Hylocomium splendens, to warming events in three consecutive winters. 3. While summertime photosynthetic performance of P. aphthosa was unaffected by the winter warming treatments, H. splendens showed significant reductions in net photosynthetic rates and growth rates (of up to 48% and 52%, respectively). Negative effects were evident already during the summer following the first winter warming event. 4. While the lichen develops without going through critical phenological stages during which vulnerable organs are produced, the moss has a seasonal rhythm, which includes initiation of growth of young, freeze-susceptible shoot apices in the early growing season; these might be damaged by breaking of dormancy during warm winter events. 5. Synthesis. Different sensitivities of the bryophyte and lichen species were unexpected, and illustrate that very little is known about the winter ecology of bryophytes and lichens from cold biomes in general. In sharp contrast to summer warming experiments that show increased vascular plant biomass and reduced lichen biomass, these results demonstrate that acute climate events in midwinter may be readily tolerated by lichens, in contrast to previously observed sensitivity of co-occurring dwarf shrubs, suggesting winter climate change may compensate for (or even reverse) predicted lichen declines resulting from summer warming.
Experimental studies have shown that deposition of reactive nitrogen is an important driver of plant community change, however, most of these experiments are of short duration with unrealistic ...treatments, and conducted in regions with elevated ambient deposition. Studies of spatial gradients of pollution can complement experimental data and indicate whether the potential impacts demonstrated by experiments are actually occurring in the ‘real world’. However, targeted surveys exist for only a very few habitats and are not readily comparable. In a coordinated campaign, we determined the species richness and plant community composition of five widespread, semi-natural habitats across Great Britain in sites stratified along gradients of climate and pollution, and related these ecological parameters to major drivers of biodiversity, including climate, pollution deposition, and local edaphic factors. In every habitat, we found reduced species richness and changed species composition associated with higher nitrogen deposition, with remarkable consistency in relative species loss across ecosystem types. Whereas the diversity of mosses, lichens, forbs, and graminoids declines with N deposition in different habitats, the cover of graminoids generally increases. Considered alongside previous experimental studies and survey work, our results provide a compelling argument that nitrogen deposition is a widespread and pervasive threat to terrestrial ecosystems.
Ecosystems limited in phosphorous (P) are widespread, yet there is limited
understanding of how these ecosystems may respond to anthropogenic
deposition of nitrogen (N) and the interconnected effects ...on the
biogeochemical cycling of carbon (C), N, and P. Here, we investigate the
consequences of enhanced N addition for the C–N–P pools of two P-limited
grasslands, one acidic and one limestone, occurring on contrasting soils, and we
explore their responses to a long-term nutrient-manipulation experiment. We
do this by combining data with an integrated C–N–P cycling model (N14CP). We
explore the role of P-access mechanisms by allowing these to vary in the
modelling framework and comparing model plant–soil C–N–P outputs to
empirical data. Combinations of organic P access and inorganic P
availability most closely representing empirical data were used to simulate
the grasslands and quantify their temporal response to nutrient
manipulation. The model suggested that access to organic P is a key
determinant of grassland nutrient limitation and responses to experimental N
and P manipulation. A high rate of organic P access allowed the acidic
grassland to overcome N-induced P limitation, increasing biomass C input to
soil and promoting soil
organic carbon (SOC) sequestration in response to N addition. Conversely,
poor accessibility of organic P for the limestone grassland meant N
provision exacerbated P limitation and reduced biomass input to the soil,
reducing soil carbon storage. Plant acquisition of organic P may therefore
play an important role in reducing P limitation and determining responses
to anthropogenic changes in nutrient availability. We conclude that
grasslands differing in their access to organic P may respond to N
deposition in contrasting ways, and where access is limited, soil organic
carbon stocks could decline.
Arctic ecosystems are under pressure from climate change and atmospheric nitrogen (N) deposition. However, knowledge of the ecology of microbial communities and their responses to such challenges in ...Arctic tundra soil remain limited, despite the central role these organisms play for ecosystem functioning. We utilised a plot-scale experiment in High Arctic tundra on Svalbard to investigate short-term variation (9 days), following simulation of a N deposition event (4 kg N ha
−1
yr
−1
), in the structure and abundance of bacterial, archaeal and fungal communities between organic and mineral soil horizons. T-RFLP analysis showed significant differences between horizons in bacterial and archaeal community structure. Q-PCR analysis showed that fungal abundance did not differ significantly between soil horizons, whilst bacterial and archaeal abundance was significantly higher in mineral than in organic horizons, despite soil water and total C and N contents being significantly greater in the organic horizon. In the organic horizon, bacterial community structure and fungal abundance varied significantly over time. In the mineral horizon, there was significant variation over time in bacterial abundance, in archaeal community structure and in both fungal community structure and abundance. In contrast, N deposition did not lead to significant variation in either the structure or the abundance of microbial communities. This research demonstrates that microbial community structure and abundance can change rapidly (over only a few days) in Arctic tundra soils and also differently between soil horizons in response to different environmental drivers. Moreover, this variability in microbial community structure and abundance is soil horizon- and taxonomic domain-specific, highlighting the importance of investigating microbial communities across all soil horizons and over short periods of time.
Estimates of vegetation carbon pools and their turnover rates are central to understanding and modelling ecosystem responses to climate change and their feedbacks to climate. In the Arctic, a region ...containing globally important stores of soil carbon, and where the most rapid climate change is expected over the coming century, plant communities have on average sixfold more biomass below ground than above ground, but knowledge of the root carbon pool sizes and turnover rates is limited. Here, we show that across eight plant communities, there is a significant positive relationship between leaf and fine root turnover rates (r2 = 0.68, P < 0.05), and that the turnover rates of both leaf (r2 = 0.63, P < 0.05) and fine root (r2 = 0.55, P < 0.05) pools are strongly correlated with leaf area index (LAI, leaf area per unit ground area). This coupling of root and leaf dynamics supports the theory of a whole‐plant economics spectrum. We also show that the size of the fine root carbon pool initially increases linearly with increasing LAI, and then levels off at LAI = 1 m2 m−2, suggesting a functional balance between investment in leaves and fine roots at the whole community scale. These ecological relationships not only demonstrate close links between above and below‐ground plant carbon dynamics but also allow plant carbon pool sizes and their turnover rates to be predicted from the single readily quantifiable (and remotely sensed) parameter of LAI, including the possibility of estimating root data from satellites.
The Arctic is experiencing the greatest climate change in winter, including increases in freeze–thaw cycles that can result in ice encasement of vegetation. Ice encasement can expose plants to ...hypoxia and greater temperature extremes, but currently the impacts of icing on plants in the field remain little understood. With this in mind, a unique field manipulation experiment was established in heathland in northern Sweden with ice encasement simulated in early March 2008, 2009 and 2010 until natural thaw each spring. In the following summers we assessed the impacts on flowering, bud phenology, shoot growth and mortality and leaf damage (measured by chlorophyll fluorescence and electrolyte leakage) of the three dominant dwarf shrub species Empetrum nigrum, Vaccinium vitis‐idaea (both evergreen) and Vaccinium myrtillus (deciduous). Two consecutive winters of icing decreased V. vitis‐idaea flowering by 57%, while flowering of V. myrtillus and E. nigrum remained unaffected. Vaccinium myrtillus showed earlier budburst but shoot growth for all species was unchanged. Shoot mortality of V. myrtillus and V. vitis‐idaea increased after the first year (by 70 and 165%, respectively) and again for V. myrtillus following the third year (by 67%), while E. nigrum shoot mortality remained unaffected, as were chlorophyll fluorescence and electrolyte leakage in all species. Overall, the sub‐arctic heathland was relatively tolerant to icing, but the considerable shoot mortality of V. myrtillus contrasting with the general tolerance of E. nigrum suggests plant community structure in the longer term could change if winters continue to see a greater frequency of icing events.
Potential land–climate feedbacks in subarctic regions, where rapid warming is driving forest expansion into the tundra, may be mediated by differences in transpiration of different plant functional ...types. Here, we assess the environmental controls of overstorey transpiration and its relevance for ecosystem evapotranspiration in subarctic deciduous woodlands. We measured overstorey transpiration of mountain birch canopies and ecosystem evapotranspiration in two locations in northern Fennoscandia, having dense (Abisko) and sparse (Kevo) overstories. For Kevo, we also upscale chamber‐measured understorey evapotranspiration from shrubs and lichen using a detailed land cover map. Subdaily evaporative fluxes were not affected by soil moisture and showed similar controls by vapour pressure deficit and radiation across sites. At the daily timescale, increases in evaporative demand led to proportionally higher contributions of overstorey transpiration to ecosystem evapotranspiration. For the entire growing season, the overstorey transpired 33% of ecosystem evapotranspiration in Abisko and only 16% in Kevo. At this latter site, the understorey had a higher leaf area index and contributed more to ecosystem evapotranspiration compared with the overstorey birch canopy. In Abisko, growing season evapotranspiration was 27% higher than precipitation, consistent with a gradual soil moisture depletion over the summer. Our results show that overstorey canopy transpiration in subarctic deciduous woodlands is not the dominant evaporative flux. However, given the observed environmental sensitivity of evapotranspiration components, the role of deciduous trees in driving ecosystem evapotranspiration may increase with the predicted increases in tree cover and evaporative demand across subarctic regions.
Summary
Winter is a period of dormancy for plants of cold environments. However, winter climate is changing, leading to an increasing frequency of stochastic warm periods (winter warming events) and ...concomitant reductions in snow cover. These conditions can break dormancy for some plants and expose them to freeze‐and‐thaw stress. Mosses are a major component of high‐latitude ecosystems, yet the longer‐term impacts of such winter warming events on mosses remain unknown.
In order to determine the longer‐term legacy effects of winter warming events on mosses, we undertook a simulation of these events over three consecutive winters in a sub‐Arctic dwarf shrub‐dominated open woodland. The mat‐forming feather moss, Hylocomium splendens (the most abundant cryptogam in this system), is one of the most widespread Arctic and boreal mosses and plays a key functional role in ecosystems. We studied the ecophysiological performance of this moss during the summers of the experimental period (2007–2009) and in the following years (2010–2013).
We show that the previously reported warming‐induced reduction in segment growth and photosynthesis during the experimental years was persistent. Four years after the last event, photosynthesis and segment growth were still 30 and 36% lower than control levels, which was only a slight improvement from 44 and 43% 4 years earlier. Winter warming did not affect segment symmetry. During the years after the last simulated event, in both warmed and control plots, chlorophyll fluorescence and segment growth, but not net photosynthesis, increased slightly. The increases were probably driven by increased summer rainfall over the study years, highlighting the sensitivity of this moss to rainfall change.
Overall, the legacy effects shown here demonstrate that this widespread and important moss is likely to be significantly disadvantaged in a future sub‐Arctic climate where frequent winter warming events may become the norm. Given the key importance of mosses for soil insulation, shelter and carbon sequestration in high‐latitude regions, such persistent impacts may ultimately affect important ecosystem functions.
A Lay Summary is available for this article.
Lay Summary
Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of ...herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub‐Arctic heath vegetation and its belowground micro‐arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro‐arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO2 fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub‐Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub‐Arctic vegetation response to multiple pressures is not easy to predict from single‐factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.
Atmospheric nitrogen deposition can cause major declines in bryophyte abundance yet the physiological basis for such declines is not fully understood. Bryophyte physiological responses may also be ...sensitive bioindicators of both the impacts of, and recovery from, N deposition. Here, responses of tissue nutrients (nitrogen (N), phosphorus (P) and potassium (K): NPK), N and P metabolism enzymes (nitrate reductase and phosphomonoesterase), photosynthetic pigments, chlorophyll fluorescence, sclerophylly and percentage cover of two common bryophytes (Pseudoscleropodium purum and Rhytidiadelphus squarrosus) to long-term (11 yr) enhanced N deposition (+3.5 and +14 g N m⁻² yr⁻¹) are reported in factorial combination with P addition. Recovery of responses 22 months after treatment cessation were also assessed. Enhanced N deposition caused up to 90% loss of bryophyte cover but no recovery was observed. Phosphomonoesterase activity and tissue N : P ratios increased up to threefold in response to N loading and showed clear recovery, particularly in P. purum. Smaller responses and recovery were also seen in all chlorophyll fluorescence measurements and altered photosynthetic pigment composition. The P limitation of growth appears to be a key mechanism driving bryophyte loss along with damage to photosystem II. Physiological measurements are more sensitive than measurements of abundance as bioindicators of N deposition impact and of recovery in particular.
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