Impact of early and late winter icing events on sub‐arctic dwarf shrubs

Polar regions are predicted to undergo large increases in winter temperature and an increased frequency of freeze–thaw cycles, which can cause ice layers in the snow pack and ice encasement of vegetation. Early or late winter timing of ice encasement could, however, modify the extent of damage caused to plants. To determine impacts of the date of ice encasement, a novel field experiment was established in sub‐arctic Sweden, with icing events simulated in January and March 2008 and 2009. In the subsequent summers, reproduction, phenology, growth and mortality, as well as physiological indicators of leaf damage were measured in the three dominant dwarf shrubs: Vaccinium uliginosum, Vaccinium vitis‐idaea and Empetrum nigrum. It was hypothesised that January icing would be more damaging compared to March icing due to the longer duration of ice encasement. Following 2 years of icing, E. nigrum berry production was 83% lower in January‐iced plots compared to controls, and V. vitis‐idaea electrolyte leakage was increased by 69%. Conversely, electrolyte leakage of E. nigrum was 25% lower and leaf emergence of V. vitis‐idaea commenced 11 days earlier in March‐iced plots compared to control plots in 2009. There was no effect of icing on any of the other parameters measured, indicating that overall these study species have moderate to high tolerance to ice encasement. Even much longer exposure under the January icing treatment does not clearly increase damage.     

Impacts of multiple extreme winter warming events on sub‐Arctic heathland: phenology,reproduction, growth,and CO2 flux responses

Extreme weather events can have strong negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme, short‐lived, winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (for instance, 2–10 °C for 2–14 days) but upon return to normal winter climate exposes the ecosystem to much colder temperatures due to the loss of insulating snow. Single events have been shown to reduce plant reproduction and increase shoot mortality, but impacts of multiple events are little understood as are the broader impacts on community structure, growth, carbon balance, and nutrient cycling. To address these issues, we simulated week‐long extreme winter warming events – using infrared heating lamps and soil warming cables – for 3 consecutive years in a sub‐Arctic heathland dominated by the dwarf shrubs Empetrum hermaphroditum, Vaccinium vitis‐idaea (both evergreen) and Vaccinium myrtillus (deciduous). During the growing seasons after the second and third winter event, spring bud burst was delayed by up to a week for E. hermaphroditum and V. myrtillus, and berry production reduced by 11–75% and 52–95% for E. hermaphroditum and V. myrtillus, respectively. Greater shoot mortality occurred in E. hermaphroditum (up to 52%), V. vitis‐idaea (51%), and V. myrtillus (80%). Root growth was reduced by more than 25% but soil nutrient availability remained unaffected. Gross primary productivity was reduced by more than 50% in the summer following the third simulation. Overall, the extent of damage was considerable, and critically plant responses were opposite in direction to the increased growth seen in long‐term summer warming simulations and the ‘greening’ seen for some arctic regions. Given the Arctic is warming more in winter than summer, and extreme events are predicted to become more frequent, this generates large uncertainty in our current understanding of arctic ecosystem responses to climate change.     

Impacts of extreme winter warming events on plant physiology in a sub‐Arctic heath community

Insulation provided by snow cover and tolerance of freezing by physiological acclimation allows Arctic plants to survive cold winter temperatures. However, both the protection mechanisms may be lost with winter climate change, especially during extreme winter warming events where loss of snow cover from snow melt results in exposure of plants to warm temperatures and then returning extreme cold in the absence of insulating snow. These events cause considerable damage to Arctic plants, but physiological responses behind such damage remain unknown. Here, we report simulations of extreme winter warming events using infrared heating lamps and soil warming cables in a sub‐Arctic heathland. During these events, we measured maximum quantum yield of photosystem II (PSII), photosynthesis, respiration, bud swelling and associated bud carbohydrate changes and lipid peroxidation to identify physiological responses during and after the winter warming events in three dwarf shrub species: Empetrum hermaphroditum, Vaccinium vitis‐idaea and Vaccinium myrtillus. Winter warming increased maximum quantum yield of PSII, and photosynthesis was initiated for E. hermaphroditum and V. vitis‐idaea. Bud swelling, bud carbohydrate decreases and lipid peroxidation were largest for E. hermaphroditum, whereas V. myrtillus and V. vitis‐idaea showed no or less strong responses. Increased physiological activity and bud swelling suggest that sub‐Arctic plants can initiate spring‐like development in response to a short winter warming event. Lipid peroxidation suggests that plants experience increased winter stress. The observed differences between species in physiological responses are broadly consistent with interspecific differences in damage seen in previous studies, with E. hermaphroditum and V. myrtillus tending to be most sensitive. This suggests that initiation of spring‐like development may be a major driver in the damage caused by winter warming events that are predicted to become more frequent in some regions of the Arctic and that may ultimately drive plant community shifts.     

Impacts of extreme winter warming in the sub‐Arctic: growing season responses of dwarf shrub heathland

Climate change scenarios predict an increased frequency of extreme climatic events. In Arctic regions, one of the most profound of these are extreme and sudden winter warming events in which temperatures increase rapidly to above freezing, often causing snow melt across whole landscapes and exposure of ecosystems to warm temperatures. Following warming, vegetation and soils no longer insulated below snow are then exposed to rapidly returning extreme cold. Using a new experimental facility established in sub‐Arctic dwarf shrub heathland in northern Sweden, we simulated an extreme winter warming event in the field and report findings on growth, phenology and reproduction during the subsequent growing season. A 1‐week long extreme winter warming event was simulated in early March using infrared heating lamps run with or without soil warming cables. Both single short events delayed bud development of Vaccinium myrtillus by up to 3 weeks in the following spring (June) and reduced flower production by more than 80%: this also led to a near‐complete elimination of berry production in mid‐summer. Empetrum hermaphroditum also showed delayed bud development. In contrast, Vaccinium vitis‐idaea showed no delay in bud development, but instead appeared to produce a greater number of actively growing vegetative buds within plots warmed by heating lamps only. Again, there was evidence of reduced flowering and berry production in this species. While bud break was delayed, growing season measurements of growth and photosynthesis did not reveal a differential response in the warmed plants for any of the species. These results demonstrate that a single, short, extreme winter warming event can have considerable impact on bud production, phenology and reproductive effort of dominant plant species within sub‐Arctic dwarf shrub heathland. Furthermore, large interspecific differences in sensitivity are seen. These findings are of considerable concern, because they suggest that repeated events may potentially impact on the biodiversity and productivity of these systems should these extreme events increase in frequency as a result of global change. Although climate change may lengthen the growing season by earlier spring snow melt, there is a profound danger for these high‐latitude ecosystems if extreme, short‐lived warming in winter exposes plants to initial warm temperatures, but then extreme cold for the rest of the winter. Work is ongoing to determine the longer term and wider impacts of these events.     

Spatial distribution of aquatic marine fungi across the western Arctic and sub‐arctic

Fungi are important parasites of primary producers and nutrient cyclers in aquatic ecosystems. In the Pacific‐Arctic domain, fungal parasitism is linked to light intensities and algal stress that can elevate disease incidence on algae and reduce diatom concentrations. Fungi are vastly understudied in the marine realm and knowledge of their function is constrained by the current understanding of fungal distribution and drivers on global scales. To investigate the spatial distribution of fungi in the western Arctic and sub‐Arctic, we used high throughput methods to sequence 18S rRNA, cloned and sequenced 28S rRNA and microscopically counted chytrid‐infected diatoms. We identified a broad distribution of fungal taxa predominated by Chytridiomycota and Dikarya. Phylogenetic analysis of our Chytridiomycota clones placed Arctic marine fungi sister to the order Lobulomycetales. This clade of fungi predominated in fungal communities under ice with low snowpack. Microscopic examination of fixed seawater and sea ice samples revealed chytrids parasitizing diatoms collected across the Arctic that notably infected 25% of a single diatom species in the Bering Sea. The Pezizomycotina comprised > 95% of eukaryotic sequence reads in Greenland, providing preliminary evidence for osmotrophs being a substitute for algae as the base of food webs.     

Contrasting seasonal overlaps between primary and secondary growth are linked to wood anatomy in Mediterranean sub‐shrubs

Whole‐plant approaches allow quantification of the temporal overlap between primary and secondary growth. If the amount of time available to grow is short, there may be a high temporal overlap between shoot growth and wood formation. We hypothesise that such overlap depends on the duration of the growing season and relates to wood anatomy. We evaluated wood anatomy, shoot longitudinal and radial growth rates, fine root production and the concentrations of non‐structural carbohydrates (NSC) in the wood of six sub‐shrub species growing in sites with contrasting climatic conditions (Lepidium subulatum, Linum suffruticosum, Salvia lavandulifolia, Satureja montana, Ononis fruticosa, Echinospartum horridum). Sub‐shrub species living in sites with a short growing season displayed a high overlap between aboveground primary and secondary growth and formed wide vessels, whereas species from the warmest and driest sites presented the reverse characteristics. The highest overlap was linked to a rapid shoot extension and thickening through the enhanced hydraulic conductivity provided by wide vessels. The reductions in NSC concentrations when growth peaked were low or moderate, indicating that sub‐shrubs accumulate NSC in excess, as do trees. The temporal overlap among primary and secondary growth in woody plants may be connected to the duration and rates of shoot and wood growth, which in turn depend on the vessel lumen area.     

Experimental icing affects growth,mortality, and flowering in a high Arctic dwarf shrub

Effects of climate change are predicted to be greatest at high latitudes, with more pronounced warming in winter than summer. Extreme mid‐winter warm spells and heavy rain‐on‐snow events are already increasing in frequency in the Arctic, with implications for snow‐pack and ground‐ice formation. These may in turn affect key components of Arctic ecosystems. However, the fitness consequences of extreme winter weather events for tundra plants are not well understood, especially in the high Arctic. We simulated an extreme mid‐winter rain‐on‐snow event at a field site in high Arctic Svalbard (78°N) by experimentally encasing tundra vegetation in ice. After the subsequent growing season, we measured the effects of icing on growth and fitness indices in the common tundra plant, Arctic bell‐heather (Cassiope tetragona). The suitability of this species for retrospective growth analysis enabled us to compare shoot growth in pre and postmanipulation years in icing treatment and control plants, as well as shoot survival and flowering. Plants from icing treatment plots had higher shoot mortality and lower flowering success than controls. At the individual sample level, heavily flowering plants invested less in shoot growth than nonflowering plants, while shoot growth was positively related to the degree of shoot mortality. Therefore, contrary to expectation, undamaged shoots showed enhanced growth in ice treatment plants. This suggests that following damage, aboveground resources were allocated to the few remaining undamaged meristems. The enhanced shoot growth measured in our icing treatment plants has implications for climate studies based on retrospective analyses of Cassiope. As shoot growth in this species responds positively to summer warming, it also highlights a potentially complex interaction between summer and winter conditions. By documenting strong effects of icing on growth and reproduction of a widespread tundra plant, our study contributes to an understanding of Arctic plant responses to projected changes in winter climatic conditions.     

Regeneration by seeds in alpine meadow and heath vegetation in sub‐arctic Finland

Abstract. This paper compares the regeneration by seeds of heath and meadow and studies relationships between the floristic composition of phases in the regeneration pathway. Seed densities in the seed rain and seed bank as well as the densities of emerged seedlings in gaps and in closed vegetation were greater in the meadow than in the heath. In the heath, environmental constraints hindered seedling emergence almost completely so seeds accumulated in the seed bank. In the meadow, the decrease in the seed bank was due to high seedling emergence. Within both plant communities, seedling emergence in gaps and in closed vegetation was comparable. In the meadow, the seed rain and seedling emergence in gaps, as well as the seed bank and seedling emergence in gaps were positively correlated. Differences in seed and adult plant sizes were reasons for the low correlation between the standing vegetation and the other phases. In DCA ordination the first axis separated the phase of seedling emergence in closed vegetation and seed bank. The second axis separated the standing vegetation from the other phases. The structure of the seed rain was more heterogeneous than that of other phases. In the heath, the standing vegetation and the seed rain were positively correlated. The ordination of these phases reflected the patchiness of standing vegetation and the ability of the diaspores of Betula nana to disperse over long distances.     

Impacts of “supermoon” events on the physiology of a wild bird

The position of the Moon in relation to the Earth and the Sun gives rise to several predictable cycles, and natural changes in nighttime light intensity are known to cause alterations to physiological processes and behaviors in many animals. The limited research undertaken to date on the physiological responses of animals to the lunar illumination has exclusively focused on the synodic lunar cycle (full moon to full moon, or moon phase) but the moon's orbit—its distance from the Earth—may also be relevant. Every month, the moon moves from apogee, its most distant point from Earth—and then to perigee, its closest point to Earth. Here, we studied wild barnacle geese (Branta leucopsis) to investigate the influence of multiple interacting lunar cycles on the physiology of diurnally active animals. Our study, which uses biologging technology to continually monitor body temperature and heart rate for an entire annual cycle, asks whether there is evidence for a physiological response to natural cycles in lunar brightness in wild birds, particularly “supermoon” phenomena, where perigee coincides with a full moon. There was a three‐way interaction between lunar phase, lunar distance, and cloud cover as predictors of nighttime mean body temperature, such that body temperature was highest on clear nights when the full moon coincided with perigee moon. Our study is the first to report the physiological responses of wild birds to “supermoon” events; the wild geese responded to the combination of two independent lunar cycles, by significantly increasing their body temperature at night. That wild birds respond to natural fluctuations in nighttime ambient light levels support the documented responses of many species to anthropogenic sources of artificial light, that birds seem unable to override. As most biological systems are arguably organized foremost by light, this suggests that any interactions between lunar cycles and local weather conditions could have significant impacts on the energy budgets of birds.     

Plant traits: Their role in the regeneration of alpine plant communities in sub‐arctic Finland

Abstract. Question: How do the relative frequencies of plant traits (clonality, growth form, seed weight, diaspore morphology) vary during the life cycle and how does this affect regeneration? Location: Alpine meadow and heath communities at Kilpisjärvi, sub‐Arctic Finland. Methods: Control plots and three treatments were used to measure relative species abundances for five life cycle stages: standing vegetation, seed rain, seed bank and seedlings emerging in gaps and in closed vegetation. Results: The relative frequencies of plant traits varied between the life cycle stages. The meadows were dominated by weakly clonal herbs, small or intermediate seeds and unappendaged diaspores, while the heaths were dominated by clonal dwarf shrubs, small seeds and fleshy fruits. In the meadows, species with small seeds dominated during the seed rain and in the seedling stage in gaps, while species with intermediate seeds dominated the seed bank and the seedling stage in closed vegetation. Species with unappendaged diaspores dominated throughout the life cycle. In the heaths, seed bank and seedling stage were practically absent. Conclusions: The observed differences in plant trait spectra between life cycle stages indicate that important environmental factors differ among the stages. Small seeds are advantageous for dispersal, whereas intermediate seeds have a greater probability of germinating and establishing in closed vegetation. Appendages facilitate dispersal, whereas unappendaged diaspores favour seed burial. Although the plant growth form spectrum largely reflects environmental constraints during the regeneration cycle, information on seed weight and diaspore morphology improves our knowledge of the relative importance of morphological adaptations of sexual structures in different stages during the life cycle.     

Effects of temperature and date of snowmelt on growth, reproduction, and flowering phenology in the arctic/alpine herb, Ranunculus glacialis

Changes in growing season temperature and duration may have profound effects on the population dynamics of arctic and alpine plant species in snow-bed and fell-field habitats. We examined how a typical herbaceous pioneer species, Ranunculus glacialis, responded to experimental climate change in open-top chambers for three seasons at an alpine site in southern Norway. Warming had no significant effect on any reproductive, growth or phenological variables, except for seed weight, which increased significantly during the first 2 ears. Despite large differences in average date of snowmelt among years, average reproductive output and ramet size differed little among years. Within-year variation in date of snowmelt had no impact on seed number or weight in either control or warmed plots. Leaf width and ramet leaf number decreased significantly with later snowmelt within a year. Experimental warming reduced the negative effect on ramet size of late snowmelt within a year to some extent. In general, R. glacialis reacts contrary to most other arctic/alpine species to experimental warming. Species with such low responsiveness to environmental conditions may be particularly vulnerable to climatic change, especially if their habitat is invaded by other species with higher phenotypic plasticity and a better competitive ability.     

Data logging of body temperatures provides precise information on phenology of reproductive events in a free-living arctic hibernator

Precise measures of phenology are critical to understanding how animals organize their annual cycles and how individuals and populations respond to climate-induced changes in physical and ecological stressors. We show that patterns of core body temperature (T _b) can be used to precisely determine the timing of key seasonal events including hibernation, mating and parturition, and immergence and emergence from the hibernacula in free-living arctic ground squirrels (Urocitellus parryii). Using temperature loggers that recorded T _b every 20 min for up to 18 months, we monitored core T _b from three females that subsequently gave birth in captivity and from 66 female and 57 male ground squirrels free-living in the northern foothills of the Brooks Range Alaska. In addition, dates of emergence from hibernation were visually confirmed for four free-living male squirrels. Average T _b in captive females decreased by 0.5–1.0°C during gestation and abruptly increased by 1–1.5°C on the day of parturition. In free-living females, similar shifts in T _b were observed in 78% (n = 9) of yearlings and 94% (n = 31) of adults; females without the shift are assumed not to have given birth. Three of four ground squirrels for which dates of emergence from hibernation were visually confirmed did not exhibit obvious diurnal rhythms in T _b until they first emerged onto the surface when T _b patterns became diurnal. In free-living males undergoing reproductive maturation, this pre-emergence euthermic interval averaged 20.4 days (n = 56). T _b-loggers represent a cost-effective and logistically feasible method to precisely investigate the phenology of reproduction and hibernation in ground squirrels.     

Impacts of bud set and lammas phenology on root:shoot biomass partitioning and carbon gain physiology in poplar

Key message

Timing of bud set and occurrence of lammas in trees can alter growth partitioning (i.e., root:shoot ratios), while only bud set effectively modifies carbon gain by increasing photosynthesis-related physiological traits.

Abstract

Bud set and lammas (second bud flushing) phenology may strongly influence growth, physiology, and biomass in trees. To test effects of these phenological events, 54 individuals from 16 genotypes of black cottonwood poplar (Populus trichocarpa) were grown in a potted trial under greenhouse conditions (with extended daylengths promoting growth), followed by open-air cultivation (with natural daylengths promoting bud set and/or lammas). Trees were monitored for phenology, repeatedly measured for photosynthesis-related traits, harvested for biomass, and assessed for growth partitioning (separating above- and belowground parts). We grouped trees by phenology for comparisons: (1) trees with early summer bud set, (2) trees with early summer bud set that underwent lammas, (3) trees with late summer bud set (August), and (4) trees with bud set occurring in autumn (September). We found that bud set timing positively affected growth partitioning where earlier bud set resulted in shorter trees with higher root:shoot biomass ratios (by increasing root mass). Lammas growth altered these ratios by significantly increasing shoot growth relative to belowground growth. Trees with bud set occurring in late summer also had higher root:shoot biomass ratios (by increasing root mass) compared to trees setting bud in autumn. Occurrence of bud set coincided with modified physiology of the existing canopy where photosynthesis-related traits were enhanced relative to trees still actively growing. These physiological changes were unaltered by occurrence of lammas. This suggests that bud set prompts a significant, coordinated mechanism of higher carbon gain physiology and belowground biomass accumulation in plants within a “post-bud set” phase.

     

Effects of growth temperature and winter duration on leaf phenology of Erythronium japonicum,a forest spring geophyte

The effects of growth temperature and winter duration on the leaf phenology of Erythronium japonicum were examined in two experiments. Bulbs wintered in the field were cultivated at 10 and 20° C and the bulbs were cultivated at 15° C after chilling treatment at 3° C for 60 and 120 days and without chilling in winter. The plants cultivated at 20° C showed significantly earlier leaf emergence, a more rapid rate of leaf extension and shorter leaf longevity than those cultivated at 10° C. The decrease in the leaf longevity at 20° C resulted from the decreases in the durations of all of the developmental, mature, and senescent phases. The bulbs without chilling treatment did not sprout leaves and those with chilling treatment sprouted leaves. The increase in the length of chilling treatment from 60 to 120 days affected leaf phenology in same manner as the increase in the growth temperature from 10 to 20° C.     

Extreme seasonality of litter breakdown in an arctic spring‐fed stream is driven by shredder phenology,not temperature

1. Using degree‐days to calculate ‘temperature‐corrected’ breakdown rates is a useful approach for comparing litter breakdown across sites with different thermal regimes. We used an alternative approach to investigate the importance of temperature by quantifying seasonal patterns in litter breakdown in an arctic spring‐fed stream (Ivishak Spring, North Slope, Alaska) that experiences extreme seasonality in light availability and energy inputs while fluctuations in water temperature are relatively small. 2. We incubated mesh bags of senesced Salix alaxensis litter in Ivishak Spring for successive c. 30‐day periods for 2 years. During our study, water temperature was very stable [5.7 ± 0.03 °C (daily mean ± 1 SE), range 3.7–7.8 °C]. Discharge was only slightly more variable (mean 112 ± 1 L s^?1, range 66–206 L s^?1), with lowest values occurring in late winter. Dissolved nutrient concentrations were low (52–133 μg L^?1, <1–3 μg L^?1, <1–6 μg L^?1 soluble reactive phosphorus) and also showed evidence of seasonality (i.e. highest values in winter). 3. Litter breakdown rates were sharply seasonal, ranging from <0.01 day^?1 in mid‐summer to >0.05 day^?1 in mid‐winter. Invertebrate assemblage structure in litter bags showed pronounced seasonal cyclicity; total invertebrate biomass peaked in summer. Biomass of two dominant shredders (the nemourid stonefly Zapada haysi and the limnephilid caddisfly Ecclisomyia conspersa) showed the opposite trend, however, with mid‐winter peaks in both population biomass and cohort growth rates that closely matched those we observed in litter mass loss. 4. Water temperature appeared to have negligible influence on litter breakdown rates in our study. Seasonal shifts in nutrient uptake may have increased rates of microbial activity in winter. The processing of litter inputs in Ivishak Spring, however, appeared to be most tightly coupled to shredder phenology. Our results demonstrate that extreme seasonality in the processing of allochthonous detritus can occur even in the absence of substantial temperature variation, driven by the activity of shredder taxa that have evolved to take advantage of pulsed organic matter inputs.     

Compensatory growth of two clonal dwarf shrubs, Arctostaphylos uva-ursi and Vaccinium uliginosum in a heavy metal polluted environment

The effect of artificial shoot clipping on the vegetative growth and sexual reproduction of the evergreen bearberry, Arctostaphylos uva-ursi, and the deciduous bog bilberry, Vaccinium uliginosum, was studied in the vicinity of a copper-nickel smelter in SW Finland. According to the research hypothesis, heavy metal induced shoot death breaks the apical dominance in the clones growing in a polluted environment. This causes activation of dormant axillary and adventitious buds and an increase in branching on the older parts of the stem. Regrowth after shoot death was studied by clipping off all the current-year shoots from experimental branches in autumn (1994) and spring (1995). Within-clone and between-clone control branches were used to test the data.Both species displayed a considerable ability to activate dormant meristems after the damage. Regrowth of the current shoots during the next growing season (1995) was about 80% compared to the within-clone control in both species after autumn clipping. Shoot clipping in early summer was more detrimental for both species, and the regrowth of A. uva-ursi was less than that of V. uliginosum. Differences in the storage reserves and source-sink mechanisms of carbon allocation between evergreen and deciduous species probably explain their distinct response. When the removed biomass was added to the living biomass of the branches, there was overcompensation in the total dry weight of A. uva-ursi after autumn clipping, and the weight was almost 90% of the control after spring clipping. The total dry weight of V. uliginosum also equalled that of the control when the removed biomass was added. No berries developed on either species in the year following the autumn treatment, because clipping removed all the flower buds. Spring clipping had no effect on the sexual reproduction of A. uva-ursi, but decreased the berry production of V. uliginosum. The degree of compensatory growth of both species was only slightly affected by the distance from the smelter. It is suggested that dormant bud activation, rapid regrowth and plastic branching contribute to the resistance mechanism to heavy metals.