Flower and seed biomass, and germination rate of arctic tundra plants in response to long term experimental warming

We provide new information on changes in tundra plant sexual reproduction in response to long-term (12 years) experimental warming in the High Arctic. Open-top chambers (OTCs) were used to increase growing season temperatures by 1-2 °C across a range of vascular plant communities. The warming enhanced reproductive effort and success in most species; shrubs and graminoids appeared to be more responsive than forbs. We found that the measured effects of warming on sexual reproduction were more consistently positive and to a greater degree in polar oasis compared with polar semidesert vascular plant communities. Our findings support predictions that long-term warming in the High Arctic will likely enhance sexual reproduction in tundra plants, which could lead to an increase in plant cover. Greater abundance of vegetation has implications for primary consumers - via increased forage availability, and the global carbon budget - as a function of changes in permafrost and vegetation acting as a carbon sink. Enhanced sexual reproduction in Arctic vascular plants may lead to increased genetic variability of offspring, and consequently improved chances of survival in a changing environment. Our findings also indicate that with future warming, polar oases may play an important role as a seed source to the surrounding polar desert landscape.

(Table 1) Effects of experimental warming on plant cover and diversity indices in an evergreen shrub at Alexandra Fiord

1. Identifying plant communities that are resistant to climate change will be critical for developing accurate, wide-scale vegetation change predictions. Most northern plant communities, especially tundra, have shown strong responses to experimental and observed warming. 2. Experimental warming is a key tool for understanding vegetation responses to climate change. We used open-top chambers to passively warm an evergreen-shrub heath by 1.0-1.3 °C for 15 years at Alexandra Fiord, Nunavut, Canada (79 °N). In 1996, 2000 and 2007, we measured height, plant composition and abundance with a point-intercept method. 3. Experimental warming did not strongly affect vascular plant cover, canopy height or species diversity, but it did increase bryophyte cover by 6.3% and decrease lichen cover by 3.5%. Temporal changes in plant cover were more frequent and of greater magnitude than changes due to experimental warming. 4. Synthesis. This evergreen-shrub heath continues to exhibit community-level resistance to long-term experimental warming, in contrast to most Arctic plant communities. Our findings support the view that only substantial climatic changes will alter unproductive ecosystems.

Response of three krill species to hypoxia and warming: An experimental approach to oxygen minimum zones expansion in coastal ecosystems

To understand the adaptation of euphausiid (krill) species to oxygen minimum zones (OMZ), respiratory response and stress experiments combining hypoxia/reoxygenation exposure with warming were conducted. Experimental krill species were obtained from the Antarctic (South Georgia area), the Humboldt Current system (HCS, Chilean coast), and the Northern California Current system (NCCS, Oregon). Euphausia mucronata from the HCS shows oxyconforming or oxygen partial pressure (pO2)-dependent respiration below 80% air saturation (18 kPa). Normoxic subsurface oxygenation in winter posed a "high oxygen stress" for this species. The NCCS krill, Euphausia pacifica, and the Antarctic krill, Euphausia superba maintain respiration rates constant down to low critical pO2 values of 6 kPa (30% air saturation) and 11 kPa (55% air saturation), respectively. Antarctic krill had the lowest antioxidant enzyme activities, but the highest concentrations of the molecular antioxidant glutathione (GSH) and was not affected by 6 h exposure to moderate hypoxia. Temperate krill species had higher SOD (superoxide dismutase) values in winter than in summer, which relate to higher winter metabolic rate (E. pacifica). In all species, antioxidant enzyme activities remained constant during hypoxic exposure at habitat temperature. Warming by 7°C above habitat temperature in summer increased SOD activities and GSH levels in E. mucronata (HCS), but no oxidative damage occurred. In winter, when the NCCS is well mixed and the OMZ is deeper, +4°C of warming combined with hypoxia represents a lethal condition for E. pacifica. In summer, when the OMZ expands upwards (100 m subsurface), antioxidant defences counteracted hypoxia and reoxygenation effects in E. pacifica, but only at mildly elevated temperature (+2°C). In this season, experimental warming by +4°C reduced antioxidant activities and the hypoxia combination again caused mortality of exposed specimens. We conclude that a climate change scenario combining warming and hypoxia represents a serious threat to E. pacifica and, as a consequence, NCCS food webs.

Initial soil properties and biomass after experimental grazing and warming in mesic and wet sites, Adventdalen valley, Spitzbergen

High-latitude ecosystems store large amounts of carbon (C); however, the C storage of these ecosystems is under threat from both climate warming and increased levels of herbivory. In this study we examined the combined role of herbivores and climate warming as. drivers of CO2 fluxes in two typical high-latitude habitats (mesic heath and wet meadow). We hypothesized that both herbivory and climate warming would reduce the C sink strength of Arctic tundra through their combined effects on plant biomass and gross ecosystem photosynthesis and on decomposition rates and the abiotic environment. To test this hypothesis we employed experimental warming (via International Tundra Experiment [ITEX] chambers) and grazing (via captive Barnacle Geese) in a three-year factorial field experiment. Ecosystem CO2 fluxes (net ecosystem exchange of CO2, ecosystem respiration, and gross ecosystem photosynthesis) were measured in all treatments at varying intensity over the three growing seasons to capture the impact of the treatments on a range of temporal scales (diurnal, seasonal, and interannual). Grazing and warming treatments had markedly different effects on CO2 fluxes in the two tundra habitats. Grazing caused a strong reduction in CO2 assimilation in the wet meadow, while warming reduced CO2 efflux from the mesic heath. Treatment effects on net ecosystem exchange largely derived from the modification of gross ecosystem photosynthesis rather than ecosystem respiration. In this study we have demonstrated that on the habitat scale, grazing by geese is a strong driver of net ecosystem exchange of CO2, with the potential to reduce the CO2 sink strength of Arctic ecosystems. Our results highlight that the large reduction in plant biomass due to goose grazing in the Arctic noted in several studies can alter the C balance of wet tundra ecosystems. We conclude that herbivory will modulate direct climate warming responses of Arctic tundra with implications for the ecosystem C balance; however, the magnitude and direction of the response will be habitat-specific.

Environmental characteristics, and growth traits and leaf chemistry of tundra plants in a warming experiment at Alexandra Fiord

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate-vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open-top chambers (OTCs) to passively warm vegetation by 1-2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf d15N. Long-term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect d15N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.

Climate and soil characteristics and soil arthropod abundance on Anchorage, Signy and Falkland Islands

Over a 2-year study, we investigated the effect of environmental change on the diversity and abundance of soil arthropod communities (Acari and Collembola) in the Maritime Antarctic and the Falkland Islands. Open Top Chambers (OTCs), as used extensively in the framework of the northern boreal International Tundra Experiment (ITEX), were used to increase the temperature in contrasting communities on three islands along a latitudinal temperature gradient, ranging from the Falkland Islands (51°S, mean annual temperature 7.5 °C) to Signy Island (60°S, -2.3°C) and Anchorage Island (67°S, -3.8°C). At each island an open and a closed plant community were studied: lichen vs. moss at the Antarctic sites, and grass vs. dwarf shrub at the Falkland Islands. The OTCs raised the soil surface temperature during most months of the year. During the summer the level of warming achieved was 1.7 °C at the Falkland Islands, 0.7 °C at Signy Island, and 1.1 °C at Anchorage Island. The native arthropod community diversity decreased with increasing latitude. In contrast with this pattern, Collembola abundance in the closed vegetation (dwarf shrub or moss) communities increased by at least an order of magnitude from the Falkland Islands (9.0 +/- 2 x 10**3 ind./m**2) to Signy (3.3 +/- 8.0 x 10**4 ind./m**2) and Anchorage Island (3.1 +/- 0.82 x 10**5 ind./m**2). The abundance of Acari did not show a latitudinal trend. Abundance and diversity of Acari and Collembola were unaffected by the warming treatment on the Falkland Islands and Anchorage Island. However, after two seasons of experimental warming, the total abundance of Collembola decreased (p < 0.05) in the lichen community on Signy Island as a result of the population decline of the isotomid Cryptopygus antarcticus. In the same lichen community there was also a decline (p < 0.05) of the mesostigmatid predatory mite Gamasellus racovitzai, and a significant increase in the total number of Prostigmata. Overall, our data suggest that the consequences of an experimental temperature increase of 1-2°C, comparable to the magnitude currently seen through recent climate change in the Antarctic Peninsula region, on soil arthropod communities in this region may not be similar for each location but is most likely to be small and initially slow to develop.

Soil characteristics and decomposition of organic matter on Anchorage, Signy and Falkland Islands

Antarctic terrestrial ecosystems have poorly developed soils and currently experience one of the greatest rates of climate warming on the globe. We investigated the responsiveness of organic matter decomposition in Maritime Antarctic terrestrial ecosystems to climate change, using two study sites in the Antarctic Peninsula region (Anchorage Island, 67°S; Signy Island, 61°S), and contrasted the responses found with those at the cool temperate Falkland Islands (52°S). Our approach consisted of two complementary methods: (1) Laboratory measurements of decomposition at different temperatures (2, 6 and 10 °C) of plant material and soil organic matter from all three locations. (2) Field measurements at all three locations on the decomposition of soil organic matter, plant material and cellulose, both under natural conditions and under experimental warming (about 0.8 °C) achieved using open top chambers. Higher temperatures led to higher organic matter breakdown in the laboratory studies, indicating that decomposition in Maritime Antarctic terrestrial ecosystems is likely to increase with increasing soil temperatures. However, both laboratory and field studies showed that decomposition was more strongly influenced by local substratum characteristics (especially soil N availability) and plant functional type composition than by large-scale temperature differences. The very small responsiveness of organic matter decomposition in the field (experimental temperature increase <1 °C) compared with the laboratory (experimental increases of 4 or 8 °C) shows that substantial warming is required before significant effects can be detected.

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.

(Table 1) Climate characteristics of the four North American Tundra Experiment (ITEX) sites

Climate warming is expected to differentially affect CO2 exchange of the diverse ecosystems in the Arctic. Quantifying responses of CO2 exchange to warming in these ecosystems will require coordinated experimentation using standard temperature manipulations and measurements. Here, we used the International Tundra Experiment (ITEX) standard warming treatment to determine CO2 flux responses to growing-season warming for ecosystems spanning natural temperature and moisture ranges across the Arctic biome. We used the four North American Arctic ITEX sites (Toolik Lake, Atqasuk, and Barrow [USA] and Alexandra Fiord [Canada]) that span 10° of latitude. At each site, we investigated the CO2 responses to warming in both dry and wet or moist ecosystems. Net ecosystem CO2 exchange (NEE), ecosystem respiration (ER), and gross ecosystem photosynthesis (GEP) were assessed using chamber techniques conducted over 24-h periods sampled regularly throughout the summers of two years at all sites. At Toolik Lake, warming increased net CO2 losses in both moist and dry ecosystems. In contrast, at Atqasuk and Barrow, warming increased net CO2 uptake in wet ecosystems but increased losses from dry ecosystems. At Alexandra Fiord, warming improved net carbon uptake in the moist ecosystem in both years, but in the wet and dry ecosystems uptake increased in one year and decreased the other. Warming generally increased ER, with the largest increases in dry ecosystems. In wet ecosystems, high soil moisture limited increases in respiration relative to increases in photosynthesis. Warming generally increased GEP, with the notable exception of the Toolik Lake moist ecosystem, where warming unexpectedly decreased GEP >25%. Overall, the respiration response determined the effect of warming on ecosystem CO2 balance. Our results provide the first multiple-site comparison of arctic tundra CO2 flux responses to standard warming treatments across a large climate gradient. These results indicate that (1) dry tundra may be initially the most responsive ecosystems to climate warming by virtue of strong increases in ER, (2) moist and wet tundra responses are dampened by higher water tables and soil water contents, and (3) both GEP and ER are responsive to climate warming, but the magnitudes and directions are ecosystem-dependent.