Jeffrey M. Welker

Ecological Dynamics Across the Arctic Associated with Recent Climate Change

Assessing the Arctic The Arctic is experiencing some of the most rapid climate change currently under way across the globe, but consequent ecological responses have not been widely reported. At the close of the Fourth International Polar Year, Post et al. (p. 1355) review observations on ecological impacts in this sensitive region. The widespread changes occurring in terrestrial, freshwater, and marine systems, presage changes at lower latitudes that will affect natural resources, food production, and future climate buffering. At the close of the Fourth International Polar Year, we take stock of the ecological consequences of recent climate change in the Arctic, focusing on effects at population, community, and ecosystem scales. Despite the buffering effect of landscape heterogeneity, Arctic ecosystems and the trophic relationships that structure them have been severely perturbed. These rapid changes may be a bellwether of changes to come at lower latitudes and have the potential to affect ecosystem services related to natural resources, food production, climate regulation, and cultural integrity. We highlight areas of ecological research that deserve priority as the Arctic continues to warm.

Responses of tundra plants to experimental warming : Meta-analysis of the international tundra experiment

The International Tundra Experiment (ITEX) is a collaborative, multisite experiment using a common temperature manipulation to examine variability in species response across climatic and geographic gradients of tundra ecosystems. ITEX was designed specifically to examine variability in arctic and alpine species response to increased temperature. We compiled from one to four years of experimental data from 13 different ITEX sites and used meta-analysis to analyze responses of plant phenology, growth, and reproduction to experimental warming. Results indicate that key phenological events such as leaf bud burst and flowering occurred earlier in warmed plots throughout the study period; however, there was little impact on growth cessation at the end of the season. Quantitative measures of vegetative growth were greatest in warmed plots in the early years of the experiment, whereas reproductive effort and success increased in later years. A shift away from vegetative growth and toward reproductive effort and success in the fourth treatment year suggests a shift from the initial response to a secondary response. The change in vegetative response may be due to depletion of stored plant reserves, whereas the lag in reproductive response may be due to the formation of flower buds one to several seasons prior to flowering. Both vegetative and reproductive responses varied among life-forms; herbaceous forms had stronger and more consistent vegetative growth responses than did woody forms. The greater responsiveness of the herbaceous forms may be attributed to their more flexible morphology and to their relatively greater proportion of stored plant reserves. Finally, warmer, low arctic sites produced the strongest growth responses, but colder sites produced a greater reproductive response. Greater resource investment in vegetative growth may be a conservative strategy in the Low Arctic, where there is more competition for light, nutrients, or water, and there may be little opportunity for successful germination or seedling development. In contrast, in the High Arctic, heavy investment in producing seed under a higher temperature scenario may provide an opportunity for species to colonize patches of unvegetated ground. The observed differential response to warming suggests that the primary forces driving the response vary across climatic zones, functional groups, and through time.

Shrub expansion in tundra ecosystems: dynamics, impacts and research priorities

Recent research using repeat photography, long-term ecological monitoring and dendrochronology has documented shrub expansion in arctic, high-latitude and alpine tundra

The role of topography on catchment-scale water residence time

62.4 km 2 ) that represent diverse geologic and geomorphic conditions in the western Cascade Mountains of Oregon. Our primary objective was to determine the dominant physical controls on catchment-scale water residence time and specifically test the hypothesis that residence time is related to the size of the basin. Residence times were estimated by simple convolution models that described the transfer of precipitation isotopic composition to the stream network. We found that base flow mean residence times for exponential distributions ranged from 0.8 to 3.3 years. Mean residence time showed no correlation to basin area (r 2 < 0.01) but instead was correlated (r 2 = 0.91) to catchment terrain indices representing the flow path distance and flow path gradient to the stream network. These results illustrate that landscape organization (i.e., topography) rather than basin area controls catchment-scale transport. Results from this study may provide a framework for describing scale-invariant transport across climatic and geologic conditions, whereby the internal form and structure of the basin defines the first-order control on base flow residence time.

Winter Biological Processes Could Help Convert Arctic Tundra to Shrubland

Abstract In arctic Alaska, air temperatures have warmed 0.5 degrees Celsius (°C) per decade for the past 30 years, with most of the warming coming in winter. Over the same period, shrub abundance has increased, perhaps a harbinger of a conversion of tundra to shrubland. Evidence suggests that winter biological processes are contributing to this conversion through a positive feedback that involves the snow-holding capacity of shrubs, the insulating properties of snow, a soil layer that has a high water content because it overlies nearly impermeable permafrost, and hardy microbes that can maintain metabolic activity at temperatures of −6°C or lower. Increasing shrub abundance leads to deeper snow, which promotes higher winter soil temperatures, greater microbial activity, and more plant-available nitrogen. High levels of soil nitrogen favor shrub growth the following summer. With climate models predicting continued warming, large areas of tundra could become converted to shrubland, with winter processes like those described here possibly playing a critical role.

Global patterns of foliar nitrogen isotopes and their relationships with climate, mycorrhizal fungi, foliar nutrient concentrations, and nitrogen availability.

Ratios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (delta(15)N), foliar N concentrations, mycorrhizal type and climate for over 11,000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar delta(15)N by 2 per thousand, 3.2 per thousand, 5.9 per thousand, respectively, relative to nonmycorrhizal plants. Foliar delta(15)N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT >or= -0.5 degrees C, but was invariant with MAT across sites with MAT < -0.5 degrees C. In independent landscape-level to regional-level studies, foliar delta(15)N increased with increasing N availability; at the global scale, foliar delta(15)N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar delta(15)N and ultimately global patterns in N cycling.

Plot-scale evidence of tundra vegetation change and links to recent summer warming

Temperature is increasing at unprecedented rates across most of the tundra biome(1). Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity ov ...

Long-term experimental manipulation of winter snow regime and summer temperature in arctic and alpine tundra

Three 60 m long, 2·8 m high snowfences have been erected to study long-term effects of changing winter snow conditions on arctic and alpine tundra. This paper describes the experimental design and short-term effects. Open-top fiberglass warming chambers are placed along the experimental snow gradients and in controls areas outside the fences; each warming plot is paired with an unwarmed plot. The purpose of the experiment is to examine short- and long-term changes to the integrated physical-biological systems under simultaneous changes of winter snow regime and summer temperature, as part of the Long-Term Ecological Research network and the International Tundra Experiment. The sites were at Niwot Ridge, Colorado, a temperate high altitude site in the Colorado Rockies, and Toolik Lake, Alaska, a high-latitude site. Initial results indicate that although experimental designs are essentially identical at the arctic and alpine sites, experimental effects are different. The drift at Niwot Ridge lasts much longer than do the Toolik Lake drifts, so that the Niwot Ridge fence affects both summer and winter conditions, whereas the Toolik Lake fence affects primarily winter conditions. The temperature experiment also differs in effect between the sites. Although the average temperature increase at the two sites is similar (daily increase 1·5 °C at Toolik and 1·9 °C at Niwot Ridge), at Toolik Lake there is only minor diurnal variation, whereas at Niwot Ridge the daytime increases are extreme on sunny days (as much as 7–10 °C), and minimum nighttime temperatures in the chambers are often slightly cooler than ambient (by about 1 °C). The experimental drifts resulted in wintertime increases in temperature and CO2 flux. Temperatures under the deep drifts were much more consistent and warmer than in control areas, and at Niwot Ridge remained very close to 0 °C all winter. These increased temperatures were likely responsible for observed increases in system carbon loss. Initial changes to the aboveground biotic system included an increase in growth in response to both snow and warming, despite a reduced growing season. This is expected to be a transient response that will eventually be replaced by reduced growth. At least one species, Kobresia myosuroides, had almost completely died at Niwot Ridge three years after fence construction, whereas other species were increasing. We expect in both the short- and long-term to see the strongest effects of snow at the Niwot Ridge site, and stronger effects of temperature at Toolik Lake. Copyright

Annual CO2 flux in dry and moist arctic tundra: field responses to increases in summer temperatures and winter snow depth.

We examined the annual exchange of CO2 between the atmosphere and moist tussock and dry heath tundra ecosystems (which together account for over one-third of the low arctic land area) under ambient field conditions and under increased winter snow deposition, increased summer temperatures, or both. Our results indicate that these two arctic tundra ecosystems were net annual sources of CO2 to the atmosphere from September 1994 to September 1996 under ambient weather conditions and under our three climate change scenarios. Carbon was lost from these ecosystems in both winter and summer, although the majority of CO2 evolution took place during the short summer. Our results indicate that (1) warmer summer temperatures will increase annual CO2 efflux from both moist and dry tundra ecosystems by 45–55% compared to current ambient temperatures; (2) deeper winter snow cover will increase winter CO2 efflux in both moist and dry tundra ecosystems, but will decrease net summer CO2 efflux; and (3) deeper winter snow cover coupled with warmer summer temperatures will nearly double the annual amount of CO2 emitted from moist tundra and will result in a 24% increase in the annual CO2 efflux of dry tundra. If, as predicted, climate change alters both winter snow deposition and summer temperatures, then shifts in CO2 exchange between the biosphere and atmosphere will likely not be uniform across the Arctic tundra landscape. Increased snow deposition in dry tundra is likely to have a larger effect on annual CO2 flux than warmer summer temperatures alone or warmer temperatures coupled with increased winter snow depth. The combined effects of increased summer temperatures and winter snow deposition on annual CO2 flux in moist tundra will be much larger than the effects of either climate change scenario alone.

Wintertime CO2 efflux from Arctic soils: Implications for annual carbon budgets

Estimates of annual carbon loss from arctic tundra ecosystems are based nearly entirely on measurements taken during the growing season in part because of methodological limitations but also reflecting the assumption that respiration during winter is near zero. Measurements of CO2 flux during winter, however, indicate significant amounts of carbon loss from tundra ecosystems throughout the 240-day nongrowing season. In our study during the 1996 and 1997 nongrowing seasons, winter carbon losses ranged from 2.0 g CO2 m−2 season−1 in moist dwarf shrub communities to 97 g CO2 m−2 season−1 in natural snowdrift communities, with an average wintertime CO2 efflux of 45 g CO2 m−2 for all Low Arctic tundra communities (0.14 Pg CO2 yr−1 worldwide). These measurements indicate that current estimates of annual carbon loss from tundra ecosystems are low. Inclusion of wintertime losses of CO2 into annual carbon budgets increases the annual carbon efflux of arctic tundra ecosystems by 17% and changes some ecosystems from net annual sinks to net sources of CO2 to the atmosphere.