Gregory H. R. Henry

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.

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 ...

Experiment, monitoring, and gradient methods used to infer climate change effects on plant communities yield consistent patterns

Significance Methodological constraints can limit our ability to quantify potential impacts of climate warming. We assessed the consistency of three approaches in estimating warming effects on plant community composition: manipulative warming experiments, repeat sampling under ambient temperature change (monitoring), and space-for-time substitution. The three approaches showed agreement in the direction of change (an increase in the relative abundance of species with a warmer thermal niche), but differed in the magnitude of change estimated. Experimental and monitoring approaches were similar in magnitude, whereas space-for-time comparisons indicated a much stronger response. These results suggest that all three approaches are valid, but experimental warming and long-term monitoring are best suited for forecasting impacts over the coming decades. Inference about future climate change impacts typically relies on one of three approaches: manipulative experiments, historical comparisons (broadly defined to include monitoring the response to ambient climate fluctuations using repeat sampling of plots, dendroecology, and paleoecology techniques), and space-for-time substitutions derived from sampling along environmental gradients. Potential limitations of all three approaches are recognized. Here we address the congruence among these three main approaches by comparing the degree to which tundra plant community composition changes (i) in response to in situ experimental warming, (ii) with interannual variability in summer temperature within sites, and (iii) over spatial gradients in summer temperature. We analyzed changes in plant community composition from repeat sampling (85 plant communities in 28 regions) and experimental warming studies (28 experiments in 14 regions) throughout arctic and alpine North America and Europe. Increases in the relative abundance of species with a warmer thermal niche were observed in response to warmer summer temperatures using all three methods; however, effect sizes were greater over broad-scale spatial gradients relative to either temporal variability in summer temperature within a site or summer temperature increases induced by experimental warming. The effect sizes for change over time within a site and with experimental warming were nearly identical. These results support the view that inferences based on space-for-time substitution overestimate the magnitude of responses to contemporary climate warming, because spatial gradients reflect long-term processes. In contrast, in situ experimental warming and monitoring approaches yield consistent estimates of the magnitude of response of plant communities to climate warming.

Phenological response of tundra plants to background climate variation tested using the International Tundra Experiment

The rapidly warming temperatures in high-latitude and alpine regions have the potential to alter the phenology of Arctic and alpine plants, affecting processes ranging from food webs to ecosystem trace gas fluxes. The International Tundra Experiment (ITEX) was initiated in 1990 to evaluate the effects of expected rapid changes in temperature on tundra plant phenology, growth and community changes using experimental warming. Here, we used the ITEX control data to test the phenological responses to background temperature variation across sites spanning latitudinal and moisture gradients. The dataset overall did not show an advance in phenology; instead, temperature variability during the years sampled and an absence of warming at some sites resulted in mixed responses. Phenological transitions of high Arctic plants clearly occurred at lower heat sum thresholds than those of low Arctic and alpine plants. However, sensitivity to temperature change was similar among plants from the different climate zones. Plants of different communities and growth forms differed for some phenological responses. Heat sums associated with flowering and greening appear to have increased over time. These results point to a complex suite of changes in plant communities and ecosystem function in high latitudes and elevations as the climate warms.

Reconstruction of Summer Temperature for a Canadian High Arctic Site from Retrospective Analysis of the Dwarf Shrub, Cassiope tetragona

Abstract We used retrospective analysis of the widespread evergreen dwarf-shrub, Cassiope tetragona, to reconstruct average summer air temperature for Alexandra Fiord, Ellesmere Island, Canada. Retrospective analysis is a technique based on dendrochronological methods. In this study, chronologies are based on the morphological characteristics of the plant stems. Two growth and two reproduction chronologies, ranging from 80 to 118 years long, were developed from each of two populations at the High Arctic site. We used multiple regression models to develop a 100-year-long (1895–1994) reconstruction of July–September average air temperature that explained 45% of the climatic variance in the instrumental record. The reconstruction revealed an increase in summer temperature from ∼1905 to the early 1960s, a cooling trend from the mid-1960 to the 1970s, and an increase in temperature after 1980. These historical temperature patterns correspond well with those from other climate proxies from sites on Ellesmere and Devon Islands. As well, the similarity between our model and an arctic-wide proxy temperature time series suggests that the Cassiope-based reconstruction contains a large-scale temperature signal. There is great potential for the development of proxy climate data using Cassiope tetragona from sites throughout the Arctic.

Dendrochronological Potential of the Arctic Dwarf-Shrub Cassiope tetragona

In this report, we describe the use of dendrochronological techniques on the circumpolar, evergreen dwarf-shrub, Cassiope tetragona. Using techniques such as crossdating and standardization, and the software programs COFECHA and ARSTAN, we developed C. tetragona growth and reproduction chronologies for sites in the Canadian High Arctic. High-resolution chronologies may be used to reconstruct past climate and phase changes in large-scale modes of atmospheric circulation (e.g. Arctic Oscillation, North Atlantic Oscillation), to investigate the growth and reproductive responses of the plant to ambient and manipulated environmental variables, and to reconstruct the plants past ecohydrology (δ18O, δD, δ13C), gas exchange (δ13C) and mineral nutrition (δ15N). As C. tetragona is a circumpolar species, chronologies may be developed throughout the Arctic at sites where no trees exist, and thus provide new information on the past climate and environmental history of sites and regions previously unstudied.

Long-term Abundance Patterns of Barren-ground Caribou Using Trampling Scars on Roots of Picea Mariana in the Northwest Territories, Canada

ABSTRACT The aim of this study was to reconstruct population dynamics of barren-ground caribou (Rangifer tarandus groenlandicus) herds from the frequency of trampling scars on tree roots of black spruce (Picea mariana [Mill.] BSP) in the forest-tundra of central Northwest Territories, Canada. Two groups of sites were sampled that roughly corresponded with the migration routes of the Bathurst and Beverly caribou herds. The caribou migrate annually for long distances from the forest to the open tundra in late spring, and return to the forest in the autumn. The scar frequency distribution was determined by careful crossdating and the influence of root age was assessed to account for the increasing underestimation of caribou abundance with the increasing age of the roots. The scar frequency distributions (dated from A.D. 1760 to 2000) from both groups of sites showed similar abundance patterns through time. Caribou numbers were high during the mid-1940s, and 1990s, and were very low during the 1920s, 1950s–1970s, and at the turn of the 21st century. These abundance patterns determined from scar frequencies correlate strongly with data obtained from traditional knowledge of Dogrib elders in the region and animal counts based on aerial photography. The scar frequency distribution developed in this study is the longest proxy record of caribou abundance to date.

Demography of three dominant sedges under contrasting grazing regimes in the High Arctic

. Tiller demography of Carex aquatilis ssp. stans, Carex membranacea, and Eriophorum angustifolium ssp. triste was investigated in ungrazed and grazed high arctic vegetation on central Ellesmere Island, Canada. Tiller birth, growth, flowering and death were studied from excavated clonal fragments, and tiller density and biomass were studied from excavated turfs. Five life-cycle stages were determined: dormant buds, juvenile, mature, flowering and dead tillers. A stage-based transition matrix model was developed to estimate the long-term dynamics of the sedge populations and to compare life-history strategies between ungrazed and grazed populations. Short-term and retrospective models, based on the growth during the sampling year and during the lifetime of the clonal fragments, respectively, were compared to see how well the short-term model can describe demography of long-lived plants. According to the short-term model, tiller populations were decreasing (λ < 1 except for C. membranacea), whereas the retrospective model indicated that the tiller populations were increasing. Tiller population growth rates did not differ between ungrazed and grazed habitats. Nevertheless, the similar growth rates may be obtained by balanced differences in the vital rates between plants of the two habitats. The plants in the ungrazed habitat tended to remain in their current life-cycle stage, whereas plants in the grazed habitat moved quickly to the next stage and died earlier. C. aquatilis ssp. stans appears to gain a competitive advantage over the other species under intensive grazing, as indicated by the higher tiller density and greater below-ground biomass in grazed vegetation. The greater amount of below-ground biomass apparently buffers C. aquatilis ssp. stans against grazing better than the other species.

Effects of Simulated Grazing in Ungrazed Wet Sedge Tundra in the High Arctic

Abstract Wet sedge tundra communities in the High Arctic are valuable sources of forage for several resident and migratory herbivores; however, the effects of grazing on these systems have been rarely studied. We simulated grazing in two wet sedge meadows at a site on Ellesmere Island that has not been affected by grazing. Over two summers, we clipped plots at four different frequencies and removed litter to assess effects on aboveground net primary production, availability of soil nitrogen, shoot concentrations of carbon and nitrogen, and soil temperature and moisture regimes. Available soil nitrate and ammonium were highest in plots with intermediate clipping frequencies. Shoot nitrogen concentrations were also greater at intermediate clipping frequencies in two of the four species studied. Aboveground net primary production decreased after clipping, regardless of frequency. Litter removal resulted in slightly increased soil moisture, but had no effect on aboveground net primary production. Soil temperature was not affected by any of our treatments. These results suggest that nitrogen cycling is stimulated by intermediate frequencies of simulated grazing, but clipping decreased aboveground net primary production in ungrazed high arctic wet sedge tundra.

Population structure of three dominant sedges under muskox herbivory in the High Arctic.

We investigated population structure of Carex aquatilis ssp. stans, Carex membranacea, and Eriophorum angustifolium ssp. triste in ungrazed and grazed (by muskoxen) high arctic sedge meadow vegetat...