Annika Hofgaard

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 ...

Structure and regeneration patterns in a virgin Picea abies forest in northern Sweden

Abstract. A high-altitude boreal Picea abies forest, with tree ages of up to 410 yr, was studied with respect to age structure, spatial regeneration patterns, and substrate. The results suggest that recruitment is primarily dependent on germination substrate but also negatively correlated with the density of the tree layer. 60 % of all spruces < 1.3 m high grew on substrates connected with tree-fall; ca. 40 % were found on decomposing logs and stumps, covering only ca. 6 % of the forest floor. Individual logs remain important as a regeneration substrate for ca. 150 yr. Continuous presence of decomposing coarse wood is a condition for the maintenance of the population structure under the prevailing climatic conditions. Peaks in the age distribution (the 1870s and the 1940s - 1950s) are probably climatically induced. The results challenge the previous assumptions that this kind of forest undergoes cyclic development. Long-term structural stability with climatically induced minor variations may be an alternative model.

Differential Regional Treeline Dynamics in the Scandes Mountains

Abstract Age structure, tree characteristics, and environmental data were used to analyze the status of the birch treeline in three regions along the Scandes Mountains from 62°10′N to 69°50′N. Aspect and estimated relative radiation explained most of the treeline altitude across studied regions, but not all variation. Main tree establishment occurred during the 1940s in the southern and northernmost regions, and during the 1960s in the middle region. Age distribution patterns at 2 m (tree size), however, showed stable or possibly progressive treelines in the southern and middle regions but recent recession in the north. Growth rates varied through time and between regions, with an apparent decrease in the north since the 1940s. Weak negative correlations between altitude and age in the south indicate recent changes favoring tree growth or increased turnover at higher, more exposed altitudes. Although Scandinavian treelines are expected to advance in response to climate warming, this was not evident as a general pattern for all regions. Seasonally different climate patterns, browsing, and abrasion are mechanisms involved in this. These regionally different patterns have to be taken into account in predictions of future responses to avoid overestimation of, e.g., ecosystem change, carbon uptake capacity, and feedbacks to climate systems.

50 years of change in a Swedish boreal old‐growth Picea abies forest

. Changes in the tree layer (> 1.3m) and sapling layer (< 1.3m, including seedlings) of a Swedish boreal old-growth Picea abies (Norway spruce) forest from the 1930s to the 1980s were studied in permanent plots. The plots were established in 1938–1939 and re-analysed in 1983–1988. Regeneration, mortality, turnover rate in the tree layer and amount of decomposing logs as well as the time required for complete decomposition of logs were investigated using the detailed data from the 1930s. Ca. 25 % of the trees present during the first analysis were no longer alive. This mortality was balanced by recruitment from the sapling layer. The rate of mortality suggests a turnover time for the tree layer of ca. 200 yr. The number of spruces in the sapling layer has increased by ca. 85 %, hypothetically in response to an increase in amount of decomposing wood that can serve as nurse logs and stumps. The mean time for total decomposition was calculated as ca. 200 yr. Spruce regeneration on logs does not occur until the log is at least ca. 50 yr old. The survival pattern in the sapling layer suggests a high mortality rate at the seedling stage (≤ 1 yr) and a low mortality rate at the sapling stage. In conclusion, it is suggested that the amount of coarse woody debris available for regeneration, the occurrence of seedlings, and seedling mortality constitute concurrent factors through which climatic fluctuations, in a long-term perspective, direct stand recruitment and density. As a consequence, these boreal forests will be kept in a dynamical equilibrium.

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.

Contrasting climate- and land-use-driven tree encroachment patterns of subarctic tundra in northern Norway and the Kola PeninsulaThis article is a contribution to the series Tree recruitment, growth, and distribution at the circumpolar forest–tundra transition.

High-latitude regions are experiencing substantial climate change, and the forest–tundra transition is assumed to sensitively track these changes through advancing treeline and increased tundra encroachment. However, herbivores may influence these responses. The present study addresses, through analyses of age structures, growth characteristics, and climate correspondence, how mountain birch (Betula pubescens Ehrh. ssp. czerepanovii (Orlova) Hamet-Ahti) treelines and sapling cohorts beyond the treeline have responded to the last decade’s warming in six North European subarctic areas with different climate and grazing characters. The results show different response patterns among areas representing advancing, stationary, and possibly retreating treelines. Recruitment was abundant over the last decades in all areas except one, with predominantly arctic conditions, where both tree and sapling cohorts were old. Areas with high annual precipitation show advancing birch populations characterized by young indivi...

Shifts in radial growth responses of coastal Picea abies induced by climatic change during the 20th century, central Norway

Abstract Climate-growth relationships since the end of the 19th century were analysed by using regional climate data and North Atlantic Oscillation index data, together with spruce (Picea abies (L) Karst.) tree-ring data from an altitudinal gradient close to the coast in central Norway (64°N). Correlation and response function analyses were used to decipher both spatial and temporal diameter growth responses. A positive response for May and June temperatures dominated along the entire gradient, but the importance of individual months shifted through time. In periods when May and June temperatures were above their long-term means, the importance of other climate factors increased. Winter climate was significantly related to radial tree growth mainly for periods up to the 1940s, but not during the second half of the 20th century. This pattern was principally caused by changes in the response to winter precipitation. The summer climate acquired increased importance in periods with warm and moist winters. The climate-growth relationship in this study changed continuously and dynamically during the studied period; this might be a general phenomenon that should be carefully considered in dendroclimatological studies. In scenarios for forest development, it is of vital importance to build on detailed knowledge of growth responses to multiple climate variables for all seasons, particularly where the oceanic influence is predicted to be spatially extended.

Comparing warming and grazing effects on birch growth in an alpine environment – a 10-year experiment

Background : Tree encroachment of arctic tundra and alpine vegetation is a generally predicted response to climate warming. However, herbivory plays an important role in structuring these ecosystems and their responses to warming. Aims : To experimentally test how grazing and increased growing season temperature influence growth, physiognomy and stature of birch in the alpine zone. Methods: Trait responses of naturally regenerated birch saplings to warming (open-top chambers), and changed grazing regime (exclosures) were compared with those growing in unmanipulated conditions over a 10-year period (1999–2008). The effect of treatment over time and differences between treatments were analysed with repeated measures GLM (Generalised Linear Model) and simple contrasts in GLM. Results: Warming alone had no major effect on trait responses, however, significantly smaller leaves and an increased number of short shoots indicated warming-related growth constraints. Grazing showed a strong controlling effect on most traits, conserving low stature sapling stage characterised by fewer shoots and larger leaves, compared with non-grazed treatments. Conclusions: Although derived from one experimental site, the results point to a grazing-controlled response to environmental change, with climate (warming) as a secondary driver. This herbivore-driven masking of expected climate-driven tree expansion emphasises the necessity to consider changes in grazing regimes along with climate change, in order to avoid misleading interpretations regarding climate-driven tundra encroachment.

Successional trends 219 years after fire in an old Pinus sylvestris stand in northern Sweden

. We present results from repeated analyses (1962, 1993) of a permanent plot established in 1947, combined with retrospective stand age structure data, in an old Pinus sylvestris stand in Muddus National Park, northern Sweden. The study points towards a successional pathway governed by concurrent disturbance effects of climate variability, reindeer grazing and fire. This is intermediate to the two often advocated ideas on dynamics in boreal forests, that is, one of disturbance-related tree regeneration/mortality and one of continuous regeneration. When the plot was established in 1947 the tree layer (> 1.3 m) consisted of 300 individuals/ha of P. sylvestris and 62/ha of Betula pubescens. Subsequently the stand has become more dense and the species dominance has shifted. In 1993, 362 P. sylvestris and 62 Picea abies individuals were present per ha, while no Betula individuals were found. The number of dead trees increased from zero in 1947 to 200/ha (Pinus) in 1993. Pinus was also the most common species in the sapling layer (< 1.3 m) throughout the study period, though the number dropped from 8912/ha in 1947 to 51% in 1993. Dead saplings decreased from 2650/ha in 1947 to ca. 50% in 1962, and only 9% in 1993. Temporal variations in mortality and sapling mean height coincided with variations in snow depth, indicating a critical period in sapling development when saplings are exposed at the snow/atmosphere interface. The number of living Picea saplings increased slowly until 1993; no dead saplings were found. Most Pinus recruited shortly after the 1774-fire, and during the second half of the 1900s. The major part of the spruce regeneration took place during the later half of the 1900s. No successful Betula recruitment has occurred after the 1930s, and no live Betula were present in 1993, which might be explained as an effect of increased reindeer browsing – the reindeer stock has grown by 50% since 1961. Although subjected to high mortality, Pinus regenerated and maintained a seedling/sapling bank. In this way Pinus remained dominant in the tree layer after more than 200 post-fire years. The importance of the shade-tolerant Picea has slowly increased, while Betula has died off. Thus, even after 219 yr since fire there is an early successional trend in the stand. This suggests that an increased chronic disturbance (grazing/browsing by reindeer) has partly succeeded earlier discrete fire-disturbance events, and maintained a continuous seedbed favouring the shade-intolerant pine recruitment.

Effects of warming on shrub abundance and chemistry drive ecosystem-level changes in a forest-tundra ecotone

Tundra vegetation is responding rapidly to on-going climate warming. The changes in plant abundance and chemistry might have cascading effects on tundra food webs, but an integrated understanding of how the responses vary between habitats and across environmental gradients is lacking. We assessed responses in plant abundance and plant chemistry to warmer climate, both at species and community levels, in two different habitats. We used a long-term and multisite warming (OTC) experiment in the Scandinavian forest–tundra ecotone to investigate (i) changes in plant community composition and (ii) responses in foliar nitrogen, phosphorus, and carbon-based secondary compound concentrations in two dominant evergreen dwarf-shrubs (Empetrum hermaphroditum and Vaccinium vitis-idaea) and two deciduous shrubs (Vaccinium myrtillus and Betula nana). We found that initial plant community composition, and the functional traits of these plants, will determine the responsiveness of the community composition, and thus community traits, to experimental warming. Although changes in plant chemistry within species were minor, alterations in plant community composition drive changes in community-level nutrient concentrations. In view of projected climate change, our results suggest that plant abundance will increase in the future, but nutrient concentrations in the tundra field layer vegetation will decrease. These effects are large enough to have knock-on consequences for major ecosystem processes like herbivory and nutrient cycling. The reduced food quality could lead to weaker trophic cascades and weaker top down control of plant community biomass and composition in the future. However, the opposite effects in forest indicate that these changes might be obscured by advancing treeline forests.