Biological responses to permafrost thaw slumping in Canadian Arctic lakes

1. Rapid environmental change occurring in high‐latitude regions has the potential to cause extensive thawing of permafrost. Retrogressive thaw slumps are a particularly spectacular form of permafrost degradation that can significantly impact lake–water chemistry; however, to date, the effects on aquatic biota have received little attention. 2. We used a diatom‐based palaeolimnological approach featuring a paired lake study design to examine the impact of thaw slumping on freshwater ecosystems in the low Arctic of western Canada. We compared biological responses in six lakes affected by permafrost degradation with six undisturbed, reference lakes. 3. Slump‐affected lakes exhibited greater biological change than the paired reference systems, although all systems have undergone ecologically significant changes over the last 200 years. Four of the six reference systems showed an increase in the relative abundance of planktonic algal taxa (diatoms and scaled chrysophytes), the earliest beginning about 1900, consistent with increased temperature trends in this region. 4. The response of sedimentary diatoms to thaw slumping was understandably variable, but primarily related to the intensity of disturbance and associated changes in aquatic habitat. Five of the slump‐affected lakes recorded increases in the abundance and diversity of periphytic diatoms at the presumed time of slump initiation, consistent with increased water clarity and subsequent development of aquatic macrophyte communities. Slump‐affected lakes generally displayed lower nutrient levels; however, in one system, thaw slumping, induced by an intense fire at the site in 1968, ostensibly led to pronounced nutrient enrichment that persists today. 5. Our results demonstrate that retrogressive thaw slumping represents an important stressor to the biological communities of lakes in the western Canadian Arctic and can result in a number of limnological changes. We also show that palaeolimnological methods are effective for inferring the timing and response of aquatic ecosystems to permafrost degradation. These findings provide the first long‐term perspective on the biological response to permafrost thaw, a stressor that will become increasingly important as northern landscapes respond to climate change.     

Identifying from recent sediment records the effects of nutrients and climate on diatom dynamics in Loch Leven

1. Changes in nutrients and climate have occurred over approximately the same timescales in many European lake catchments. Here, we attempt to interpret the sedimentary diatom record of a large shallow lake, Loch Leven, in relation to these pressures using information gained from analysis of long‐term data sets of water quality, climate and planktonic diatoms. 2. The core data indicate the enrichment of Loch Leven starting in c. 1800–1850, most likely from agricultural practices in the catchment, with a more marked phase since c. 1940–1950 caused by increased phosphorus inputs from sewage treatment works, land drainage and a woollen mill. 3. While the recent diatom plankton remains are dominated by taxa associated with nutrient‐rich conditions, an increase in Aulacoseira subarctica relative to Stephanodiscus taxa since the mid‐1980s suggests that reductions in external catchment sources of nutrients (since 1985) may have resulted in partial recovery. This observation accords well with the long‐term monitoring series of water chemistry and phytoplankton. 4. On a decadal‐centennial scale, the eutrophication signal in the sediment record outweighs any evidence of climate as a control on the diatom community. However, at an inter‐annual scale, while the diatom data exhibit high variability, there are several changes in species composition in the recent fossil record that may be attributed to climatic controls. 5. The study highlights the value of a palaeolimnological approach, particularly when coupled with long‐term data sets, for developing our understanding of environmental change at a range of temporal scales. The diatom record in the sediment can be used effectively to track recovery from eutrophication, but requires greater understanding of contemporary ecology to fully interpret climate impacts. 6. The study illustrates the complexity of ecosystem response to synchronous changes in nutrients and climate, and the difficulty of disentangling the effects of these multiple, interacting pressures.     

Variable sensitivity of plant communities in Iceland to experimental warming

Facing an increased threat of rapid climate change in cold‐climate regions, it is important to understand the sensitivity of plant communities both in terms of degree and direction of community change. We studied responses to 3–5 years of moderate experimental warming by open‐top chambers in two widespread but contrasting tundra communities in Iceland. In a species‐poor and nutrient‐deficient moss heath, dominated by Racomitrium lanuginosum, mean daily air temperatures at surface were 1–2°C higher in the warmed plots than the controls whereas soil temperatures tended to be lower in the warmed plots throughout the season. In a species‐rich dwarf shrub heath on relatively rich soils at a cooler site, dominated by Betula nana and R. lanuginosum, temperature changes were in the same direction although more moderate. In the moss heath, there were no detectable community changes while significant changes were detected in the dwarf shrub heath: the abundance of deciduous and evergreen dwarf shrubs significantly increased (>50%), bryophytes decreased (18%) and canopy height increased (100%). Contrary to some other studies of tundra communities, we detected no changes in species richness or other diversity measures in either community and the abundance of lichens did not change. It is concluded that the sensitivity of Icelandic tundra communities to climate warming varies greatly depending on initial conditions in terms of species diversity, dominant species, soil and climatic conditions as well as land‐use history.     

Responses of High Arctic wet sedge tundra to climate warming since 1980

The global climate is changing rapidly and Arctic regions are showing responses to recent warming. Responses of tundra ecosystems to climate change have been examined primarily through short‐term experimental manipulations, with few studies of long‐term ambient change. We investigated changes in above‐ and belowground biomass of wet sedge tundra to the warming climate of the Canadian High Arctic over the past 25 years. Aboveground standing crop was harvested from five sedge meadow sites and belowground biomass was sampled from one of the sites in the early 1980s and in 2005 using the same methods. Aboveground biomass was on average 158% greater in 2005 than in the early 1980s. The belowground biomass was also much greater in 2005: root biomass increased by 67% and rhizome biomass by 139% since the early 1980s. Dominant species from each functional group (graminoids, shrubs and forbs) showed significant increases in aboveground biomass. Responsive species included the dominant sedge species Carex aquatilis stans, C. membranacea, and Eriophorum angustifolium, as well as the dwarf shrub Salix arctica and the forb Polygonum viviparum. However, diversity measures were not different between the sample years. The greater biomass correlated strongly with increased annual and summer temperatures over the same time period, and was significantly greater than the annual variation in biomass measured in 1980–1983. Increased decomposition and mineralization rates, stimulated by warmer soils, were likely a major cause of the elevated productivity, as no differences in the mass of litter were found between sample periods. Our results are corroborated by published short‐term experimental studies, conducted in other wet sedge tundra communities which link warming and fertilization with elevated decomposition, mineralization and tundra productivity. We believe that this is the first study to show responses in High Arctic wet sedge tundra to recent climate change.     

Global warming will affect the genetic diversity and uniqueness of Lycaena helle populations

The climate warming of the postglacial has strongly reduced the distribution of cold‐adapted species over most of Central Europe. Such taxa have therefore become extinct over most of the lowlands and shifted to higher altitudes where they have survived to the present day. The lycaenid butterfly Lycaena helle follows this pattern of former widespread distribution and later restriction to mountain areas such as the European middle mountains. We sampled 203 individuals from 10 populations representing six mountain ranges (Pyrenees, Jura, Massif Central, Morvan, Vosges and Ardennes) over the species' western distribution. Allozyme and microsatellite polymorphisms were analysed to study the genetic status of these highly fragmented populations. Both molecular marker systems revealed a strong genetic differentiation among the analysed populations, coinciding with the orographic structure and highly restricted gene flow among them. The large‐scale genetic differentiation is more pronounced in allozymes (F_CT: 0.326) than in microsatellites (R_CT: 0.113), but microsatellites show a higher resolution on the regional scale (R_SC: 0.082) compared with allozymes (F_SC: n.s.). For both analytical tools, we found private alleles occurring exclusively in a single mountain area. The highly fragmented and isolated occurrence of populations is supported by the distribution pattern of potentially suitable climate suggested by species distribution models. Model projections under two climate warming scenarios predict a decline of climatically suitable areas, which will result in the extinction of most of the populations showing unique genetic characteristics.     

Global warming is changing the dynamics of Arctic host-parasite systems

Global climate change is altering the ecology of infectious agents and driving the emergence of disease in people, domestic animals, and wildlife. We present a novel, empirically based, predictive model for the impact of climate warming on development rates and availability of an important parasitic nematode of muskoxen in the Canadian Arctic, a region that is particularly vulnerable to climate change. Using this model, we show that warming in the Arctic may have already radically altered the transmission dynamics of this parasite, escalating infection pressure for muskoxen, and that this trend is expected to continue. This work establishes a foundation for understanding responses to climate change of other host-parasite systems, in the Arctic and globally.     

Early pest development and loss of biological control are associated with urban warming

Climate warming is predicted to cause many changes in ectotherm communities, one of which is phenological mismatch, wherein one species'' development advances relative to an associated species or community. Phenological mismatches already lead to loss of pollination services, and we predict that they also cause loss of biological control. Here, we provide evidence that a pest develops earlier due to urban warming but that phenology of its parasitoid community does not similarly advance. This mismatch is associated with greater egg production that likely leads to more pests on trees.     

Ecosystem change and stability over multiple decades in the Swedish subarctic: complex processes and multiple drivers

The subarctic environment of northernmost Sweden has changed over the past century, particularly elements of climate and cryosphere. This paper presents a unique geo-referenced record of environmental and ecosystem observations from the area since 1913. Abiotic changes have been substantial. Vegetation changes include not only increases in growth and range extension but also counterintuitive decreases, and stability: all three possible responses. Changes in species composition within the major plant communities have ranged between almost no changes to almost a 50 per cent increase in the number of species. Changes in plant species abundance also vary with particularly large increases in trees and shrubs (up to 600%). There has been an increase in abundance of aspen and large changes in other plant communities responding to wetland area increases resulting from permafrost thaw. Populations of herbivores have responded to varying management practices and climate regimes, particularly changing snow conditions. While it is difficult to generalize and scale-up the site-specific changes in ecosystems, this very site-specificity, combined with projections of change, is of immediate relevance to local stakeholders who need to adapt to new opportunities and to respond to challenges. Furthermore, the relatively small area and its unique datasets are a microcosm of the complexity of Arctic landscapes in transition that remains to be documented.     

Global warming and excess nitrogen may induce butterfly decline by microclimatic cooling

Global warming may explain the current poleward shift of species distributions. However, paradoxically, climatic warming can lead to microclimatic cooling in spring by advancing plant growth, an effect worsened by excess nitrogen. We suggest that spring-developing but thermophilous organisms, such as butterflies hibernating as egg or larva, are particularly sensitive to the cooling of microclimates. Using published data on butterfly trends in distribution, we report a comparatively greater decline in egg–larva hibernators in European countries with oceanic climates and high nitrogen deposition, which supports this explanation. Furthermore, trends in abundance from a nationwide butterfly monitoring scheme reveal a 63% decrease over 13 years (1992–2004) for egg–larva hibernators in the Netherlands, contrasting with a nonsignificant trend in adult–pupa hibernators. This evidence supports the hypothesis that these environmental changes pose new threats to spring-developing, thermophilous species. We underline the threat of climate change to biodiversity, as previously suggested on the basis of mobility, habitat fragmentation and evolutionary adaptation, but we here emphasize a different ecological axis of change in habitat quality.     

Persistent effects of a discrete warming event on a polar desert ecosystem

A discrete warming event (December 21, 2001–January 12, 2002) in the McMurdo Dry Valleys, Antarctica, enhanced glacier melt, stream flow, and melting of permafrost. Effects of this warming included a rapid rise in lake levels and widespread increases in soil water availability resulting from melting of subsurface ice. These increases in liquid water offset hydrologic responses to a cooling trend experienced over the previous decade and altered ecosystem properties in both aquatic and terrestrial ecosystems. Here, we present hydrological and meteorological data from the McMurdo Dry Valleys Long Term Ecological Research project to examine the influence of a discrete climate event (warming of >2 °C) on terrestrial environments and soil biotic communities. Increases in soil moisture following this event stimulated populations of a subordinate soil invertebrate species (Eudorylaimus antarcticus, Nematoda). The pulse of melt-water had significant influences on Taylor Valley ecosystems that persisted for several years, and illustrates that the importance of discrete climate events, long recognized in hot deserts, are also significant drivers of soil and aquatic ecosystems in polar deserts. Thus, predictions of Antarctic ecosystem responses to climate change which focus on linear temperature trends may miss the potentially significant influence of infrequent climate events on hydrology and linked ecological processes.     

Rapid micro-evolution and loss of chromosomal diversity in Drosophila in response to climate warming

Concern regarding the ecological impact of rapid global warming has encouraged research on climate-induced changes in biological systems. Critical problems, still poorly understood, are the potential for rapid adaptive responses and their genetic costs to populations. The O chromosomal polymorphisms of Drosophila subobscura have been monitored at a southern Palearctic locality experiencing sustained climate warming since the mid-1970s. Observations suggest that the population is rapidly evolving in response to the new environmental conditions, and has lost a significant amount of chromosomal diversity (18.3% in 16 years). These findings are consistent with results from another population of D. subobscura, which is also undergoing climate warming, and are in accord with what would be expected from latitudinal and seasonal patterns of the various inversions. In addition, data on the O chromosomal polymorphisms from other localities throughout t he range of this species suggest that other populations vary similarly.     

Keeping up with a warming world; assessing the rate of adaptation to climate change

The pivotal question in the debate on the ecological effects of climate change is whether species will be able to adapt fast enough to keep up with their changing environment. If we establish the maximal rate of adaptation, this will set an upper limit to the rate at which temperatures can increase without loss of biodiversity.The rate of adaptation will primarily be set by the rate of microevolution since (i) phenotypic plasticity alone is not sufficient as reaction norms will no longer be adaptive and hence microevolution on the reaction norm is needed, (ii) learning will be favourable to the individual but cannot be passed on to the next generations, (iii) maternal effects may play a role but, as with other forms of phenotypic plasticity, the response of offspring to the maternal cues will no longer be adaptive in a changing environment, and (iv) adaptation via immigration of individuals with genotypes adapted to warmer environments also involves microevolution as these genotypes are better adapted in terms of temperature, but not in terms of, for instance, photoperiod.Long-term studies on wild populations with individually known animals play an essential role in detecting and understanding the temporal trends in life-history traits, and to estimate the heritability of, and selection pressures on, life-history traits. However, additional measurements on other trophic levels and on the mechanisms underlying phenotypic plasticity are needed to predict the rate of microevolution, especially under changing conditions.Using this knowledge on heritability of, and selection on, life-history traits, in combination with climate scenarios, we will be able to predict the rate of adaptation for different climate scenarios. The final step is to use ecoevolutionary dynamical models to make the link to population viability and from there to biodiversity loss for those scenarios where the rate of adaptation is insufficient.     

Effect of warming and grazing on litter mass loss and temperature sensitivity of litter and dung mass loss on the Tibetan plateau

Knowledge about the role of litter and dung decomposition in nutrient cycling and response to climate change and grazing in alpine ecosystems is still rudimentary. We conducted two separate studies to assess the relative role of warming and grazing on litter mass loss and on the temperature sensitivity of litter and dung mass loss. Experiments were conducted for 1–2 years under a controlled warming–grazing system and along an elevation gradient from 3200 to 3800 m. A free‐air temperature enhancement system (FATE) using infrared heaters and grazing significantly increased soil temperatures (average 0.5–1.6 °C) from 0 to 40 cm depth, but neither warming nor grazing affected soil moisture except early in the growing seasons at 30 cm soil depth. Heaters caused greater soil warming at night‐time compared with daytime, but grazing resulted in greater soil warming during daytime compared with night‐time. Annual average values of the soil temperature at 5 cm were 3.2, 2.4 and 0.3 °C at 3200, 3600 and 3800 m, respectively. Neither warming nor grazing caused changes of litter quality for the first year of the controlled warming–grazing experiment. The effects of warming and grazing on litter mass losses were additive, increasing litter mass losses by about 19.3% and 8.3%, respectively, for the 2‐year decomposition periods. The temperature sensitivity of litter mass losses was approximately 11% °C^?1 based on the controlled warming–grazing experiment. The annual cumulative litter mass loss was approximately 2.5 times that of dung along the elevation gradient. However, the temperature sensitivity (about 18% °C^?1) of the dung mass loss was about three times that of the litter mass loss. These results suggest greater warming at night‐time compared with daytime may accelerate litter mass loss, and grazing will enhance carbon loss to atmosphere in the region through a decrease of litter biomass and an increase of dung production with an increase of stocking rate in future warmer conditions.     

Russian Arctic warming and ‘greening’ are closely tracked by tundra shrub willows

Growth in arctic vegetation is generally expected to increase under a warming climate, particularly among deciduous shrubs. We analyzed annual ring growth for an abundant and nearly circumpolar erect willow (Salix lanata L.) from the coastal zone of the northwest Russian Arctic (Nenets Autonomous Okrug). The resulting chronology is strongly related to summer temperature for the period 1942–2005. Remarkably high correlations occur at long distances (>1600 km) across the tundra and taiga zones of West Siberia and Eastern Europe. We also found a clear relationship with photosynthetic activity for upland vegetation at a regional scale for the period 1981–2005, confirming a parallel ‘greening’ trend reported for similarly warming North American portions of the tundra biome. The standardized growth curve suggests a significant increase in shrub willow growth over the last six decades. These findings are in line with field and remote sensing studies that have assigned a strong shrub component to the reported greening signal since the early 1980s. Furthermore, the growth trend agrees with qualitative observations by nomadic Nenets reindeer herders of recent increases in willow size in the region. The quality of the chronology as a climate proxy is exceptional. Given its wide geographic distribution and the ready preservation of wood in permafrost, S. lanata L. has great potential for extended temperature reconstructions in remote areas across the Arctic.     

Validation of a diatom–phosphorus calibration set for Sweden

1. A weighted averaging (WA) regression and calibration model for diatoms and total phosphorus (TP) was developed from a dataset of 45 surface‐sediment samples from Swedish lakes. Jack‐knifed error statistics were comparable with those for similar diatom–TP datasets: r²_jack=0.47, root mean squared error of prediction (RMSEP)=0.24 log₁₀μg TP L^–1 and mean bias=–0.002 log₁₀ μg TP L^–1 for the simple WA model; r²_jack=0.36, RMSEP=0.27 log₁₀ μg TP L^–1 and mean bias=0.017 log₁₀ μg TP L^–1 for WA with tolerance downweighting. 2. The model was used to estimate TP concentrations for the Ekoln basin of Lake Mälaren using a ²¹⁰Pb‐dated sediment core record. Highly eutrophic conditions developed in the basin in the 1960s as a result of nutrient inputs from cultivated land and the city of Uppsala. A reduction in the supply of phosphorus from sewage outlets in the late 1960s resulted in less eutrophic conditions. 3. The model results indicated levels of 50–60 μg TP L^–1 prior to 1900. The rapid eutrophication of the lake basin after the 1950s and the subsequent recovery were evident from the diatom data. 4. Diatom‐inferred TP (DI–TP) values were validated by comparison with monitored data for the period 1966–95. The diatom model tended to underestimate TP at high levels (> 80 μg L^–1) but overestimate at lower concentrations. 5. A good agreement was observed between the trends in TP concentration and the DI–TP concentration and the timing of the maximum was well reflected by the diatom‐based reconstruction. A significant correlation (r²=0.69, P < 0.01) was found between DI–TP and measured TP at this site.     

The response of Alaskan arctic tundra to experimental warming: differences between short- and long-term responses

Global climate models predict continued rapid warming for most of the Arctic throughout the next century. To further understand the response of arctic tundra to climate warming, four sites in northern Alaska were warmed for five to seven consecutive growing seasons using open‐top chambers. Sites were located in dry heath and wet meadow communities near Barrow (71°18′N, 156°40′W) and Atqasuk (70°29′N, 157°25′W). Change in plant community composition was measured using a point frame method. During the period of observation, species richness declined in control plots by up to 2.7 species plot^?1. Responses to warming varied by site but similar trends included increased canopy height (?0.1 to 2.3 cm) and relative cover of standing dead plant matter (1.5–6.0%) and graminoids (1.8–5.8%) and decreased species diversity (0.1–1.7 species plot^?1) and relative cover of lichens (0.2–9.1%) and bryophytes (1.4–4.6%) (parentheses enclose the range of average values for the sites). The response to warming was separated into an initial short‐term response assessed after two growing seasons of warming and a secondary longer‐term response assessed after an additional three to five growing seasons of warming. The initial responses to warming were similar in the four sites, while the secondary responses varied by site. The response to warming was greater at Barrow than Atqasuk because of a greater initial response at Barrow. However, the long‐term response to warming was projected to be greater at Atqasuk because of a greater secondary response at Atqasuk. These findings show that predictions of vegetation change due to climate warming based on manipulative experiments will differ depending on both the duration and plant community on which the study focuses.     

The recent palaeolimnology of a remote Scottish loch with special reference to the relative impacts of regional warming and atmospheric contamination

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