Plant community dynamics in a calcareous grassland under climate change manipulations

This study investigates the effects of field manipulations of local climate to determine the potential impact of climate change on plant community dynamics in a calcareous grassland. The experimental site is located in a grassland at the Wytham estate, Oxfordshire, UK. The one hectare study area is within a 10 ha abandoned arable field on Jurassic corallian limestone. Two climate change scenarios were used: warmer winters with increased summer rainfall and warmer winters with summer drought. Plant cover and species richness were significantly increased in plots receiving supplemented summer rainfall, while the amount of litter was significantly reduced. Litter formation was significantly increased by winter warming and drought. The responses of the plant community to the climate manipulations were related to the life-history attributes of the dominant species. Seedling recruitment was limited by microsite availability, which also varied in the different climate manipulations. The results are discussed in terms of successional dynamics. They suggest that warmer winters may delay succession, as gap formation in the sward will provide sites for colonisation of annuals, thereby enabling their persistence in the sward. Under wetter conditions during summer, perennial grasses tend to close the sward, thereby inhibiting the establishment of later successional species.     

Mycorrhizal fungal abundance is affected by long-term climatic manipulations in the field

Climate change treatments – winter warming, summer drought and increased summer precipitation – have been imposed on an upland grassland continuously for 7 years. The vegetation was surveyed yearly. In the seventh year, soil samples were collected on four occasions through the growing season in order to assess mycorrhizal fungal abundance. Mycorrhizal fungal colonisation of roots and extraradical mycorrhizal hyphal (EMH) density in the soil were both affected by the climatic manipulations, especially by summer drought. Both winter warming and summer drought increased the proportion of root length colonised (RLC) and decreased the density of external mycorrhizal hyphal. Much of the response of mycorrhizal fungi to climate change could be attributed to climate‐induced changes in the vegetation, especially plant species relative abundance. However, it is possible that some of the mycorrhizal response to the climatic manipulations was direct – for example, the response of the EMH density to the drought treatment. Future work should address the likely change in mycorrhizal functioning under warmer and drier conditions.     

Increasing hydrologic variability threatens depleted anadromous fish populations

Predicting effects of climate change on species and ecosystems depend on understanding responses to shifts in means (such as trends in global temperatures), but also shifts in climate variability. To evaluate potential responses of anadromous fish populations to an increasingly variable environment, we performed a hierarchical analysis of 21 Chinook salmon populations from the Pacific Northwest, examining support for changes in river flows and flow variability on population growth. More than half of the rivers analyzed have already experienced significant increases in flow variability over the last 60 years, and this study shows that this increase in variability in freshwater flows has a more negative effect than any other climate signal included in our model. Climate change models predict that this region will experience warmer winters and more variable flows, which may limit the ability of these populations to recover.     

Flying in the face of climate change: a review of climate change, past, present and future

The distribution and abundance of birds is known to depend critically upon climate variability at a range of temporal and spatial scales. In this paper we review historical changes in climate in the context of what is known about climate variability over the last millennium, with particular reference to the British Isles. The climate of Britain is now warmer than it has been in at least 340 years, with the 1990s decade 0.5 °C warmer than the 1961–1990 average. In addition, the frequency of cold days (mean temperature below 0 °C), particularly during March and November, has declined and there has been a marked shift in the seasonality of precipitation, with winters becoming substantially wetter and summers becoming slightly drier. Current understanding is that the rate of future warming is likely to accelerate with more frequent and more intense summer heatwaves, milder winters, an increase in winter rainfall, an increased risk of winter river floods, and an increase in mean sea-level and associated coastal flooding. All of these aspects of climate change are likely to impact on coastal birds. A range of potential future climate scenarios for the British Isles are presented derived from recently completed global climate model experiments. For migrant bird species, changes in the British climate have also to be seen within the context of remote climate change in both the breeding and the overwintering grounds.     

Direct and indirect effects of climate change on insect herbivores: Auchenorrhyncha (Homoptera)

1. Novel manipulations of local climate were employed to investigate how warmer winters with either wetter or drier summers would affect the Auchenorrhyncha, a major component of the insect fauna of grasslands. Direct and indirect effects of climate manipulation were found.
2. Supplemented summer rainfall resulted in an increase in vegetation cover, leading to an increase in the abundance of the Auchenorrhyncha.
3. Summer drought, however, caused a decrease in vegetation cover, but this did not lead to a corresponding decrease in the abundance of the Auchenorrhyncha.
4. Egg hatch and the termination of nymphal hibernation occurred earlier in winter warmed plots; however, the rate of nymphal development was unaffected.     

Macro‐ and microclimatic interactions can drive variation in species' habitat associations

Many species are more restricted in their habitat associations at the leading edges of their range margins, but some species have broadened their habitat associations in these regions during recent climate change. We examine the effects of multiple, interacting climatic variables on spatial and temporal patterns of species' habitat associations, using the speckled wood butterfly, Pararge aegeria, in Britain, as our model taxon. Our analyses reveal that this species, traditionally regarded as a woodland‐dependent insect, is less restricted to woodland in regions with warmer winters and warmer and wetter summers. In addition, over the past 40 years of climate change, the species has become less restricted to woodland in locations where temperature and summer rainfall have increased most. We show that these patterns arise mechanistically because larval growth rates are slower in open (i.e. nonwoodland) habitats associated with colder microclimates in winter and greater host plant desiccation in summer. We conclude that macro‐ and microclimatic interactions drive variation in species' habitat associations, which for our study species resulted predominantly in a widening of habitat associations under climate change. However, species vary in their climatic and nonclimatic requirements, and so complex spatial and temporal patterns of changes in habitat associations are likely to be observed in future as the climate changes.     

Effects of changing climate on aquatic habitat and connectivity for remnant populations of a wide‐ranging frog species in an arid landscape

Amphibian species persisting in isolated streams and wetlands in desert environments can be susceptible to low connectivity, genetic isolation, and climate changes. We evaluated the past (1900–1930), recent (1981–2010), and future (2071–2100) climate suitability of the arid Great Basin (USA) for the Columbia spotted frog (Rana luteiventris) and assessed whether changes in surface water may affect connectivity for remaining populations. We developed a predictive model of current climate suitability and used it to predict the historic and future distribution of suitable climates. We then modeled changes in surface water availability at each time period. Finally, we quantified connectivity among existing populations on the basis of hydrology and correlated it with interpopulation genetic distance. We found that the area of the Great Basin with suitable climate conditions has declined by approximately 49% over the last century and will likely continue to decline under future climate scenarios. Climate conditions at currently occupied locations have been relatively stable over the last century, which may explain persistence at these sites. However, future climates at these currently occupied locations are predicted to become warmer throughout the year and drier during the frog's activity period (May – September). Fall and winter precipitation may increase, but as rain instead of snow. Earlier runoff and lower summer base flows may reduce connectivity between neighboring populations, which is already limited. Many of these changes could have negative effects on remaining populations over the next 50–80 years, but milder winters, longer growing seasons, and wetter falls might positively affect survival and dispersal. Collectively, however, seasonal shifts in temperature, precipitation, and stream flow patterns could reduce habitat suitability and connectivity for frogs and possibly other aquatic species inhabiting streams in this arid region.     

Differential sensitivity of planktonic trophic levels to extreme summer temperatures in boreal lakes

The stress–size hypothesis predicts that smaller organisms will be less sensitive to stress. Consequently, climate warming is expected to favour smaller taxa from lower trophic levels and smaller individuals within populations. To test these hypotheses, we surveyed zooplankton communities in 20 boreal lakes in Killarney Provincial Park, Canada during 2005 (an anomalously warm summer) and 2006 (a normal summer). Higher trophic levels had larger responses to warm temperatures supporting the stress–size hypothesis; however, rather than imposing negative effects, higher density and biomass were observed under warmer temperatures. As a result, larger taxa from higher trophic levels were disproportionately favoured with warming, precluding an expected shift towards smaller species. Proportionately greater increases in metabolic rates of larger organisms or altered biotic interactions (e.g. predation and competition) are possible explanations for shifts in biomass distribution. Warmer temperatures also favoured smaller individuals of the two most common species, in agreement with the stress–size hypothesis. Despite this, these populations had higher biomass in the warm summer. Therefore, reduced adult survivorship may have triggered these species to invest in reproduction over growth. Hence, warmer epilimnions, higher zooplankton biomass and smaller individuals within zooplankton populations may function as sensitive indicators of climate warming in boreal lakes.     

A Race for Space? How Sphagnum fuscum stabilizes vegetation composition during long‐term climate manipulations

Strong climate warming is predicted at higher latitudes this century, with potentially major consequences for productivity and carbon sequestration. Although northern peatlands contain one‐third of the world's soil organic carbon, little is known about the long‐term responses to experimental climate change of vascular plant communities in these Sphagnum‐dominated ecosystems. We aimed to see how long‐term experimental climate manipulations, relevant to different predicted future climate scenarios, affect total vascular plant abundance and species composition when the community is dominated by mosses. During 8 years, we investigated how the vascular plant community of a Sphagnum fuscum‐dominated subarctic peat bog responded to six experimental climate regimes, including factorial combinations of summer as well as spring warming and a thicker snow cover. Vascular plant species composition in our peat bog was more stable than is typically observed in (sub)arctic experiments: neither changes in total vascular plant abundance, nor in individual species abundances, Shannon's diversity or evenness were found in response to the climate manipulations. For three key species (Empetrum hermaphroditum, Betula nana and S. fuscum) we also measured whether the treatments had a sustained effect on plant length growth responses and how these responses interacted. Contrasting with the stability at the community level, both key shrubs and the peatmoss showed sustained positive growth responses at the plant level to the climate treatments. However, a higher percentage of moss‐encroached E. hermaphroditum shoots and a lack of change in B. nana net shrub height indicated encroachment by S. fuscum, resulting in long‐term stability of the vascular community composition: in a warmer world, vascular species of subarctic peat bogs appear to just keep pace with growing Sphagnum in their race for space. Our findings contribute to general ecological theory by demonstrating that community resistance to environmental changes does not necessarily mean inertia in vegetation response.     

Asymmetrical thermal constraints on the parapatric species boundaries of two widespread generalist butterflies

Abstract.  1. The sibling species Papilio glaucus and Papilio canadensis meet in a narrow hybrid zone believed to be maintained by temperature thresholds acting independently on both species. The present study tests if this assertion is true for the cold-adapted species, P. canadensis , which is presumed to be limited by the effect of high temperatures in late summer and/or autumn on pupal survival.
2. Three experiments were conducted examining the effects of: (i) short periods of high temperature stress in autumn, (ii) prolonged warm temperatures in autumn, and (iii) temperatures simulating warmer winters/longer springs upon the survival of P. canadensis and P. glaucus.
3. Results indicated that short periods of high temperatures did not induce the high mortality rates required to be the key factor limiting the range of P. canadensis . However, P. canadensis did exhibit a considerably lower tolerance to high temperature extremes, prolonged warm temperatures in autumn, and conditions simulating shorter/warmer winters than P. glaucus .
4. Differences in temperature tolerance throughout the pupal stage are likely to be a significant factor in maintaining the southern range limit of P. canadensis . Further warming as may occur during climate change, particularly in winter and spring, will likely affect the dynamics of southerly populations of P. canadensis.     

Climate change trade-offs in the side-blotched lizard (Uta stansburiana): effects of growing-season length and mild temperatures on winter survival

An expanding body of literature has demonstrated that global climate change continues to adversely affect many populations, species, and ecosystems. However, life-history theory also predicts possible benefits from longer growing seasons and less severe winters, particularly for ectotherms. To test the idea that climate change will have benefits as well as costs, I studied the impacts of growing-season length on growth and overwintering conditions on survival time using side-blotched lizards (Uta stansburiana). Experiments in replicate field enclosures revealed that fall growing-season length has a direct effect on overwintering body size. Laboratory experiments revealed that both size and overwintering temperature have direct effects on winter survival time. Larger lizards are more likely to survive longer regardless of winter temperature. Furthermore, animals in colder (but still mild) winter microenvironments are more likely to survive longer than those in warmer winter environments. These results indicate that warmer winters caused by global climate change have the potential to negatively affect ectotherm populations. However, longer growing seasons may offset losses by allowing additional growth and energy storage. Thus, environmental alterations associated with climate change may be simultaneously beneficial and detrimental, and the long-term persistence of certain organisms may depend on the relative strength of their effects.     

Effects of Warming, Summer Drought, and CO2 Enrichment on Aboveground Biomass Production, Flowering Phenology, and Community Structure in an Upland Grassland Ecosystem

Future climate scenarios predict simultaneous changes in environmental conditions, but the impacts of multiple climate change drivers on ecosystem structure and function remain unclear. We used a novel experimental approach to examine the responses of an upland grassland ecosystem to the 2080 climate scenario predicted for the study area (3.5°C temperature increase, 20% reduction in summer precipitation, atmospheric CO₂ levels of 600 ppm) over three growing seasons. We also assessed whether patterns of grassland response to a combination of climate change treatments could be forecast by ecosystem responses to single climate change drivers. Effects of climate change on aboveground production showed considerable seasonal and interannual variation; April biomass increased in response to both warming and the simultaneous application of warming, summer drought, and CO₂ enrichment, whereas October biomass responses were either non-significant or negative depending on the year. Negative impacts of summer drought on production were only observed in combination with a below-average rainfall regime, and showed lagged effects on spring biomass. Elevated CO₂ had no significant effect on aboveground biomass during this study. Both warming and the 2080 climate change scenario were associated with a significant advance in flowering time for the dominant grass species studied. However, flowering phenology showed no significant response to either summer drought or elevated CO₂. Species diversity and equitability showed no response to climate change treatments throughout this study. Overall, our data suggest that single-factor warming experiments may provide valuable information for projections of future ecosystem changes in cool temperate grasslands.     

Predicting potential responses to future climate in an alpine ungulate: interspecific interactions exceed climate effects

The altitudinal shifts of many montane populations are lagging behind climate change. Understanding habitual, daily behavioural rhythms, and their climatic and environmental influences, could shed light on the constraints on long‐term upslope range‐shifts. In addition, behavioural rhythms can be affected by interspecific interactions, which can ameliorate or exacerbate climate‐driven effects on ecology. Here, we investigate the relative influences of ambient temperature and an interaction with domestic sheep (Ovis aries) on the altitude use and activity budgets of a mountain ungulate, the Alpine chamois (Rupicapra rupicapra). Chamois moved upslope when it was hotter but this effect was modest compared to that of the presence of sheep, to which they reacted by moving 89–103 m upslope, into an entirely novel altitudinal range. Across the European Alps, a range‐shift of this magnitude corresponds to a 46% decrease in the availability of suitable foraging habitat. This highlights the importance of understanding how factors such as competition and disturbance shape a given species’ realised niche when predicting potential future responses to change. Furthermore, it exposes the potential for manipulations of species interactions to ameliorate the impacts of climate change, in this case by the careful management of livestock. Such manipulations could be particularly appropriate for species where competition or disturbance already strongly restricts their available niche. Our results also reveal the potential role of behavioural flexibility in responses to climate change. Chamois reduced their activity when it was warmer, which could explain their modest altitudinal migrations. Considering this behavioural flexibility, our model predicts a small 15–30 m upslope shift by 2100 in response to climate change, less than 4% of the altitudinal shift that would be predicted using a traditional species distribution model‐type approach (SDM), which assumes that species’ behaviour remains unchanged as climate changes. Behavioural modifications could strongly affect how species respond to a changing climate.     

Range position and climate sensitivity: The structure of among‐population demographic responses to climatic variation

Species’ distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species’ climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long‐term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long‐term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species’ climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species‐interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.     

Geographic variation and thermal plasticity shape salamander metabolic rates under current and future climates

Predicted changes in global temperature are expected to increase extinction risk for ectotherms, primarily through increased metabolic rates. Higher metabolic rates generate increased maintenance energy costs which are a major component of energy budgets. Organisms often employ plastic or evolutionary (e.g., local adaptation) mechanisms to optimize metabolic rate with respect to their environment. We examined relationships between temperature and standard metabolic rate across four populations of a widespread amphibian species to determine if populations vary in metabolic response and if their metabolic rates are plastic to seasonal thermal cues. Populations from warmer climates lowered metabolic rates when acclimating to summer temperatures as compared to spring temperatures. This may act as an energy saving mechanism during the warmest time of the year. No such plasticity was evident in populations from cooler climates. Both juvenile and adult salamanders exhibited metabolic plasticity. Although some populations responded to historic climate thermal cues, no populations showed plastic metabolic rate responses to future climate temperatures, indicating there are constraints on plastic responses. We postulate that impacts of warming will likely impact the energy budgets of salamanders, potentially affecting key demographic rates, such as individual growth and investment in reproduction.     

Plasticity in functional traits in the context of climate change: a case study of the subalpine forb Boechera stricta (Brassicaceae)

Environmental variation often induces shifts in functional traits, yet we know little about whether plasticity will reduce extinction risks under climate change. As climate change proceeds, phenotypic plasticity could enable species with limited dispersal capacity to persist in situ, and migrating populations of other species to establish in new sites at higher elevations or latitudes. Alternatively, climate change could induce maladaptive plasticity, reducing fitness, and potentially stalling adaptation and migration. Here, we quantified plasticity in life history, foliar morphology, and ecophysiology in Boechera stricta (Brassicaceae), a perennial forb native to the Rocky Mountains. In this region, warming winters are reducing snowpack and warming springs are advancing the timing of snow melt. We hypothesized that traits that were historically advantageous in hot and dry, low‐elevation locations will be favored at higher elevation sites due to climate change. To test this hypothesis, we quantified trait variation in natural populations across an elevational gradient. We then estimated plasticity and genetic variation in common gardens at two elevations. Finally, we tested whether climatic manipulations induce plasticity, with the prediction that plants exposed to early snow removal would resemble individuals from lower elevation populations. In natural populations, foliar morphology and ecophysiology varied with elevation in the predicted directions. In the common gardens, trait plasticity was generally concordant with phenotypic clines from the natural populations. Experimental snow removal advanced flowering phenology by 7 days, which is similar in magnitude to flowering time shifts over 2–3 decades of climate change. Therefore, snow manipulations in this system can be used to predict eco‐evolutionary responses to global change. Snow removal also altered foliar morphology, but in unexpected ways. Extensive plasticity could buffer against immediate fitness declines due to changing climates.     

Response of an arctic predator guild to collapsing lemming cycles

Alpine and arctic lemming populations appear to be highly sensitive to climate change, and when faced with warmer and shorter winters, their well-known high-amplitude population cycles may collapse. Being keystone species in tundra ecosystems, changed lemming dynamics may convey significant knock-on effects on trophically linked species. Here, we analyse long-term (1988–2010), community-wide monitoring data from two sites in high-arctic Greenland and document how a collapse in collared lemming cyclicity affects the population dynamics of the predator guild. Dramatic changes were observed in two highly specialized lemming predators: snowy owl and stoat. Following the lemming cycle collapse, snowy owl fledgling production declined by 98 per cent, and there was indication of a severe population decline of stoats at one site. The less specialized long-tailed skua and the generalist arctic fox were more loosely coupled to the lemming dynamics. Still, the lemming collapse had noticeable effects on their reproductive performance. Predator responses differed somewhat between sites in all species and could arise from site-specific differences in lemming dynamics, intra-guild interactions or subsidies from other resources. Nevertheless, population extinctions and community restructuring of this arctic endemic predator guild are likely if the lemming dynamics are maintained at the current non-cyclic, low-density state.     

Increase in body size is correlated to warmer winters in a passerine bird as inferred from time series data

Climate change is expected to affect natural populations in many ways. One way of getting an understanding of the effects of a changing climate is to analyze time series of natural populations. Therefore, we analyzed time series of 25 and 20 years, respectively, in two populations of the citril finch (Carduelis citrinella) to understand the background of a dramatic increase in wing length in this species over this period, ranging between 1.3 and 2.9 phenotypic standard deviations. We found that the increase in wing length is closely correlated to warmer winters and in one case to rain in relation to temperature in the summer. In order to understand the process of change, we implemented seven simulation models, ranging from two nonadaptive models (drift and sampling), and five adaptive models with selection and/or phenotypic plasticity involved and tested these models against the time series of males and females from the two population separately. The nonadaptive models were rejected in each case, but the results were mixed when it comes to the adaptive models. The difference in fit of the models was sometimes not significant indicating that the models were not different enough. In conclusion, the dramatic change in mean wing length can best be explained as an adaptive response to a changing climate.     

Effects of climate warming and species richness on photochemistry of grasslands

In view of the projected climatic changes and the global decrease in plant species diversity, it is critical to understand the effects of elevated air temperature (T_air) and species richness (S) on physiological processes in plant communities. Therefore, an experiment of artificially assembled grassland ecosystems, with different S (one, three or nine species), growing in sunlit climate-controlled chambers at ambient T_air and ambient T_air + 3°C was established. We investigated whether grassland species would be more affected by midday high-temperature stress during summer in a warmer climate scenario. The effect of elevated T_air was expected to differ with S. This was tested in the second and third experimental years by means of chlorophyll a fluorescence. Because acclimation to elevated T_air would affect the plant's stress response, the hypothesis of photosynthetic acclimation to elevated T_air was tested in the third year by gas exchange measurements in the monocultures. Plants in the elevated T_air chambers suffered more from midday stress on warm summer days than those in ambient chambers. In absence of severe drought, the quantum yield of PSII was not affected by elevated T_air. Our results further indicate that species had not photosynthetically acclimated to a temperature increase of 3°C after 3 years exposure to a warmer climate. Although effects of S and T_air × S interactions were mostly not significant in our study, we expect that combined effects of T_air and S would be important in conditions of severe drought events.     

Heating up relations between cold fish: competition modifies responses to climate change

Most predictions about species responses to climate change ignore species interactions. Helland and colleagues (2011) test whether this assumption is valid by evaluating whether ice cover affects competition between brown trout [Salmo trutta (L.)] and Arctic charr [Salvelinus alpines (L.)]. They show that increasing ice cover correlates with lower trout biomass when Arctic charr co-occur, but not in charr's absence. In experiments, charr grew better in the cold, dark environments that typify ice-covered lakes. Decreasing ice cover with warmer winters could mean more trout and fewer charr. More generally, their results provide an excellent example, suggesting that species interactions can strongly modify responses to climate change.