Alpine flora dynamics – a critical review of responses to climate change in the Swedish Scandes since the early 1950s

Reports about changes of alpine plant species richness over the past 60 years in the Swedish Scandes are reviewed, synthesized and updated with data from recent reinventories. Methodologically, this endeavour is based on resurveys of the floristic composition on the uppermost 20 m of four high‐mountain summits. The key finding is that the species pool has increased by 60–170% since the 1950s and later. Some of the invading species are new to the alpine tundra, with more silvine and thermophilic properties than the extant alpine flora. Not a single species of the original flora has disappeared from any of the summits. This circumstance is discussed in perspective of widespread expectations of pending temperature‐driven extinction of alpine species in an alleged future warmer climate. These progressive changes coincided with distinct warming (summer and winter) since the late 1980s. During a short cooler period (1974–1994), the species numbers decreased and the upper elevational limits of some ground cover species descended. Thus, discernible responses, concurrent with both warming and cooling intervals, sustain a strong causal link between climate variability and alpine plant species richness. Methodologically, plot‐less revisitation studies of the present kind are beset with substantial uncertainties, which may overstate floristic changes over time. However, it is argued here that carefully executed and critically interpreted, no other method can equally effectively sense the earliest phases of plant invasions into alpine vegetation.     

Short-term variation in species richness across an altitudinal gradient of alpine summits

In response to climate warming, high altitude alpine vegetation may be replaced by typically lower altitude species, as species re-assemble and migrate to new areas. However, empirical evidence showing vegetation change in response to climate warming is largely unavailable for Australian alpine areas. Here, we examine changes in species richness with respect to climate and altitude over a 7?year period at a range of spatial scales in a re-survey of five alpine summits that are part of the Global Observation Research Initiative in Alpine Environments monitoring network. Eighty species were recorded in 2011 across all summits, an increase of 6 species since 2004. Mean species richness increased at the whole-of-summit scale from 45 to 50 species (about 12?%). At this scale, the rate of species richness increase was almost one new species per year, with 15 new species recorded at one summit. Here, shrub and graminoid species showed the largest increases. At the smaller spatial scales, changes in species richness were less pronounced. Turnover at the species and community level was typically moderate at all spatial scales and on all summits. The strength and direction of species richness change (the difference in species richness between the two sample periods, +/?) was not related to altitude nor variation in climate. Future re-surveys of the summits will confirm whether these short-term variations in species richness, particularly increases in shrubs, are indeed signals of longer-term trends and interactions with a changing climate.     

Long‐term geobotanical observations of climate change impacts in the Scandes of West‐Central Sweden

In the context of projected future human‐caused climate warming, the present study reports and analyses the performance of subalpine/alpine plants, vegetation and phytogeographical patterns during the past century of about 1 °C temperature rise. Historical baseline data of altitudinal limits of woody and non‐woody plants in the southern Scandes of Sweden are compared with recent assessments of these limits at the same locations. The methodological approach also includes repeat photography, individual age determinations and analyses of permanent plots. At all levels, from trees to tiny herbs, and from high to low altitudes, the results converge to indicate a causal association between temperature rise and biotic evolution. The importance of snow cover phenology is particularly evident. Treeline advance since the early‐20th century varies between 75 and 130 m, depending on species and site. Tendencies and potentials for further upshift in a near future are evident from the appearance of young saplings of all tree species, growing 400–700 m atop of the treeline. Subalpine/alpine plant species have shifted upslope by average 200 m. In addition, present‐day repetitions of floristic inventories on two alpine mountain summits reveal increases of plant species richness by 58 and 67%, respectively, since the early‐1950s. Obviously, many plants adjust their altitudinal ranges to new climatic regimes much faster than generally assumed. Nevertheless, plants have migrated upslope with widely different rates. This produces non‐analogous alpine plant communities, i.e. peculiar mixtures of alpine and silvine species. The alpine region is shrinking (higher treeline), and the character of the remaining alpine vegetation landscape is changing. For example, extensive alpine grasslands are replacing snow bed plant communities.     

Snowbeds are more affected than other subalpine–alpine plant communities by climate change in the Swiss Alps

While the upward shift of plant species has been observed on many alpine and nival summits, the reaction of the subalpine and lower alpine plant communities to the current warming and lower snow precipitation has been little investigated so far. To this aim, 63 old, exhaustive plant inventories, distributed along a subalpine–alpine elevation gradient of the Swiss Alps and covering different plant community types (acidic and calcareous grasslands; windy ridges; snowbeds), were revisited after 25–50 years. Old and recent inventories were compared in terms of species diversity with Simpson diversity and Bray–Curtis dissimilarity indices, and in terms of community composition with principal component analysis. Changes in ecological conditions were inferred from the ecological indicator values. The alpha‐diversity increased in every plant community, likely because of the arrival of new species. As observed on mountain summits, the new species led to a homogenization of community compositions. The grasslands were quite stable in terms of species composition, whatever the bedrock type. Indeed, the newly arrived species were part of the typical species pool of the colonized community. In contrast, snowbed communities showed pronounced vegetation changes and a clear shift toward dryer conditions and shorter snow cover, evidenced by their colonization by species from surrounding grasslands. Longer growing seasons allow alpine grassland species, which are taller and hence more competitive, to colonize the snowbeds. This study showed that subalpine–alpine plant communities reacted differently to the ongoing climate changes. Lower snow/rain ratio and longer growing seasons seem to have a higher impact than warming, at least on plant communities dependent on long snow cover. Consequently, they are the most vulnerable to climate change and their persistence in the near future is seriously threatened. Subalpine and alpine grasslands are more stable, and, until now, they do not seem to be affected by a warmer climate.     

Upward shift of alpine plants increases floristic similarity of mountain summits

Question: Does the upward shift of species and accompanied increase in species richness, induced by climate change, lead to homogenization of Alpine summit vegetation? Location: Bernina region of the Swiss Alps. Methods: Based on a data set from previous literature we expand the analysis from species richness to beta‐diversity and spatial heterogeneity. Species compositions of mountain summits are compared using a two‐component heterogeneity concept including the mean and the variance of Sørensen similarities calculated between the summits. Non‐metric multidimensional scaling is applied to explore developments of single summits in detail. Results: Both heterogeneity components (mean dissimilarity and variance) decrease over time, indicating a trend towards more homogeneous vegetation among Alpine summits. However, the development on single summits is not strictly unidirectional. Conclusions: The upward shift of plant species leads to homogenization of alpine summit regions. Thus, increasing alpha‐diversity is accompanied by decreasing beta‐diversity. Beta‐diversity demands higher recognition by scientists as well as nature conservationists as it detects changes which cannot be described using species richness alone.     

Effects of experimentally imposed climate scenarios on flowering phenology and flower production of subarctic bog species

Climate scenarios for high‐latitude areas predict not only increased summer temperatures, but also larger variation in snowfall and winter temperatures. By using open‐top chambers, we experimentally manipulated both summer temperatures and winter and spring snow accumulations and temperatures independently in a blanket bog in subarctic Sweden, yielding six climate scenarios. We studied the effects of these scenarios on flowering phenology and flower production of Andromeda polifolia (woody evergreen) and Rubus chamaemorus (perennial herb) during 2 years. The second year of our study (2002) was characterized by unusually high spring and early summer temperatures. Our winter manipulations led to consistent increases in winter snow cover. As a result, average and minimum air and soil temperatures in the high snow cover treatments were higher than in the winter ambient treatments, whereas temperature fluctuations were smaller. Spring warming resulted in higher average, minimum, and maximum soil temperatures. Summer warming led to higher air and soil temperatures in mid‐summer (June–July), but not in late summer (August–September). The unusually high temperatures in 2002 advanced the median flowering date by 2 weeks for both species in all treatments. Superimposed on this effect, we found that for both Andromeda and Rubus, all our climate treatments (except summer warming for Rubus) advanced flowering by 1–4 days. The total flower production of both species showed a more or less similar response: flower production in the warm year 2002 exceeded that in 2001 by far. However, in both species flower production was only stimulated by the spring‐warming treatments. Our results show that the reproductive ecology of both species is very responsive to climate change but this response is very dependent on specific climate events, especially those that occur in winter and spring. This suggests that high‐latitude climate change experiments should focus more on winter and spring events than has been the case so far.     

Impact of climate change on alpine vegetation of mountain summits in Norway

Climate change is affecting the composition and functioning of ecosystems across the globe. Mountain ecosystems are particularly sensitive to climate warming since their biota is generally limited by low temperatures. Cryptogams such as lichens and bryophytes are important for the biodiversity and functioning of these ecosystems, but have not often been incorporated in vegetation resurvey studies. Hence, we lack a good understanding of how vascular plants, lichens and bryophytes respond interactively to climate warming in alpine communities. Here we quantified long-term changes in species richness, cover, composition and thermophilization (i.e. the increasing dominance of warm-adapted species) of vascular plants, lichens and bryophytes on four summits at Dovrefjell, Norway. These summits are situated along an elevational gradient from the low alpine to high alpine zone and were surveyed for all species in 2001, 2008 and 2015. During the 15-year period, a decline in lichen richness and increase in bryophyte richness was detected, whereas no change in vascular plant richness was found. Dwarf-shrub abundance progressively increased at the expense of lichens, and thermophilization was most pronounced for vascular plants, but occurred only on the lowest summits and northern aspects. Lichens showed less thermophilization and, for the bryophytes, no significant thermophilization was found. Although recent climate change may have primarily caused the observed changes in vegetation, combined effects with non-climatic factors (e.g. grazing and trampling) are likely important as well. At a larger scale, alpine vegetation shifts could have a profound impact on biosphere functioning with feedbacks to the global climate.     

Studies of the alpine flora along an east-west gradient in central Western Norway

Central Western Norway was divided up into seven mountain areas, representing an east-west gradient from Hardangervidda (area A) westwards to the islands of Tysnes and Stord (area G). The alpine flora of totally 138 species thins out gradually where A has the greatest number (137 species) and G has the fewest (55 species). A drop of 83 species from east to west must be due to ecological barriers which prevent growth on the coastal mountains. Several factors have been taken into account and discussed. Rich Cambro-Ordovician rocks are present in all the explored areas, but the zone narrows towards the west due to the overlying base-poor thrusted rocks in the Caledonian nappe system. West of Hardangervidda, the summits of the mountains are almost entirely composed of acid rocks, and habitats for basiphilous species are therefore not present at the highest altitudes. Climate during the season of winter dormancy differs much more than during the growing season between east and west. Alpine plants require climatic conditions conducive to the termination of growth in autumn, a state of quiescence due to cold winters, and stable snow conditions which give a rapid transition from winter to summer. As the oceanic climate does not comply with these requirements, the number of alpine species declines. The alpine flora of Westem Norway is closely related to that of Great Britain. This is probably explained historically by a common origin during the Late Weichselian period.     

Foliar freezing resistance of Australian alpine plants over the growing season

We assessed the freezing resistance of leaves ex situ of 25 Australian alpine plant species. We compared the freezing resistance of forb, graminoid and shrub species from three alpine summits of different altitudes; from a low altitude site just above treeline, to a fully alpine tundra site. Foliar freezing resistance (LT₅₀) in spring varied from ?5.9°C to ?18.7°C and standardized LT₅₀ values within species were significantly related to site altitude. Additionally, when comparing all the species in the study, freezing resistance was significantly related to site; the LT₅₀ of samples from a low‐altitude summit (1696 m) were significantly lower than those of samples from mid‐ (1805 m) and high‐altitude (1860 m) summits. The LT₅₀ of juvenile foliage did not differ significantly from that of adult foliage. Shrubs were highly resistant to freezing. At the highest summit, we examined the course of seasonal freezing resistance from early summer to early autumn across three alpine plant communities that differed in the time of natural snowmelt; from sheltered (snowpatch) to exposed (open heath). No differences in freezing resistance over the growing season were detected for exposed or sheltered communities and there were no consistent trends indicating frost hardening over the growing season. Overall, the common Australian alpine species we investigated appear well adapted to freezing conditions throughout the snow‐free growing season. We have no evidence to suggest that freezing temperatures soon after snowmelt in spring are especially damaging to the alpine plants at these summits.     

Carbon sequestration in a high-elevation, subalpine forest

We studied net ecosystem CO₂ exchange (NEE) dynamics in a high‐elevation, subalpine forest in Colorado, USA, over a two‐year period. Annual carbon sequestration for the forest was 6.71 mol C m^?2 (80.5 g C m^?2) for the year between November 1, 1998 and October 31, 1999, and 4.80 mol C m^?2 (57.6 g C m^?2) for the year between November 1, 1999 and October 31, 2000. Despite its evergreen nature, the forest did not exhibit net CO₂ uptake during the winter, even during periods of favourable weather. The largest fraction of annual carbon sequestration occurred in the early growing‐season; during the first 30 days of both years. Reductions in the rate of carbon sequestration after the first 30 days were due to higher ecosystem respiration rates when mid‐summer moisture was adequate (as in the first year of the study) or lower mid‐day photosynthesis rates when mid‐summer moisture was not adequate (as in the second year of the study). The lower annual rate of carbon sequestration during the second year of the study was due to lower rates of CO₂ uptake during both the first 30 days of the growing season and the mid‐summer months. The reduction in CO₂ uptake during the first 30 days of the second year was due to an earlier‐than‐normal spring warm‐up, which caused snow melt during a period when air temperatures were lower and atmospheric vapour pressure deficits were higher, compared to the first 30 days of the first year. The reduction in CO₂ uptake during the mid‐summer of the second year was due to an extended drought, which was accompanied by reduced latent heat exchange and increased sensible heat exchange. Day‐to‐day variation in the daily integrated NEE during the summers of both years was high, and was correlated with frequent convective storm clouds and concomitant variation in the photosynthetic photon flux density (PPFD). Carbon sequestration rates were highest when some cloud cover was present, which tended to diffuse the photosynthetic photon flux, compared to periods with completely clear weather. The results of this study are in contrast to those of other studies that have reported increased annual NEE during years with earlier‐than‐normal spring warming. In the current study, the lower annual NEE during 2000, the year with the earlier spring warm‐up, was due to (1) coupling of the highest seasonal rates of carbon sequestration to the spring climate, rather than the summer climate as in other forest ecosystems that have been studied, and (2) delivery of snow melt water to the soil when the spring climate was cooler and the atmosphere drier than in years with a later spring warm‐up. Furthermore, the strong influence of mid‐summer precipitation on CO₂ uptake rates make it clear that water supplied by the spring snow melt is a seasonally limited resource, and summer rains are critical for sustaining high rates of annual carbon sequestration.     

Historical vs. ecological factors influencing global patterns of scarabaeine dung beetle diversity

Abstract. There has been much debate concerning the relative influence on biodiversity of historical vs. current ecological factors. Although both are important, we suggest that historical influences might be greater at higher taxonomic level, since one is looking further back into evolutionary history than at lower taxonomic level. Although we are unable to separate ecological from historical effects in the present global study on scarabaeine dung beetles, we are able to demonstrate differences in correlations between major environmental influences (climatic area, numbers of dung types) and major components of diversity (taxon richness, taxon diversity, functional composition) at different taxonomic levels (tribe, genus, species). Current global variation in taxon richness is correlated strongly to current biogeographical variation in the area of suitable climate at all three taxonomic levels. However, generic and species richness is correlated most strongly to climatic combinations which include tropical and warm summer rainfall climate types (I, II). In contrast, tribal richness is correlated most strongly to climatic combinations which include both warm summer rainfall and temperate climate types (II, VI, X). Regional variation in the number of available dung types shows a strong positive correlation to regional variation in taxon richness at higher tribal level but not at lower generic and species levels. Similarly, biogeographical differences in the number of available dung types show a strong negative correlation to dominance indices for taxon diversity at tribal level (distribution of generic numbers between tribes) but none at generic level (species numbers per genus). As functional diversification is linked closely to taxonomic diversification at tribal level, proportions of both ball‐rolling genera and ball‐rolling species also show strong negative correlations to the number of dung types available in each region. In conclusion, the presence of dung type correlations only at higher taxonomic level may reflect historical effects on scarabaeine taxon diversification, whereas differences in correlations to climate type with taxonomic level may reflect both current ecological and historical effects.     

Short-term signals of climate change along an altitudinal gradient in the South Alps

Short-term changes in plant species number, frequency and composition were studied along an altitudinal gradient crossing four summits from the treeline ecotone to the subnival zone in the South Alps (Dolomites, Italy). Large-scale (summit areas) and small-scale patterns (16 plots of 1 m2/summit) were monitored. After 5 years, a re-visitation of the summit areas revealed a considerable increase of species richness at the upper alpine and subnival zone (10% and 9%, respectively) and relatively modest increases at the lower alpine zone and the treeline ecotone (3% and 1%, respectively). At the small scale, the results were partly different, with species richness decreasing at the lower summits and increasing at the higher summits. The changes can most likely be attributed to climate warming effects and to competitive interactions. The main newcomers at the lower three summits were species from the treeline and the lower altitudinal zones. Only at the highest summit, the newcomers came from the alpine species pool. At the treeline ecotone, the abundance of Pinus cembra, of dwarf shrubs and clonal graminoid species increased. Here, displacements of alpine species may be predicted for the near future. At the higher summits, expansions of the established alpine species and further invasions of species from lower altitudes are forecasted.     

Effects of grazing abandonment and climate change on mountain summits flora: a case study in the Tatra Mts

Changes in the local flora of mountains are often explained by climate warming, but changes in grazing regimes may also be important. The aim of this study was to evaluate whether the alpine flora on summits in the Tatra Mts, Poland and Slovakia, has changed over the last 100 years, and if the observed changes are better explained by changes in sheep grazing or climate. We resurveyed the flora of 14 mountain summits initially investigated in the years 1878–1948. We used ordination methods to quantify changes in species composition. We tested whether changes in plant species composition could be explained by cessation of grazing and climate change, and whether these factors have influenced shifts in Ellenberg’s plant ecological indicator values and Raunkiaer’s life forms. Changes in alpine flora were greater on lower elevation summits, and lower on summits less accessible for sheep. More accessible summits were associated with a decrease in mean values of plant species’ light ecological indicator values over time, and a concurrent increase in temperature and nitrogen ecological indicator values. No significant relationships were found between accessibility for sheep and changes in Raunkiaer’s life-forms. Greater accessibility for sheep (meaning high historical grazing pressure) led to greater compositional changes of mountain summits compared with summits with low accessibility. Our results suggest that cessation of sheep grazing was the main factor causing changes in the species composition of resurveyed mountain summits in the Tatra Mts, while climate change played a more minor role.     

Latitudinal and altitudinal patterns of plant community diversity on mountain summits across the tropical Andes

The high tropical Andes host one of the richest alpine floras of the world, with exceptionally high levels of endemism and turnover rates. Yet, little is known about the patterns and processes that structure altitudinal and latitudinal variation in plant community diversity. Herein we present the first continental‐scale comparative study of plant community diversity on summits of the tropical Andes. Data were obtained from 792 permanent vegetation plots (1 m²) within 50 summits, distributed along a 4200 km transect; summit elevations ranged between 3220 and 5498 m a.s.l. We analyzed the plant community data to assess: 1) differences in species abundance patterns in summits across the region, 2) the role of geographic distance in explaining floristic similarity and 3) the importance of altitudinal and latitudinal environmental gradients in explaining plant community composition and richness. On the basis of species abundance patterns, our summit communities were separated into two major groups: Puna and Páramo. Floristic similarity declined with increasing geographic distance between study‐sites, the correlation being stronger in the more insular Páramo than in the Puna (corresponding to higher species turnover rates within the Páramo). Ordination analysis (CCA) showed that precipitation, maximum temperature and rock cover were the strongest predictors of community similarity across all summits. Generalized linear model (GLM) quasi‐Poisson regression indicated that across all summits species richness increased with maximum air temperature and above‐ground necromass and decreased on summits where scree was the dominant substrate. Our results point to different environmental variables as key factors for explaining vertical and latitudinal species turnover and species richness patterns on high Andean summits, offering a powerful tool to detect contrasting latitudinal and altitudinal effects of climate change across the tropical Andes.     

Diaspore traits discriminate good from weak colonisers on high-elevation summits

The richness of plant species in Swiss alpine-nival summits increased during the climate warming of the 20th century. Thirty-seven summits (2797–3418 m a.s.l.) with both old (～1900–1920) and recent (～2000) plant inventories were used to test whether biological species traits can explain the observed rates of summit colonisation. Species were classified into two groups: good colonisers (colonising five or more summits) and weak colonisers (fewer than five new summits). We compared species traits related to growth, reproduction and dispersal between these two groups and between the good colonisers and a group of high alpine grassland species. The observed colonisation pattern was subsequently compared with a simulated random colonisation pattern.The distribution of new species on the summits was not random, and 16 species exhibited a colonisation rate higher than expected by chance. Taraxacum alpinum aggr. and Cardamine resedifolia were the best colonisers. Results showed that diaspore traits enhancing long-distance dispersal were more frequent among good colonisers than among weak colonisers. Good colonisers were mostly characterised by pappi or narrow wings on their diaspores. Both groups were able to grow on soils more bare and rocky than species from the alpine grasslands. All other biological traits that we considered were similar among the three alpine species groups. These results are important for improving predictive models of species distribution under climate change.     

Vascular plant diversity and climate change in the alpine zone of the Lefka Ori,Crete

The aim of this study is to analyse the vascular flora and the local climate along an altitudinal gradient in the Lefka Ori massif Crete and to evaluate the potential effects of climate change on the plant diversity of the sub-alpine and alpine zones. It provides a quantitative/qualitative analysis of vegetation-environment relationships for four summits along an altitude gradient on the Lefka Ori massif Crete (1664–2339 m). The GLORIA multi-summit approach was used to provide vegetation and floristic data together with temperature records for every summit. Species richness and species turnover was calculated together with floristic similarity between the summits. 70 species were recorded, 20 of which were endemic, belonging to 23 different families. Cretan endemics dominate at these high altitudes. Species richness and turnover decreased with altitude. The two highest summits showed greater floristic similarity. Only 20% of the total flora recorded reaches the highest summit while 10% is common among summits. Overall there was a 4.96°C decrease in temperature along the 675 m gradient. Given a scenario of temperature increase the ecotone between the sub-alpine and alpine zone would be likely to have the greatest species turnover. Southern exposures are likely to be invaded first by thermophilous species while northern exposures are likely to be more resistant to changes. Species distribution shifts will also depend on habitat availability. Many, already threatened, local endemic species will be affected first.     

Experimental icing affects growth,mortality, and flowering in a high Arctic dwarf shrub

Effects of climate change are predicted to be greatest at high latitudes, with more pronounced warming in winter than summer. Extreme mid‐winter warm spells and heavy rain‐on‐snow events are already increasing in frequency in the Arctic, with implications for snow‐pack and ground‐ice formation. These may in turn affect key components of Arctic ecosystems. However, the fitness consequences of extreme winter weather events for tundra plants are not well understood, especially in the high Arctic. We simulated an extreme mid‐winter rain‐on‐snow event at a field site in high Arctic Svalbard (78°N) by experimentally encasing tundra vegetation in ice. After the subsequent growing season, we measured the effects of icing on growth and fitness indices in the common tundra plant, Arctic bell‐heather (Cassiope tetragona). The suitability of this species for retrospective growth analysis enabled us to compare shoot growth in pre and postmanipulation years in icing treatment and control plants, as well as shoot survival and flowering. Plants from icing treatment plots had higher shoot mortality and lower flowering success than controls. At the individual sample level, heavily flowering plants invested less in shoot growth than nonflowering plants, while shoot growth was positively related to the degree of shoot mortality. Therefore, contrary to expectation, undamaged shoots showed enhanced growth in ice treatment plants. This suggests that following damage, aboveground resources were allocated to the few remaining undamaged meristems. The enhanced shoot growth measured in our icing treatment plants has implications for climate studies based on retrospective analyses of Cassiope. As shoot growth in this species responds positively to summer warming, it also highlights a potentially complex interaction between summer and winter conditions. By documenting strong effects of icing on growth and reproduction of a widespread tundra plant, our study contributes to an understanding of Arctic plant responses to projected changes in winter climatic conditions.     

The influence of life history traits on the phenological response of British butterflies to climate variability since the late‐19th century

Many species of plants and animals have advanced their phenology in response to climate warming in recent decades. Most of the evidence available for these shifts is based on data from the last few decades, a period coinciding with rapid climate warming. Baseline data is required to put these recent phenological changes in a long‐term context. We analysed the phenological response of 51 resident British butterfly species using data from 83 500 specimens in the collections of the Natural History Museum, London, covering the period 1880–1970. Our analysis shows that only three species significantly advanced their phenology between 1880 and 1970, probably reflecting the relatively small increase in spring temperature over this period. However, the phenology of all but one of the species we analysed showed phenological sensitivity to inter‐annual climate variability and a significant advancement in phenology in years in which spring or summer temperatures were warm and dry. The phenologies of butterfly species were more sensitive to weather if the butterfly species was early flying, southerly distributed, and a generalist in terms of larval diet. This observation is consistent with the hypothesis that species with greater niche breadth may be more phenologically sensitive than species with important niche constraints. Comparison of our results with post‐1976 data from the UK Butterfly Monitoring Scheme show that species flying early in the year had a greater rate of phenological advancement prior to the mid‐1970s. Additionally, prior to the mid‐1970s, phenology was influenced by temperatures in March or April, whereas since 1976, February temperature had a stronger influence on the phenology. These results suggest that early flying species may be approaching the limits of phenological advancement in response to recent climate warming.     

The influences of snow cover,vegetation and topography on the upper range limit of common wombats Vombatus ursinus in the subalpine zone,Australia

Aim In subalpine and alpine environments, range shifts of species to higher altitudes are predicted to occur in response to reductions in the snow cover from climate change. However, the distribution of key resources may constrain the range of animal populations and prevent any upward migration. This study examined the local resource constraints on the upper range limit of a large, native herbivore in Australia, the common wombat Vombatus ursinus. Location The subalpine zone of the Snowy Mountains, Australia. Methods Logistic regression analyses of snow and habitat predictors were conducted on the presence/absence of wombat signs recorded along an altitudinal gradient during winter and summer, using parametric and nonparametric methods. Results Wombats responded strongly to the altitudinal gradient, but snow cover alone did not fully explain their upper range limit. Wombat occurrence in the subalpine zone was influenced by local habitat features in combination with maximum snow depth. More rugged, high‐relief terrain was important to wombats in winter, allowing individuals access to a wider range of altitudes, snow depths and shelter sites. During summer, high soil bulk density was an important predictor of occurrence, and in both seasons, occurrence declined in response to a higher cover of burnt grass. Main conclusions These models demonstrate that local habitat factors play a role even where there are strong regulating environmental factors. For wombats, this may limit future range expansion into the alpine zone despite the potential for an increase in abundance at their present range limit. These findings show the need for local ecological studies to be conducted in parallel with broad scale climate modelling if we are to understand shifts in species distributions as the climate rapidly changes.     

Effects of decadal climate change on zooplankton over the last 50 years in the western subarctic North Pacific

Decadal‐ to multi‐decadal variations have been reported in many regional ecosystems in the North Pacific, resulting in an increasing demand to elucidate the link between long‐term climatic forcing and marine ecosystems. We detected phenological and quantitative changes in the copepod community in response to the decadal climatic variation in the western subarctic North Pacific by analyzing the extensive zooplankton collection taken since the 1950s, the Odate Collection. Copepod species were classified into five seasonal groups depending on the timing of the annual peak in abundance. The abundance of the spring community gradually increased for the period 1960–2002. The spring–summer community also showed an increasing trend in May, but a decadal oscillation pattern of quasi‐30‐year cycles in July. Phenological changes coincided with the climate regime shift in the mid‐1970s, indicated by the Pacific decadal oscillation index (PDO). After the regime shift, the timing of the peak abundance was delayed one month, from March–April to April–May, in the spring community, whereas it peaked earlier, from June–July to May–June, in the spring–summer community, resulting in an overlap of the high productivity period for the two communities in May. Wintertime cooling, followed by rapid summertime warming, was considered to be responsible for delayed initiation and early termination of the productive season after the mid‐1970s. Another phenological shift, quite different from the previous decade, was observed in the mid‐1990s, when warm winters followed by cool summers lengthened the productive season. The results suggest that climatic forcing with different decadal cycles may operate independently during winter–spring and spring–summer to create seasonal and interannual variations in hydrographic conditions; thus, combinations of these seasonal processes may determine the annual biological productivity.