You are on page 1of 9

J. Nat. Conserv.

11, 1523 (2003) Urban & Fischer Verlag

Journal for

http://www.urbanfischer.de/journals/jnc

Nature Conservation

The sensitivity and vulnerability of terrestrial habitats and species in Britain and Ireland to climate change
Pam M. Berry*, Terry P. Dawson, Paula A. Harrison, Richard Pearson & Nathalie Butt
Environmental Change Institute, 1A Mansfield Road, Oxford, OX1 3SZ, UK; e-mail: pam.berry@eci.ox.ac.uk

Abstract
Climate change is having an increasing impact on the distribution and functioning of species and habitats. This has important implications for conservation practice and policy. The aim of this study was to model the direct impacts of climate change on terrestrial environments in Britain and Ireland in order to understand the possible changes in the distribution of species and the composition of habitats. A model, based on an artificial neural network, was used to predict changes in the bioclimate envelope of species, under the UKCIP98 climate change scenarios. A total of 50 species, representing several taxa, were modelled. Many species demonstrated a consistent response to climate change, either increasing or losing suitable climate space, although some had a variable response with losses starting to occur under the high scenarios. The percentage change in the bioclimate envelope of the species was calculated. This showed that montane species and habitats were the most sensitive to climate change. Other habitats from upland areas or species with northern distributions were also sensitive to losses, while species gaining suitable climate space represented a variety of habitats. Sensitivity needs to be viewed alongside vulnerability, the ability of the species or habitat to adapt to climate change. Montane species and habitats were the most vulnerable, with limited adaptation possibilities. Other vulnerable habitats, for which species modelling was carried out, include lowland raised bog, lowland calcareous grassland and native pine woodland. The potential impacts of climate change should be taken into account when planning conservation measures for these sensitive and vulnerable species and habitats. Key words: Adaptive capacity, artificial neural network, bioclimate envelopes, conservation.

Introduction
The earths climate has warmed by 0.6 C over the past century (IPCC 2001a), and recent studies show that it is possible to detect the effects of a changing climate on ecological systems (IPCC 2001b). Research on climate change impacts have demonstrated that many ecosystems, habitats and species are already being affected by climate change (Parmesan et al. 1999; Thomas & Lennon 1999; Both & Visser 2001; McCarty 2001; Thomas et al. 2001; Warren et al. 2001; Doran et al. 2002) and have the potential to be affected in the future (Grime et al. 2000; Berry et al. 2001a, b; Berry et al. 2002a). Changes in recent decades can be seen at all levels of ecological organisation: population and life-history changes, shifts in geographical range, changes in species composition of communities, and changes in the structure and functioning of ecosystems (McCarty 2001). It is no longer safe to assume that all of a species historic range will remain suitable.

*corresponding author

1617-1381/03/11/01-015 $ 15.00/0

16

P. M. Berry et al.

The objective of the MONARCH project (Modelling natural resource responses to climate change) was to evaluate the direct impacts of climate change on the nature conservation resources of Britain and Ireland and to assess the implications for conservation practice and policy (Hossell et al. 2003). Habitats of conservation interest were chosen and these formed the basis for selecting species for modelling. A total of 33 plant, four insect, two amphibian, ten bird (Harrison et al. 2003) and one mammal species of conservation importance, representing 12 habitats, were investigated. The results also help to address the need for the quantification of sensitivity, vulnerability and adaptive capacity of systems to climate change (IPCC 2001b).

Method
The SPECIESv1 (Spatial Estimator of Climate Impacts on the Envelope of Species), model was developed during the REGIS project (Regional climate change impact and response studies in East Anglia and North West) described elsewhere (Berry et al. 2001a, b; Pearson et al. 2002) and the version for birds is described by Harrison et al. (2003). It is based on an artificial neural network which integrates bioclimate variables for predicting the distribution of species through the characterisation of their climate space. The five input variables for all species, apart from birds, were: Accumulated annual soil water surplus Accumulated annual soil water deficit Absolute minimum temperature expected over a 20year period Annual maximum temperature Growing degree days above a base temperature of 5 C. The model was trained using existing empirical data on the European distributions of species to enable a wide climate space to be characterised, which includes the climatic range of the future scenarios. Once the model was trained, validated and tested at the European scale, it was then used to produce a map of the simulated distribution for current baseline climate and to estimate the potential re-distribution of a species suitable climate space under different UKCIP98 climate change scenarios (Hulme & Jenkins 1998) at a finer spatial resolution in Britain and Ireland. The output from the trained neural network was a suitability surface for each species for each climate change scenario showing the potentially available suitable climate space across Europe, and Britain and Ireland. The suitability surface was converted into a presence/absence map by assuming that all areas with probabilities greater than a calculated

kappa threshold value contained presences. The kappa statistic is used to test the similarity of distributions, in this case the actual distribution and the suitability surface (Monserud 1990). It was calculated for the different probability distributions for each species and the highest kappa was taken as the threshold value. In order to assess the sensitivity of species to climate change the percentage change in their suitable climate space was calculated, both as a percentage of the land area of Britain and Ireland in order to assess the significance of the species changes in an international context, and also as a percentage of its simulated current suitable climate space. The simulated current suitable climate space was used instead of actual distribution because a number of species are not in equilibrium with climate, for historical or land cover reasons, and thus the actual distribution does not represent the potential climate space of the species. The simulated climate space, however, is a model output and thus is directly comparable between the present and future time slices.

Results
The SPECIESv1 model results for the individual species are summarised in Table 1 (full details in Berry et al. 2001b). Species show an individualistic response (Huntley 1991; Huntley et al. 1995), but can be categorised into those that overall are gaining suitable climate space (winners), those losing suitable climate space (losers) and those showing little or no change (Berry et al. 2002a). For most species this is a consistent response but several of them show a variable response and this included some of the birds (Harrison et al. 2003). Blackstonia perfoliata (L.) Huds., for example, shows a total gain in suitable climate space under the Low scenarios and a loss under the High; Bufo calamita Laurenti loses under all scenarios apart from the 2050s High when it displays a potential gain. Several other species start to show a decreased net gain of suitable climate space under the certain scenarios. Dryopteris aemula (Ait.) Kuntze, for example, increasingly gains suitable climate space up until the 2020s High scenario and then experiences a decrease. Sensitivity of species and habitats Winners Sensitivity is defined as the degree to which a system is affected, either adversely or beneficially, by climate change (IPCC 2001b). The species showing the greatest gain in suitable climate space in Britain and Ireland

The sensitivity and vulnerability of terrestrial habitats and species in Britain and Ireland to climate change

17

Table 1. Summary of results from the SPECIES model. Habitat Montane heath Species losing climate space Species gaining climate space Comments All species show a loss of climate space. Most sensitive of the habitats modelled.

Erebia epiphron1 Carex bigelowii Salix herbacea Loiseleuria procumbens Trollius europaeus Geranium sylvaticum Orthilia secunda Sanguisorba officinalis Dryopteris aemula2 Parus montanus2 Sitta europaea2 Luscinia megarhynchos2 Cirsium acaulon2 Blackstonia perfoliata3 Helianthemum nummularium Motacilla flava Streptopelia turtur Erodium cicutarium3 Silene otites

Upland hay meadows Upland oak woodland

Likely to have changing species composition.

Blechnum spicant shows no change. Species of the herb layer likely to change.

Lowland calcareous grassland

All three plant species only show a small response.

Drought-prone acid grassland Beech woodland Pine woodland

Thought to be a relatively sensitive habitat and it is not known to what extent these species reflect the habitat response. Associated species Taxus baccata and Sanicula europaea show little change. Potentially sensitive habitat.

Fagus sylvatica2 Sciurus vulgaris3 Linnaea borealis Tetrao urogallus Gentiana pneumonanthe Coenonympha tullia Andromeda polifolia Rubus chamaemorus Eriophorum vaginatum Potamogeton filiformis Blysmus rufus Puccinellia maritima Haematopus ostragalus3 Equisetum variegatum Bufo calamita4 Artemesia norvegica1 Gavia stellata Rhynchospora alba Myrica gale2 Sphagnum papillosum2 Ophioglossum vaginatum Ranunculus scleratus2 Atriplex portulacoides

Wet heath Blanket/raised bogs

Robust habitat since dominant Erica tetralix shows no change. Currently widespread species show little change, but some elements of this habitat are sensitive. Variable species response. Results need to be combined with effects of coastal processes on habitat availability. Variable species response.

Wetlands Salt marshes

Coastal dune slacks No association


1 2

Epipactis palustris2 Ochlodes venata Coenagrion puella

Triturus cristatus shows little change.

Indicates that the species may become extinct. Indicates that species generally expands its suitable climate space, but suitability is lost from the southern and/or eastern limit of its distribution under severe climate change. 3 Indicates that species is predicted to lose suitable climate space from some part of its current distribution, but it simultaneously expands its range into other areas. 4 Indicates that species retreats under Low climate change scenarios and then expands under High scenarios.

18

P. M. Berry et al.

of Wales with scattered occurrences elsewhere. Under the climate change scenarios it displays a small loss of suitable climate space in the north of Scotland and a concomitant expansion from the south of England. 25% to < 50% gain 25% to <50% loss This unusual response, which has not been seen with any other species, is a function of its competition with Atriplex portulacoides Coenonympha tullia Sciurus caroliniensis L. This results in the European Erodium cicutarium Geranium sylvaticum distribution, on which the model is trained, showing it Ochlodes venata Loiseleuria procumbens as absent from much of England. This is, however, Rhynchospora alba Rubus chamaemorus suitable climate space and the climate change scenarSciurus vulgaris ios gradually pick this up. The distribution of this species, therefore, is not so sensitive to climate change, but rather to competition. The other two (Tables 2 and 3) is Atriplex portulacoides L. species showing over a 25% increase in suitable cli(Figure 1). This salt marsh plant is found around the mate space in Britain and Ireland, Erodium cicutarium coasts of England and Wales and the southern and east- (L.) LHrit. and Ochlodes venata Turati, could expand ern coasts of Ireland, with several occurrences in northwards under the various climate scenarios. Five species have the potential to more than double Dumfriesshire. Under the 2050s High scenario almost all the coast of Britain and Ireland represents suitable their currently potentially suitable climate space climate space. The second highest increase is for Rhyn- (Table 3). In the case of Silene otites L. Wibel and Bufo chospora alba L. Vahl. It is currently uncommon in calamita this is because they currently have only a raised peat bogs, but it could increase its range so that small simulated distribution. For the remainder it repreunder the 2020s High scenario most of Ireland and sents a significant expansion; for two of them (RhynchEngland, and all of Wales are suitable. Under the spora alba and S. vulgaris) this increase is also signifi2050s High scenario only the east of Scotland and cant at the scale of Britain and Ireland, although the north east England do not offer suitable climate space. caveat above concerning the Sciurus vulgaris results Sciurus vulgaris L. is the third most sensitive modelled should be noted. Fagus sylvatica L. gains suitable clispecies. In Britain it is currently confined almost en- mate space in the northern part of Great Britain, while tirely to Scotland, northern England and higher parts losing it from the south and east of England (Berry et al. 2002b), with the net effect being a greater than a 100% increase in suitable climate Table 3. Sensitivity of species with changes in their suitable climate space for the space. The species that could have a 50% to 2050s High scenario expressed as a percentage of their currently suitable climate 75% gain in their suitable climate space space. (Erodium cicutarium and Ochlodes venata) a) Gains also are amongst the highest percentage gains at the Britain and Ireland scale.
50% to <75% 75% to <100% None 100% +

Table 2. Sensitivity of species with changes in their suitable climate space for the 2050s High scenario expressed as a percentage of the area of Britain and Ireland.

Erodium cicutarium Ochlodes venata

Atriplex portulacoides Bufo calamita Fagus sylvatica Rhynchospora alba Sciurus vulgaris Silene otites

b) Losses 50% to <75% 75% to <100% 100%

Carex bigelowii Equisetum variegatum Salix herbacea

Andromeda perfoliata Geranium sylvaticum Linnaea borealis Loiseleuria procumbens Orthilia secunda Rubus chamaemorus Trollius europaeus

Erebia epiphron

Losers The most negatively affected species is Erebia epiphron Knoch, which loses all suitable climate space under the High scenarios. Currently it only occurs in the Lake District and higher parts of Scotland and thus it does not cover a significant part of Britain and Ireland and does not show up in Table 2. The species exhibiting more than a 25% loss of potentially suitable climate space in Britain and Ireland all tend to be found in northern and/or upland areas, thus highlighting their potential sensitivity to climate change. The most sensitive species in terms of percentage losses in current potentially suitable climate space is Erebia epiphron, which has already been seen to lose all

The sensitivity and vulnerability of terrestrial habitats and species in Britain and Ireland to climate change

19

suitable climate space under the High scenarios. The other montane heath species and species with northern distributions, such as Loiseleuria procumbens (L.) Desv. (Figure 2), Trollius europaeus L. (Figure 3) and Orthilia secunda (L) House, are the next most sensitive species (Table 3). All the montane heath species lose over half their currently suitable climate space. Habitats effects If the species are related back to the habitats that they were chosen to represent (Tables 1 to 3), then montane heaths emerge as the most sensitive habitat to climate

change, since all the species modelled for this habitat lose suitable climate space. They are all species with a northern distribution in Britain and Ireland. On other habitats the species modelled show a variable response and thus it is more difficult to ascribe an overall sensitivity value to the habitat. However, upland hay meadows, pine woodland, upland oak woodland and beech woodland could be considered the next most sensitive. In upland hay meadows, for example, Geranium sylvaticum L. and Trollius europaeus are predicted to lose suitable climate space, although previous modelling work has shown that Anthoxanthum odoratum

Figure 1. SPECIES model results for Atriplex portulacoides L.: (a) current climate (19611990); (b) 2020s Low scenario; (c) 2020s High scenario; (d) 2050s Low scenario; (e) 2050s High scenario.

Figure 2. SPECIES model results for Loiseleuria procumbens (L.) Desv.: (a) current climate (19611990); (b) 2020s Low scenario; (c) 2020s High scenario; (d) 2050s Low scenario; (e) 2050s High scenario.

20

P. M. Berry et al.

L. would not lose climate space (Berry et al. 2001a). In pine and upland oak woods too, the dominant (tree) species are unlikely to lose suitable climate space (Berry et al. 2001b), but it is the associated ground flora that might do so. In the case of beech woodland, however, it is the dominant species, Fagus sylvatica, that is adversely af-

fected, in southern and eastern England. If Fagus sylvatica is lost from these areas due to factors such as higher temperatures, summer drought and lack of winter chilling then significant changes in canopy composition could occur (Berry et al. 2002b). The modelled species associated with it, Taxus baccata L. and Sanicula europaea L., however, show little change.

Figure 3. SPECIES model results for Trollius europaeus L.: (a) simulated current distribution (19611990); (b) 2020s Low scenario; (c) 2020s High scenario; (d) 2050s Low scenario; and (e) 2050s High scenario.

Figure 4. SPECIES model results for Cirsium acaulon (L.) Scop.: (a) current climate (19611990); (b) 2020s Low scenario; (c) 2020s High scenario; (d) 2050s Low scenario; (e) 2050s High scenario.

The sensitivity and vulnerability of terrestrial habitats and species in Britain and Ireland to climate change

21

In none of the habitats did all the modelled species show a net gain in climate space. In calcareous grassland two species showed a net gain and Blackstonia perfoliata increased under the Low scenarios. However, two of the calcareous grassland species (Cirsium acaulon (L). Scop. and Helianthemum nummularium L.) start to lose suitable climate space just from the south of their range, which suggests that they are sensitive either to the predicted temperature increases and/or to the increased soil moisture deficits.

Discussion
The modelling of the changing suitable climate space of species has shown that there are species specific, and thus habitat specific, responses to climate change. As the magnitude of the climate change increases so the number of systems affected and the geographical extent of losses will increase (IPCC 2001b). Under the UKCIP98 scenarios a number of species show a consistent response with the changes in their climate space increasing with the magnitude of the change. Several species, however, show a variable response and as the magnitude increases, so the extent of climate space gains may decrease, even to the point of a net loss as in the case of Blackstonia perfoliata. More recent predictions of global climate change (IPCC 2001a; Hulme et al. 2002) suggest that even greater warming may be experienced in the future, so the results presented here may be conservative. The SPECIESv1 model outputs only show changes in suitable climate space, they do not explicitly address how or whether this might be fulfilled. Retractions in the suitable climate space for a species, as is the case for the montane heath species and some upland hay meadow species, are likely to be realised. These may, however, occur with a time lag, dependent on the ability of the species to withstand the stresses resulting from the unfavourable climate and the pressure from competitors. Where a species is shown to have an expansion in suitable climate space, the realisation of this will depend on the migration ability of the species involved and the availability of intervening habitat. Silene otites and Cirsium acaulon are plants with southern distributions, which could experience an increase in potentially suitable climate space, although nothing is known of their dispersal abilities, while the insects modelled that could expand their range, Ochlodes venata and Coenagrion puella L., could start to realise this, given their potential mobility. The latter has been shown to be able to disperse at least 860 m in a season (Conrad et al. 1999), although this would not be sufficient to fill its potentially suitable climate space in the time span predicted. Species mobility should be en-

couraged in order that they may realise their future climate space (Hossell et al. 2003). Vulnerability was not a direct outcome of the MONARCH results but research by Hossell et al. (2000) produced expert-based vulnerability matrices for habitats and species and these will be used as the basis for an assessment of habitat vulnerability. Vulnerability is defined as the degree to which a system is susceptible to, or unable to cope with, adverse effects of climate change. It is a function of the character, magnitude and rate of climate change to which a system is exposed, its sensitivity and its adaptive capacity (IPCC 2001b). The vulnerability of the modelled species will depend on their ability to realise their new climate space. Those species that are losing climate space are amongst the most vulnerable, because this loss is more likely to be realised and it will lead to fragmented habitats and decreased populations. For example, Erebia epiphron could become extinct in Britain and Artemesia norvegica Fr. would be severely threatened. Studies on the possible effects of climate change on Erebia epiphron on Ben Lawers have shown that rises of 1 C to 2 C could reduce current habitat by up to 40% (French 1995). Vulnerability can also occur where there will be little or no overlap between the current and future distribution, as is the case with Bufo calamita. For species that have the opportunity to expand their climate space, the issue is their ability to migrate and the availability of suitable habitat. This will be more difficult for species, such as some of the lowland calcareous grassland plants, which have specific habitat requirements that are not easily met in new areas further north. Species, such as Cirsium acaulon, which lose climate space in the southern part of Britain and Ireland, while gaining it further north, will probably not be vulnerable to direct climate change, providing there is an adequate overlap with their current distribution. It could, however, be vulnerable through a lack of habitat availability for migration northwards. Montane heaths have been identified as the most sensitive habitat and they are also considered highly vulnerable as they are restricted in extent and have limited possibility of adapting to climate change (Hossell et al. 2000). They may also be adversely affected by increasing levels of nitrogen deposition (Baddeley et al. 1994). The other habitat that is considered highly vulnerable and for which species were modelled, is lowland raised bog (Hossell et al. 2000). It could be susceptible to future summer drying, especially in south eastern England and East Anglia (Dawson et al. 2003). Although many of the species associated with it would continue to find suitable climate space, some, such as

22

P. M. Berry et al.

Eriophorum vaginatum L., could lose it under the high scenarios in these areas. Native pine woodland and lowland calcareous grassland were considered to be of medium-high vulnerability (Hossell et al. 2000). Climate change could lead to the spread of pine woodland to higher altitudes, aided by an increased growth rate for Pinus sylvestris L. (Hossell et al. 2000), but the modelled species associated with it (Linnaea borealis L. and Tetrao urogallus) could lose suitable climate space. The modelled species for lowland calcareous grassland showed a generally positive response to climate change, in terms of their climate space, but other work has shown that this habitat is sensitive to temperature increases (Duckworth et al. 2000). Climate change could lead to forbs replacing grasses as conditions become drier. The other potentially sensitive habitats: upland hay meadows, upland oak woodland and beech woodland have medium vulnerability. The adaptive capacity of a system is its ability to adjust to climate change to moderate potential damages, to take advantage of opportunities, or to cope with the consequences (IPCC 2001b). Species and natural and semi-natural habitats will, to some degree, adapt autonomously to climate change, although management (planned adaptation) can provide opportunities for their protection. In the case of Erebia epiphron little probably can be done in situ to prevent such loss, but in less severe instances maintaining or improving habitat quality can help species survival. In upland hay meadows, for example, the survival of species, such as Trollius europaeus, could be facilitated by the maintenance of traditional hay making practices. For the most vulnerable species translocations are an option, but these are timeconsuming and require good knowledge of the autecology of a species and the availability of suitable habitats in appropriate areas. They have already been undertaken for Bufo calamita; six new populations were established in the 1980s (Banks & Beebee 1987; Denton & Beebee 1994; Rowe et al. 1998). The conservation of habitats and their maintenance in appropriate condition is essential for species trying to migrate. There are few real options in the case of montane species and habitats, as opportunities for migration to higher altitudes ultimately are limited. Lowland raised bogs also have limited adaptation capacity, as they will be negatively affected by summer drought, although it might be possible to manage water resources at a few sites of high conservation importance. Native pine woodland is currently affected by fragmentation and poor regeneration as a result of grazing. The lowering of grazing pressure could help natural regeneration, while the use of Pinus sylvestris seedlings of a more southerly provenance could help overcome

some of the adverse effects of climate change by maintaining the dominant species in this habitat, but the habitats overall species composition is likely to change. Lowland calcareous grassland species could spread north into similar grasslands in the uplands, but in the lowlands they are restricted by geology. The possibility for natural adaptation in these areas, therefore, is low, though there is limited potential of re-creation of lowland calcareous grassland from arable uses (Hossell et al. 2000). Overall, therefore, they are moderately vulnerable with a limited adaptive capacity. The species and habitat specific response has been seen to apply also to sensitivity, vulnerability and adaptive capacity. Montane heaths and their species are of high conservation concern on all these accounts and are, therefore, of high conservation priority. However, given their limited adaptive capacity it may be necessary in the long-term to accept some loss and to focus resources on other sensitive and vulnerable habitats where some mitigation of climate change impacts may be possible. These results can be used to inform conservation policy and practice, and to develop appropriate adaptation strategies and management options for vulnerable species and habitats (Hossell et al. 2003).
Acknowledgements: The SPECIES model was developed in the REGIS project (Regional climate change impact and response studies in East Anglia and North West England), which was jointly funded between the UKs MAFF, DETR (now DEFRA) and UKWIR. MONARCH was funded by a consortium of government and non- government nature conservation organisations, led by English Nature and we would like to thank them for their support. We would also like to thank Raino Lampinen, University of Helsinki, who supplied some of the European plant distributions from the Atlas Flora Europaea in electronic format, the Biological Records Centre who supplied some British data and Florence Miller for her help with the modelling.

References
Baddeley JA, Thompson DBA & Lee JA (1994) Regional and historical variation in the nitrogen content of Racomitrium lanuginosum in Britain in relation to atmospheric nitrogen deposition. Environmental Pollution 84: 189196. Banks B & Beebee TJC (1987) Factors influencing breeding site choice by the pioneering amphibian Bufo calamita. Holarctic Ecology 10: 1421. Berry PM, Dawson TP, Harrison PA & Pearson RG (2001a) Integrated impacts on biodiversity. In: Regional Climate Impact Studies in East Anglia and North West England (eds Loveland P & Holman I). Final Report to MAFF,

The sensitivity and vulnerability of terrestrial habitats and species in Britain and Ireland to climate change

23

DETR & UKWIR, Soil Survey and Land Research Centre, Silsoe. Berry PM, Vanhinsbergh D, Viles HA, Harrison PA, Pearson RG, Fuller R, Butt N & Miller F (2001b) Impacts on terrestrial environments. In: Climate Change and Nature Conservation in the Britain and Ireland: Modelling natural resource responses to climate change (the MONARCH project) (eds Harrison PA, Berry PM & Dawson TP) UKCIP Technical Report, Oxford. Berry PM, Dawson TP, Harrison PA & Pearson RG (2002a) Modelling potential impacts of climate change on the bioclimatic envelope of species in Britain and Ireland. Global Ecology and Biogeography 11: 453-462. Berry PM, Dawson TP, Harrison PA & Pearson RG (2002b) Impacts on native woodland dynamics and distribution. In: Climate change and UK forests (ed Broadmeadow MSJ): 169180. Forestry Commission Bulletin 124 Forestry Commission, Edinburgh. Both C & Visser ME (2001) Adjustment to climate change is constrained by arrival date in a long-distance migrant bird. Nature 411: 296298. Conrad KF, Willson KH, Harvey IF, Thomas CJ & Sherratt TN (1999) Dispersal characteristics of seven odonate species in an agricultural landscape. Ecography 22: 524531. Dawson TP, Berry PM & Kampa E (2003) Climate change impacts on freshwater wetland habitats. Journal for Nature Conservation 11: 2530. Denton JS & Beebee TJC (1994) The basis of niche separation during terrestrial life between two species of toad (Bufo bufo and Bufo calamita): Competition or specialisation? Oecologia 97: 390398. Doran PT, Priscu JC, Berry Lyons W, Walsh JE, Fountain AG, McKnight DM, Moorhead DL, Virginia RA, Wall DH, Clow GD, Fritsen CH, McKay, Andrew N, Parsons CP (2002) Antarctic climate cooling and terrestrial ecosystem response. Nature 415: 517520. Duckworth JC, Bunce RGH & Malloch AJC (2000) Modelling the potential effects of climate change on calcareous grasslands in Atlantic Europe. Journal of Biogeography 27: 347358. French DD (1995) Use of GIS to predict effects of environmental change on the small mountain ringlet at Ben Lawers. Institute of Terrestrial Ecology. Unpublished report to Scottish Natural Heritage. Grime JP, Brown VK, Thompson K, Masters GJ, Hillier SH, Clarke IP, Askew AP, Corker D & Kielty JP (2000) The response of two contrasting limestone grasslands to simulated climate change. Science 29: 762765. Harrison PA, Vanhinsbergh D, Fuller RJ & Berry PM (2003) Modelling climate change impacts on the distribution of breeding birds in Britain and Ireland. Journal for Nature Conservation 11: 3142. Hossell JE, Ellis N, Harley M & Hepburn IR (2003) Climate change and nature conservation: Implications for policy and practice in Britain and Ireland. Journal for Nature Conservation 11: 6773. Hossell JE, Briggs B & Hepburn IR (2000) Climate Change and Nature Conservation: A Review of the Impact of Cli-

mate Change on UK Species and Habitat Conservation Policy. HMSO, DETR, MAFF, London. Hulme M & Jenkins GJ (1998) Climate Change Scenarios for the United Kingdom: Scientific Report. UK Climate Impacts Programme Technical Report No. 1, Climatic Research Unit, Norwich, 80 pp. Hulme M, Jenkins GJ, Lu X, Turnpenny JR, Mitchell TD, Jones RG, Lowe J, Murphy JM, Hassell D, Boorman P, McDonald R & Hill S (2002) Climate Change Scenarios for the United Kingdom: The UKCIP02Scientific Report. Tyndall Centre for Climate Change Research, School of Environmental Sciences, University of East Anglia, Norwich, UK 120 pp. Huntley B (1991) How plants respond to climate change: migration rates, individualism and the consequences for plant communities. Annals of Botany 67: 1522. Huntley B, Berry PM, Cramer W & Mc Donald AP (1995) Modelling present and potential future ranges of some European higher plants using climate response surfaces. Journal of Biogeography 22: 9671001. IPCC (2001a) Climate Change 2001: The Scientific Basis. Intergovernmental Panel on Climate Change, Cambridge University Press, Cambridge. IPCC (2001b) Climate Change 2001: Impacts Adaptation and Vulnerability. Summary for Policymakers. Intergovernmental Panel on Climate Change, Cambridge University Press, Cambridge. McCarty JP (2001) Ecological consequences of recent climate change. Conservation Biology 15: 320326. Monserud RA (1990). Methods for Comparing Global Vegetation Maps. WP-90-40, IIASA, Luxembourg. Parmesan C, Ryrholm N, Stefanescu C, Hill JK, Thomas CD, Descimon H, Huntley B, Kaila L, Kullberg J, Tammaru T, Tennent WJ, Thomas JA & Warren M (1999) Poleward shifts in geographical ranges of butterfly species associated with regional warming. Nature 399: 579583. Pearson RG, Dawson TP, Berry PM & Harrison PA (2002) SPECIES: a Spatial Evaluation of Climate Impact on the Envelope of Species. Ecological Modelling 154: 289300. Rowe G, Beebee TJC & Burke T (1998) Phylogeography of the natterjack toad Bufo calamita in Britain: Genetic differentiation of native and translocated populations. Molecular Ecology 7: 751760. Thomas CD & Lennon J (1999) Birds extend their range northwards. Nature 399: 213214. Thomas CD, Bodsworth EJ, Wilson RJ, Simmons AD, Davies ZG, Musche M & Conradt L (2001) Ecological and evolutionary processes at expanding range margins. Nature 411: 577581. Warren MS, Hill JK, Thomas JA, Asher J, Fox R, Huntley B, Roy DB, Telfer MG, Jeffcoate S, Harding P, Jeffcoate G, Willis SG, Greatorex-Davies JN, Moss D & Thomas CD (2001) Rapid responses of British butterflies to opposing forces of climate and habitat change. Nature 414: 6569. Received 31. 03. 02 Accepted 30. 09. 02