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Methodology Statement Generalist species more able to cope than specialist species: snow-bed species Indicator Frequency of specialist and generalist species in snow-bed communities Indicator type: Risk/opportunity Impact Action X SCCAP Theme SCCAP Objective CCRA risk/opportunity Natural Environment N2: Support a healthy and diverse natural environment with capacity to adapt. Generalist species more able to adapt than specialists Author(s) Chris Ellis, Royal Botanic Garden Edinburgh. Acknowledgements Dave Genney, SNH. Why are snow-bed species of interest? A British vegetation type which is strongly associated with areas of late-lying snow occurs in the Scottish mountains, and is described in the UK’s National Vegetation Classification [1] as comprising three community-types: (a) U8, Carex bigelowii-Polytrichum alpinum sedgeheath; (b) U11, Polytrichum sexangulare-Kiaeria starkei snow-bed; (c) U12 Salix herbaceaRacomitrium heterostichum snow-bed. These three communities are priority features for conservation; they are sub-types nested within the broad Annex I habitat ‘Siliceous alpine and boreal grasslands - 6150’ [2], and form an important constituent within protected sites such as the Cairngorms Special Area of Conservation [3]. The snow-bed vegetation is characterised by mosses and liverworts (bryophytes) which are ‘snow-bed specialists’ and might be considered as indicator species for the habitat [4]. The snow-bed bryophyte community includes BAP/priority species such as Andreaea nivalis and Marsupella arctica, and the Schedule 8 species Gymnotrium apiculatum. Information on the status of snow-bed bryophytes would provide a valuable assessment of the condition of snow-bed vegetation, while also being representative of wider challenges in biodiversity and climate change: (1) Snow-bed species are niche specialists, and monitoring their status is especially relevant during a period when observational data indicates an increasing homogenisation of the mountain flora, through an expansion of generalists [5,6]; (2) Conceptually, snow-bed species occur at a latitudinal and altitudinal ‘range edge’ in Scotland, capturing a scenario in which species are unable to track climate change through migration along a continuous environmental gradient via inter-connected habitat. (3) Areas of late-lying snow provide ecosystem services in addition to their unique biodiversity, for example by sustaining the baseflow of streams and rivers following the main period of snowmelt [7]. References [1] Averis, A., Averis, B., Birks, J., Horsfield, D., Thompson, D. & Yeo, M. (2004) An Illustrated Guide to British Upland Vegetation. Joint Nature Conservation Committee, Peterborough. [2] JNCC Habitat Account – 6150, Siliceous alpine and boreal grasslands. http://jncc.defra.gov.uk/protectedsites/sacselection/habitat.asp?FeatureIntCode=H6150 (accessed 14th June 2013) [3] JNCC Cairngorms SAC Account. http://jncc.defra.gov.uk/protectedsites/sacselection/sac.asp?EUCode=UK0016412 (accessed 14th June 2013) [4] Woolgrove, C.E. &Woodin, S.J. (1994) Relationships between the duration of snowlie and the distribution of bryophyte communities within snowbeds in Scotland. Journal of Bryology, 18: 253-260. [5] Britton, A.J., Beale, C.M., Towers, W. &Hewison, R.L. (2009) Biodiversity gains and losses: evidence for homogenisation of Scottish alpine vegetation. Biological Conservation, 142, 1728-1739. [6] Ross, L.C., Woodin, S.J., Hester, A.J., Thompsom, D.B.A. & Birks, H.J.B. (2012) Biotic homogenization of upland vegetation: patterns and drivers at mutiple scales over five decades. Journal of Vegetation Science, 23, 755-770. [7] Dunn, S.M., Langan, S.J., Colohan, R.J.E. (2001) The impact of variable snow pack accumulation on a major Scottish water resource. The Science of the Total Environment, 265: 181-194. What are the key climate hazard(s) affecting snow-bed species? Snow-bed vegetation and associated specialist species may be dependent on late-laying snow for a variety of reasons [1], for example: (a) intolerance of extreme low temperatures, (b) a demand for soil moisture throughout the growing season, (c) specific adaptation for physiological activity during an ephemeral period of snowmelt and an associated nutrient pulse, or (d) reduced competition from co-occurring species which require a longer growing season. The link between the number of late-lying snow-beds (e.g. beyond July [2]) and their size, and the climate, are broadly established, being dependent on winter and spring temperatures, and factors of wind and topography affecting drift [2]. Alongside warming temperatures, UKCP09 scenarios project a decline in mean snowfall rate of 65-80 % over montane areas generally by the 2080s [3], with regional assessments also strongly suggesting a reduction in snow cover across altitudinal bands in Scotland [4].These climate projections underpin bioclimatic models which indicate a loss of suitable niche space for specialist snow-bed species and a transition to generalists [5]. It is expected that a long-term trend towards a reduction in the spatial extent (snow-bed size) and length of snow-lie (snow-bed duration) will shift snow-bed vegetation composition, with conservation-priority snow-bed communities transitioning towards a widespread community-type typical of more open montane conditions. Nevertheless, given the potential variety of ecological factors controlling each species’ association with and distribution within snow-beds [1,6], the signature of change away from a snow-bed community may not be as simple as a linear decline across all snow-bed associated species. References [1] Björk, R.G. &Molau, U. (2007) Ecology of alpine snowbeds and the impact of global change. Arctic, Antarctic and Alpine Research, 39: 34-43. [2] Watson, A., Davison, R.W. & French, D.D. (1994) Summer snow patches and climate in northeast Scotland, U.K. Arctic and Alpine Research, 26: 141-151. [3] Jenkins, G., Murphy, J.M., Sexton, D.M.H., Lowe, J.A., Jones, P. &Kilsby, C. (2010) UK Climate Projections: Briefing Report. Met Office. [4] Trivedi, M.R., Browne, M.K., Berry, P.M., Dawson, T.P. &Morecroft, M. D. (2007) Projecting climate change impacts on mountain snow cover in Central Scotland from historical patterns. Arctic, Antarctic and Alpine Research, 39: 488-499. [5] Trivedi, M.R., Morecroft, M.D., Berry, P.M. & Dawson, T.P. (2008) Potential effects of climate change on plant communities in three montane nature reserves in Scotland, UK. Biological Conservation, 141: 1665-1675. [6] Woolgrove, C.E. &Woodin, S.J. (1994) Relationships between the duration of snowlie and the distribution of bryophyte communities within snowbeds in Scotland. Journal of Bryology, 18: 253-260. What is the exposure of snow-bed species to climate change? A comprehensive survey of snow-bed vegetation was carried out in 1989-1990, for 58 areas of late-lying snow across Scotland [1]. The area of Scotland’s snow-bed vegetation is expected to total c. 160-250 ha across these and other isolated patches. Scottish Natural Heritage commissioned a repeat survey of the original 58 snow-beds during 2007-2008, and 22 were successfully resurveyed (Fig. 1, Table 1); these were located across western and central mountain ranges, and within the eastern and relatively more continental Cairngorm Mountains [2,3]. A report to SNH compared the composition of bryophyte communities, and the frequency of species, between the two survey periods [2,3]. The results demonstrated only very tentative evidence for change in snow-bed communities [2] and a shift in the frequency of certain species. Notably, this included a statistically significant 13% decline in the frequency of the snow-bed specialist liverwort Moerckia blytii [3]. However, trends in frequency shifts were not consistent across the snow-bed bryophytes, with some species showing a decrease (e.g. Kiaeria starkei: -4%) and some an increase (Polytrichum sexangulare: +11 %) [3]. This lack of a clear trend may demonstrate individualistic species responses consistent with a reorganisation of snow-bed vegetation structure, or may reflect the fact that the repeat sampled plots were not strictly replicates, but were resampled for equivalent vegetationtypes across the time-periods. For the purpose of future monitoring, permanent plots have been established. For the purposes of this indicator, the focus was less on species composition, and instead on broad measures of vegetation structure: vascular plants, with grasses as a specific example guild, and bryophytes generally, with NVC snow-bed species as indicators. Accumulated cover values within these groups were analysed separately for western/central and eastern snow-beds (Fig. 1) for those plots within which a NVC dominant snow-bed bryophyte had been recorded in 1989-1990: (a) Polytrichum alpinum (b) Polytrichum sexangulare (c) Kiaeria starkei (d) Racomitrium heterostichum This takes into account possible decadal-scale differences in the rate, magnitude and intraannual direction of climate change between these regions [4]. Analysis demonstrated a decline in the frequency of snow-bed NVC indicators and bryophytes generally, for the western snow-beds, implying structural change in the vegetation (Fig. 2). In contrast, cover values for bryophytes increased across the Cairngorm snow-beds (Fig. 2). There is preliminary evidence for structural change in the snow-bed vegetation, which may show regional variation. When available Snow Survey of Great Britain data will be provided in this section. References [1] Rothero, G.P. (1991) Bryophyte-dominated snow-beds in the Scottish Highlands. Unpublished MSc thesis, University of Glasgow. [2] Rothero, G.P., Grytnes, J.-A., Birks, H.J.B. & Genney, D. (2008) Effects of climate change on bryophyte-dominated snowbed vegetation. Scottish Natural Heritage 104 pp. [3] Rothero, G., Birks, J., Genney, D., Grytnes, J.-A. & Long, D. (2011) Climate change and its consequences on bryophyte-dominated snowbed vegetation. The Changing Nature of Scotland (ed. by S.J. Marrs, S. Foster, C. Hendrie, E.C. Mackey and D.B.A. Thompsom), pp. 435-440. TSO Scotland, Edinburgh. [4] Jenkins, G., Murphy, J.M., Sexton, D.M.H., Lowe, J.A., Jones, P. &Kilsby, C. (2010) UK Climate Projections: Briefing Report. Met Office. What are the key factors affecting the vulnerability of snow-bed species to climate change? There is direct vulnerability of snow-bed associated species if climate change causes a decline in montane snow cover with impacts extending to areas of late-lying snow. There are various sources of information addressing temporal variability in the extent of snowcover in the Scottish mountains [1]. These data have been collected for a variety of purposes, and would require careful meta-analysis to provide a synthesis. Recently, Met Office snow survey data for a period 1945-2007 has been digitised [1], and will become publically available on request [2], with recommendations for improving the quality of snow survey information into the future [1]. In addition to direct impacts on late-lying snow, a range of additional existing pressures can be identified: 1. Air pollutants can accumulate at high levels in the late-lying snow-pack, leading to physiological damage for sensitive snow-bed bryophytes [3,4]; 2. Human activity can result in localised pollution impacts through waste [5], or trampling of the sensitive vegetation. References [1] Spencer, M. (2012) Scottish snowline observations – the past and the future. BHS 11th National Symposium, Dundee. http://www.hydrology.org.uk/assets/2012%20papers/Spencer_49.pdf (accessed 17th June 2013) [2] Met Office Integrated Data Archive System (MIDAS) Land and Marine Surface Stations Data (1853-current). http://badc.nerc.ac.uk/view/badc.nerc.ac.uk__ATOM__dataent_ukmo-midas (accessed 17th June 2013) [3] Woolgrove, C.E. &Woodin, S.J. (1996) Effects of pollutants in snowmelt on Kiaeriastarkei, a characteristic species of late snowbed bryophyte dominated vegetation. New Phytologist, 133: 519-529. [4] Woolgrove, C.E. &Woodin, S.J. (1996) Ecophysiology of a snow-bed bryophyte Kiaeriastarkei during snowmelt and uptake of nitrate from meltwater. Canadian Journal of Botany, 74: 1095-1103. [5] Cairngorm Mountain Ranger Service – ‘Keep Cairngorm Snow White’. http://www.cairngormmountain.org/sites/default/files/Snow%20White%20Project_0.pdf (accessed 14th June 2013) as reported on the BBC: http://news.bbc.co.uk/1/hi/scotland/highlands_and_islands/7170227.stm (accessed 14th June 2013). Are there any current or planned adaptation actions to address the climate change risk to snow-bed species? Currently adaptation to address climate change risk is focussed on policy objectives. For example, bryophyte assemblages are a notified feature of the Eastern Cairngorms SSSI, and a component of this feature (snow-bed bryophytes) were considered to be in unfavourable condition during the last round of Site Condition Monitoring (SCM), possibly as a direct consequence of climate change. The condition of snow-bed bryophytes will be assessed over several cycles of SCM, and if further decline in condition is confirmed as a consequence of external pressures (climate change) it could trigger removal from the SSSI citation. This is an example of a policy adaptation consistent with managed retreat. Alternative adaptation options include the reduction of other environmental pressures (e.g. reducing direct human impacts through the reduction of waste, see above), through to experimental infrastructure projects, including: (a) snow-fences to encourage drifting and retain late-lying snow, as demonstrated in an experimental framework[1,2], (b) textile blankets as used in the Alps to prevent glacier melt[3], or (c) artificial snow blowing to augment natural deposits. Currently, none of these options are under consideration as a formal approach to conservation. References [1] Scott, P.A. & Rouse, W.R. (1995) Impacts of increased winter snow cover on upland tundra vegetation: a case example. Climate Research, 5: 25-30. [2] Wipf, S. &Rixen, C. (2010) A review of snow manipulation experiments in Arctic and alpine tundra ecosystems. Polar Research, 29: 95-109. [3] ‘Researchers cover up melting glaciers’ – as reported in USA Today: http://usatoday30.usatoday.com/weather/climate/2005-07-17-glacier-blankets_x.htm (accessed 14th June 2013). Supporting graphics Fig. 1: The location of 22 repeat-survey snow-beds for which data was made available by Scottish Natural Heritage. Snow-beds were grouped into two regions for analysis here: Region 1 (red box), with snow-beds 1.-8., representing more western and central mountain ranges, and Region 2 (blue circle), with snow-beds 9.-22., representing the eastern and relatively more continental Cairngorm snow-beds (cf. Table 1). Table 1. Location data for 22 snow-bed survey sites (cf. Fig. 1) for which vegetation data was made available by Scottish Natural Heritage. Region Site Code SNH Reference Site Name Location # plots S1. S2. S3. S4. S5. S6. S7. S8. 33 30 34 44 45 54 57 58 Beinn Dearg Ben Wyvis, North Coire Mam Sodhail, Lochan Uaine White Mounth, Coire Boidheach White Mounth, Glas Allt Creag Meagaidh, West Coire Ben Lawers, An stuc Ben Lawers, NE Face NH256815 NH466686 NH120255 NN233845 NN241846 NN408871 NN637433 NN636413 4 4 1 5 3 1 2 1 S9. S10. S11. S12. S13. S14. S15. S16. S17. S18. S19. S20. S21. S22. 1 3 19 4 5 6 17 18 13 14 7 8 9 10 Cairngorm, Ciste Mhearad Carn Lochan, Coire Domhain Carn Lochan, Coire Domhain Ben Macdui, Lower Garbh Uisge Beag Ben Macdui, Upper Garbh Uisge Beag Ben Macdui, North Slope Ben MacDui, Upper Garbh Uisge Mor Ben MacDui, Bealach 1232m Braeriach, Garbh Coire Mor Braeriach, Garbh Coire Mor E Gully Beinn a Bhuird, Coire an t'Snaechda Beinn a Bhuird, Dubh Lochan Beinn a Bhuird, North Top Beinn a Bhuird, Garbh Coire NJ011045 NH992023 NH986100 NH994008 NH999004 NH990001 NN994992 NN999989 NN940980 NN942983 NO094979 NO091991 NJ095009 NJ104016 18 11 9 10 6 4 10 1 4 2 5 3 7 4 Western Eastern Fig. 2: Time-series comparison (1989/90-2007/8) in the percent cover of four vegetation parameters – (i) grasses, (ii) all vascular plants (including ferns), (iii) NVC snow-bed dominant bryophytes (Kiaeria starkei, Polytrichum alpinum, Polytrichum sexangulare and Racomitrium heterostichum), and (iv) all bryophytes. Statistical values are for a comparison between sets of paired plots within each of the two contrasting regions (western and eastern), using a Wilcoxon paired sample test. Eastern Snow-beds (sites 9.-22., n = 83 plots) Western Snow-beds (sites 1.-8., n = 20 plots) 80 60 40 20 0 -20 -40 V = 173, P = 0.008 N VC s ry op hy te s Al lb hy te ts br yo p pl an ed sn ow -b Va sc ul ar ra ss es V = 367, V = 1386, V = 1357, V = 584, P = 0.97 P = 0.99 P = 0.154 P < 0.001 G s ry op hy te s hy te Al lb br yo p pl an ed sn ow -b Va sc ul ar ra ss es -80 ts V = 37.5, V = 77, V = 142, P = 0.6 P = 0.99 P = 0.061 N VC -60 G Change in percent cover (1989/90 - 2007) 100