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Ecological Indicators
journal homepage: www.elsevier.com/locate/ecolind
Review
The future of upland water ecosystems of the UK in the 21st century: A synthesis
Chris J. Curtis a,b, , Richard W. Battarbee b , Donald T. Monteith c , Ewan M. Shilland b
a
School of Geography, Archaeology & Environmental Studies, University of the Witwatersrand, Private Bag 3, Wits 2050, Johannesburg, South Africa Environmental Change Research Centre, University College London, Gower Street, London WC1E 6BT, UK c NERC Centre for Ecology & Hydrology (Lancaster), Lancaster Environment Centre, Library Avenue, Bailrigg, Lancaster LA1 4AP, UK
b
a r t i c l e
i n f o
a b s t r a c t
Upland waters are located upstream of the areas of direct human disturbance and intensive land use through industry, agriculture and urbanisation and are valued particularly as sources of potable water and as biodiverse freshwater habitats. Nonetheless, the impact of human activity can be detected even in some of the most remote lakes on the planet due to the effects of long-range transport of air pollutants and global climate change. In the UK, upland waters are threatened by a range of pressures including not only atmospheric deposition of acidic compounds, trace metals and organic pollutants, but also climate change, nutrient enrichment by deposited nitrogen and changing land-use or land management regimes. The threat from acid deposition has declined sharply since the 1980s but its legacy remains a major concern; recovery is taking place but even assuming a complete chemical and biological recovery to a pre-industrial baseline were possible, dynamic models suggest that this could take another 100 years. However, current climate change projections suggest that by then, UK upland waters will become much warmer, with lower summer streamows, higher winter streamows and a much reduced or even absent inuence of snowfall and lake ice-cover. Hence there is no rationale for aspiring to the return of pre-industrial reference conditions because climate change is likely to shift the climatic baseline at an unprecedented rate. Meanwhile, expansion of forest planting, changing grazing regimes and management measures for peatland restoration, among other land-use pressure, will affect upland catchments in the UK with potentially major repercussions for aquatic ecosystems, both positive and negative. The challenge for scientists, policymakers and other stakeholders in the uplands is to determine not only the trajectory of change in upland waters, but also to dene the ecological endpoints needed to provide the optimum range of ecosystem services for society. Integrated monitoring of the kind exemplied by the papers in this volume is a fundamental prerequisite in this regard. 2013 Elsevier Ltd. All rights reserved.
Article history: Received 28 July 2013 Received in revised form 25 September 2013 Accepted 8 October 2013 Keywords: Uplands Freshwater ecosystems Acidication Climate change Toxic substances Nutrients Land-use
Contents 1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.1. Upland waters and environmental change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2. Upland waters: values and pressures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Future threats to upland waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1. Air pollution and upland waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1.1. Acid deposition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1.2. Nutrient enrichment (N deposition) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1.3. Trace metals and POPs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2. Land-use change in the uplands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2.1. Forestry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2.2. Moorland management (burning, drain blocking) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2.3. Grazing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 00 00 00 00 00 00 00 00 00 00 00 00
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Corresponding author at: School of Geography, Archaeology & Environmental Studies, University of the Witwatersrand, Private Bag 3, Wits 2050, Johannesburg, South Africa. Tel.: +27 11 717 6505; fax: +27 11 717 6529. E-mail addresses: christopher.curtis@wits.ac.za, c.curtis@ucl.ac.uk (C.J. Curtis). 1470-160X/$ see front matter 2013 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.ecolind.2013.10.012
Please cite this article in press as: Curtis, C.J., et al., The future of upland water ecosystems of the UK in the 21st century: A synthesis. Ecol. Indicat. (2013), http://dx.doi.org/10.1016/j.ecolind.2013.10.012
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Climate change impacts on upland waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.1. Direct impacts of climate change in the uplands: hydrology, temperature and aquatic ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.2. Climate change impacts on catchment biogeochemical cycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synthesis and recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1. The legacy of acid deposition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2. Emerging pressures on upland waters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3. Shifting baselines and novel recovery endpoints . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4. The requirements for integrated monitoring of upland waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2.3.
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1. Introduction 1.1. Upland waters and environmental change In his book The End of Nature, McKibben (1990) evocatively describes the human need to believe that there are still parts of our planet unspoiled by human activities, true wilderness areas where the imprint of human activity has not yet reached. He quotes George Orwells encouraging thought, from The Road to Wigan Pier; In spite of hard trying, man has not yet succeeded in doing his dirt everywhere. . .. Perhaps if you looked for them you might nd streams with live sh in them instead of salmon tins. Orwell might have been less encouraged to learn that human impacts would in fact extend to the most remote parts of the planet well before the end of the 20th century. Stratigraphic changes in the sediments of lakes, even very remote ones, have been proposed as one of the key indicators of the new, human-dominated geological epoch termed the Anthropocene (e.g. Zalasiewicz et al., 2008; Holmgren et al., 2010; Hobbs et al., 2010; Holtgrieve et al., 2011). Indeed, remote upland lakes have been proposed as ideal sentinel ecosystems for large scale global change, due to their physical isolation from local human impacts (Battarbee et al., 2002; Schindler, 2009; Mueller et al., 2009). Furthermore, the smallest headwater streams in upland areas may have a far greater role in biogeochemical cycling than previously recognised and are often poorly represented on maps (Benstead and Leigh, 2012). The dearth of information on not only the functional role but even the extent of rst order headwater streams led Bishop et al. (2008) to coin the term Aqua incognita the unknown waters. Visually at least, it may seem that the streams and lakes of upland areas still represent relatively pristine ecosystems, unharmed by human activity, but in fact several decades of research have demonstrated that this is not the case. Many papers and indeed several journal special issues have been dedicated to the subject of environmental change in remote, upland lakes and streams (e.g. Battarbee et al., 2002; Psenner et al., 2002; Livingstone, 2005; Catalan et al., 2009; Kernan et al., 2009; Tetzlaff et al., 2013a). The best known of these impacts is acidication from long-distance transported air pollution, but there are other threats to the health of aquatic ecosystems in the uplands, including eutrophication from nitrogen deposition, contamination by toxic metals and trace organic compounds as well as from land-use change and climate change (Battarbee et al., 2009). Upland waters in the UK have been affected by most if not all of these disturbances and are far from pristine. We dene them as those situated beyond the limit of enclosed agricultural land, draining predominantly moorland or heathland catchments, corresponding largely to the mountain, moorland and heath (MMH) category of the UK National Ecosystem Assessment (Van der Wal et al., 2011). With very low human occupancy in their catchments, they are generally not subjected to the direct pressures of urban efuent inputs or fertiliser runoff from agricultural land experienced by many water bodies in more densely populated lowland regions.
1.2. Upland waters: values and pressures MMH habitat covers around 18% of the UK (Fig. 1a) and represents the great majority of its semi-natural habitats (Van der Wal et al., 2011). Due to their low fertility, these areas are used for livestock grazing and eld sports (deer hunting and grouse shooting) although they are also increasingly being used for tourism and recreation, providing important cultural landscapes with iconic imagery central to the landscape quality of many National Parks and Areas of Outstanding Natural Beauty in the UK. Major ecosystem services linked to the role of MMH areas in the water cycle include: 1. provisioning services: 68% of the UKs drinking water supply is estimated to come from surface waters (Van der Wal et al., 2011) for which MMH areas are the most important sources; 2. regulating services: these include water purication (i.e. buffering against the effects of atmospheric, diffuse and point source pollution, since MMH soils and biota retain a large proportion of deposition inputs), carbon storage (since around 40% of all UK soil carbon is found in MMH soils; Bradley et al., 2005; Van der Wal et al., 2011), carbon sequestration and ood regulation; 3. cultural services, which include aesthetic values of MMH landscapes, outdoor pursuits, archaeology, biodiversity and geological diversity. In particular, MMH habitats contribute signicantly to the biological and genetic diversity of the UKs native ora and fauna. Many upland waters are located in areas with national or international conservation designations, such as Sites of Special Scientic Interest, National Nature Reserves, RAMSAR sites, International Biosphere Reserves, Special Areas of Conservation (EU Habitats Directive) and Special Protection Areas (EU Birds Directive). A large number are Biodiversity Action Plan priority habitats (e.g. rivers and streams, oligotrophic and dystrophic lakes), host priority species (e.g. otter, freshwater pearl mussel, common scoter) and support a substantial proportion of important salmonid sh populations including sensitive spring salmon stocks. Despite their relative distance from centres of population all upland waters in the UK have, to a greater or lesser extent, been modied by human activity and are vulnerable to a range of pressures, including: elevated, although declining, levels of acid (sulphur and nitrogen) deposition which have caused widespread acidication of upland waters, with subsequent increases in dissolved organic carbon as surface waters recover from acidication; potential increases in nitrate leaching from catchment soils leading to acidication and/or eutrophication; climate change, directly through changes in temperature, precipitation and wind strength, and indirectly through alterations to catchment biogeochemistry (especially the carbon and nitrogen
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Fig. 1. (a) Extent of MMH habitat within the UK predominantly uplands but also including very fragmented lowland heaths (used with permission from the United Nations Environment Programme World Conservation Monitoring Centre and NERC. This material is based upon Crown Copyright and is reproduced with the permission of Land & Property Services under delegated authority from the Controller of Her Majestys Stationery Ofce, Crown copyright and database rights, EMOU206.2). (b) Exceedance of acidity critical loads for freshwaters for 198688 using a critical ANC value of 20 equiv. L1 , showing wide extent of critical load exceedance during the 1980s (Source: Jane Hall, CEH Bangor). The dataset comprises a grid-based national survey of the most sensitive headwater lakes or streams in each 10 km square (20 km in non-sensitive lowland regions), combined with later regional surveys from some of the most impacted upland areas (Curtis et al., in press-a). Note the close correspondence between the distribution of exceeded sites and MMH habitat.
cycles), soil erosion and pollutant behaviour (specically trace metals and POPs); land-use and land management change related to changes in forestry practices, animal husbandry and arable expansion, associated with socio-economic and environmental pressures, potentially causing changes in acidity, nutrient status, thermal regime and soil erosion; and invasive species, both aquatic and riparian. These have been recognised as potential threats to upland water ecosystems in other parts of the world, e.g. North America (Rahel and Olden, 2008) and present a threat in the UK but as yet there is no evidence for their presence in UK upland waters. The spatial extent of potential impact from one of the biggest historical threats, acid deposition, is best expressed in terms of the exceedance of critical loads for acidity (Fig. 1b). Critical load models for surface waters employ empirically based chemical relationships between hydrochemistry and deposition along with mass balance principles to determine the maximum load of acid deposition that will not cause a critical chemical threshold to be crossed (Henriksen and Posch, 2001). For the UK the critical chemical threshold is a mean annual acid neutralising
capacity of 20 equiv. L1 for the great majority of sites (both lakes and streams), linked to a 90% probability of undamaged brown trout populations (Lien et al., 1996; Curtis et al., 2000, in press-a). Exceedance of critical loads indicates that at long-term steady state between deposition and surface waters, ANC will decline below the critical value (Fig. 1). Critical loads data for UK surface waters feed into the international integrated assessment modelling efforts carried out under the auspices of the UN ECE Convention on Long Range Transboundary Air Pollution and the UK is one of only ve countries which currently submit freshwater critical loads (Curtis et al., in press-a). Almost all of the most sensitive water bodies modelled in the UK are located in MMH habitats. Here we consider the evidence from this Special Issue and elsewhere for continuing and changing threats to the chemical and biological quality of upland freshwater ecosystems and their structure and function. We identify knowledge gaps and data requirements to assist policymakers, landscape and environmental managers in protecting upland water ecosystems and maintaining their healthy functioning for the benets of both biodiversity and the human populations downstream.
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Fig. 2. Schematic diagram showing environmental pressures on upland waters (climate change, air pollution and land-use) and their key interactions. All pressures ultimately impact on macronutrient cycling, structure and function of terrestrial and aquatic ecosystems, and affect the form and export uxes of macronutrients as well as pollutants from catchments.
2. Future threats to upland waters Most of the pressures or drivers identied above interact with each other and hence are not independent in their effects on upland water ecosystems and their catchments (Fig. 2). For example, changes in rainfall, temperature, storminess and cloud cover affect photochemical reactions as well as the relative balance between dry, occult and wet deposition of pollutants. Climate and land-use factors (e.g. overgrazing, commercial forestry) will affect catchment hydrology and vulnerability to soil erosion. Air pollution and land use or land cover may interact, for example by increasing pollutant deposition due to scavenging of air pollutants by forests, or by causing a reduction in cover of certain plants such as bryophytes and lichens due to acid deposition or excess nutrient inputs (Curtis et al., 2005). 2.1. Air pollution and upland waters Since upland waters occur upstream of the most intensive agricultural and industrial activity, contamination of their catchments is usually dominated by atmospherically transported pollutants including sulphur and nitrogen compounds and toxic substances. Recently, industrially derived hydrochloric acid has also been shown to have made a signicant contribution to the atmospheric pollution load over much wider areas of the uplands than previously thought (Evans et al., this volume). Nitrogen deposition may act as an agent of acidication and/or of nutrient enrichment of naturally nutrient poor ecosystems in the uplands; it is therefore considered in both potential roles below. 2.1.1. Acid deposition Levels of acid deposition to upland waters have declined greatly in the UK and many parts of Europe and North America, but the problem has not disappeared. Major reductions in acid deposition in the UK over the last 2040 years have been described in the recently published RoTAP Report (2012) as well as the paper by Curtis and Simpson (this volume). Sulphur emissions decreased by 94% from a 1970 peak up to 2010, with the UK meeting its target under the EU National Emission Ceilings Directive (NECD) (RoTAP, 2012). Across the UK as a whole, resulting decreases in total sulphur deposition from the start of monitoring in 1986 to 2006 amounted to 80%, comprising a 93% reduction in dry deposition and a 57% decrease in wet deposition. Dry deposition shifted from the
dominant component of total S deposition in the 1980s to a minor component by 2008 (RoTAP, 2012). Reductions in non-marine sulphate in bulk deposition appear to be the main cause of increasing rainfall pH since the late 1990s (Curtis and Simpson, this volume) and while not always linear, show a broadly consistent decline. Since wet deposition has decreased less than dry deposition, and because total deposition reductions tend to be largest closer to emission sources in central and southeast England, the smallest reductions in total S deposition are found in the uplands where wet deposition is dominant (RoTAP, 2012; Curtis and Simpson, this volume). Emissions of oxidised nitrogen (NOx ) decreased by 58% between 1970 and 2010, meeting the NECD target for 2010 (RoTAP, 2012). Much smaller reductions in emissions of ammonia (NH3 ) were achieved from a later baseline, just 21% between 1990 and 2010. Reductions in total deposition of nitrogen have been much smaller than for sulphur, with only a 13% decline between 1988 and 2008, despite emission reductions of 50% for NOx and 18% for NH3 over the same period. The non-linearities between emissions and deposition of nitrogen compounds are poorly understood, but are thought to be related to changes in chemical processing of NOx in the atmosphere (RoTAP, 2012). Temporal patterns in the bulk deposition of nitrate and ammonium across the UK uplands include periods of both signicant increases and decreases over the monitoring period, with few sites showing signicant, monotonic trends (Curtis and Simpson, this volume). Responses in the chemistry and ecology of upland waters to the decreases in acid deposition are described in a number of the accompanying papers in this special issue and summarised in Battarbee et al. (this volume-a). Indicators of chemical recovery include major reductions in non-marine sulphate, reductions in labile aluminium and increases in ANC, while changes in nitrate are more heterogeneous and dissolved organic carbon (DOC) levels are increasing (Fig. 3: Monteith et al., this volume). There has been considerable debate about the reasons for increasing DOC, but the decline in acid deposition is known to be a major factor (Monteith et al., 2007; Evans et al., 2012). As a result of the chemical improvement, aquatic plant and animal communities are now beginning to recover. Benthic diatom populations show statistically signicant trends away from acid tolerant taxa (Flower et al., 2010). New aquatic plant species have appeared in seven of the lake sites and ve of the stream sites in the Acid Waters Monitoring Network. However, species composition at ten sites has not changed signicantly during the 20 year monitoring period, including some sites with signicant improving trends in surface water chemistry (Shilland and Monteith, 2010). Benthic invertebrate changes at most sites remain fairly modest but community composition has changed signicantly at about half of the sites in the Network (Murphy et al., this volume). For some sites the invertebrate community has yet to respond to chemical improvement. Salmonid sh populations are also showing some signs of improvement, with labile aluminium being the single best predictor of the presence of fry (Malcolm et al., this volume). Dynamic modelling of the water chemistry of sites in the Acid Waters Monitoring Network has provided valuable insights into the baseline or reference chemical conditions at these sites as well as future predictions of timescales of recovery (Helliwell et al., this volume-a, this volume-b). The degree of recovery in ANC at each AWMN site is shown relative to the MAGIC-modelled reference ANC value (using 1860 as the pre-industrial reference year) in Fig. 4. In order to control for effects of short-term variation, three year means are used for the initial (generally 19891991) and most recent (20082010) three years of monitoring (cf. Kernan et al., 2010). While all sites show an increase in ANC towards the reference value over the period of monitoring, recent average ANC values at most sites still fall short of the reference value (Fig. 4).
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Fig. 3. Annual mean DOC concentrations in nine AWMN lakes (19892011). Reductions in concentrations since 2006 largely reect a return from particularly elevated levels in 2006 following a drought. The longer term trend remains signicantly upward.
Diatom data strongly support these results. Comparison of lake diatom species composition in recent sediment trap samples with pre-acidication diatom assemblages from sediment cores shows that while there has been some recovery in acid-sensitive diatom species (as noted above from epilithon samples), current assemblages are still very different from the diatom assemblages that occurred in the acidied lakes prior to the rst signs of acidication in the 19th century (Battarbee et al., this volume-b). The patterns of chemical and biological recovery observed in upland waters of the UK are consistent with broader scale patterns around Europe and North America reported for long term monitoring sites within the ICP Waters Programme (Stoddard et al., 1999; Skjelkvle et al., 2005; Skjelkvle and De Wit, 2011). The evidence
for only modest recovery from acidication in the UK is consistent with national scale modelling of critical loads for ve north-west European countries that shows critical load exceedances will still be widespread following the full implementation of the Gothenburg Protocol by 2020, including recent revisions to the Protocol in 2012 (Fig. 5, Curtis et al., in press-a). Hence it is clear that while some improvement in water quality and, to a lesser degree, biology has occurred in response to large reductions in acid deposition over the last 20 years, upland lakes and streams in the most impacted and acid-sensitive regions of the UK and other northern European countries will remain acidied into the foreseeable future. It is likely that over the long term (decades) only a global switch away from fossil fuels as a major
Fig. 4. Initial (c. 19891991, measured), recent (20082010, measured) and reference (1860, MAGIC modelled) values of acid neutralising capacity equiv. L1 in sites of the UK Acid Waters Monitoring Network with continuous recent data (after Kernan et al., 2010). Note that initial data do not reect the peak of acidication in the UK which occurred some years prior to the onset on monitoring. See Battarbee et al. (this volume-a) for further details of AWMN sites. Sites are ordered by decreasing reference ANC values.
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nitrate concentrations are apparent in upland waters monitored as part of the UK Acid Waters Monitoring Network (Monteith et al., this volume) suggesting potential impacts on productivity in these systems, particularly in high deposition areas (Curtis and Simpson, 2011). Contemporary studies of nutrient limitation in upland lakes have employed nutrient enrichment and bioassay methods to show that on average, freshwater phytoplankton are as commonly N-limited as P-limited and addition of both nutrients typically produces the strongest response (Maberly et al., 2002; Curtis and Simpson, 2007, 2011; Elser et al., 2007; Lewis and Wurtsbaugh, 2008). These studies include 43 upland lakes and 3 upland streams in the UK (Maberly et al., 2002; Curtis and Simpson, 2007, 2011). Bergstrm and Jansson (2006) suggested that in the northern hemisphere, atmospheric N deposition has driven many lakes from a natural N-limited state towards P-limitation: i.e. prior to increases in anthropogenic N inputs via atmospheric deposition, most lakes in the northern hemisphere could have been N-limited. This idea has been supported by more recent experimental work (Bergstrm et al., 2008; Elser et al., 2009a,b; Liess et al., 2009) and observations of lake chemistry and its relationship with foliar N:P ratios in trees in catchments of the northeastern US (Crowley et al., 2012). On a global scale, upland waters have been instrumental in revealing the potential effects of N deposition as an agent of nutrient enrichment in surface waters (Lepori and Keck, 2012), through both historical and contemporary studies of aquatic ecosystem change. The historical impact of increased N deposition has been demonstrated in palaeolimnological studies from remote lakes in North America and the Arctic, generally employing two strands of evidence: 1. natural abundance of stable isotopes of N (15 N) as an indicator of anthropogenic N deposition (e.g. Holtgrieve et al., 2011); 2. corresponding changes in planktonic diatom communities linked to nutrient enrichment (Wolfe et al., 2001, 2003, 2006; Saros et al., 2003, 2005, 2011; Hobbs et al., 2010). Similar changes have been observed in upland lakes in the UK, both from nutrient limitation studies and from palaeolimnological studies of 15 N trends (Fig. 6, Curtis and Simpson, 2007, 2011). Analysis of bulk sediment 15 N in cores from 19 upland lakes found a decline in 15 N in recent decades (Curtis and Simpson, 2011), as observed in boreal and arctic lakes by Holtgrieve et al. (2011). While other studies have attributed the decline in sediment 15 N at high latitudes to increasing inputs of isotopically light N deposition, studies comparing spatial variation in 15 N of surface sediments in UK upland lakes with N deposition uxes and bulk deposition 15 N were equivocal. Highly signicant, though weak, relationships were found between the surface sediment 15 N of 83 upland lakes and deposition of N species, with NOx deposition showing the strongest relationship (Curtis and Simpson, 2007). However, a later study directly comparing bulk deposition and sediment trap organic matter 15 N at 12 sites found no signicant relationships (Curtis and Simpson, 2011). Hence it seems likely that while down-core changes in bulk sediment 15 N in lakes are indicative of changes in biogeochemical processing of N, and that this is also likely to be driven at least in part by N deposition, there is no simple relationship between sediment 15 N and either the magnitude or isotopic signature of deposited N. Site specic factors play a major role in determining the 15 N of lake sediment organic matter. Further work is therefore required to determine the extent to which, for a given site, temporal increases in N deposition may be responsible for observed decreases in lake sediment 15 N. The extent to which there have been related changes in phytoplankton and aquatic macrophyte communities is currently being investigated. Previous studies investigating the potential inuence
Fig. 5. Average accumulated exceedance (AAE) of critical loads of acidity for surface waters in north-west Europe for S and N depositions in the year 2020 as projected under the revised (2012) Gothenburg Protocol. Grey shading indicates 10 km grid cells containing surface waters which will not exceed critical loads in 2020; other colours represent levels of exceedance which are linked to long term acidication and the suppression of ANC below a critical value, for example ANCcrit = 20 equiv. L1 for most of the UK. AAE is a function of the number of water bodies within a grid cell (selected on differing criteria for each country see Curtis et al. (in press-a)), their catchment areas and individual site exceedances see Posch et al. (2001) for further explanation. (For interpretation of the references to color in this gure legend, the reader is referred to the web version of the article.) Data and gure provided by Max Posch, CCE, The Netherlands.
energy source will allow the acidity of these systems to decline to levels conducive for the sustainable re-establishment of some of the UKs most acid-sensitive aquatic species. Furthermore, chronic base cation depletion of upland soils (Helliwell et al., this volume-a, this volume-b) may persist for centuries, while other drivers of change (e.g. climate) are likely to prevent a complete return to preindustrial biological assemblages and ecological functioning. 2.1.2. Nutrient enrichment (N deposition) Whereas most emphasis over the last few decades has been on the acidication potential of N deposition, there has been a growing concern about the role of N deposition in promoting the eutrophication of upland waters (e.g. De Wit and Lindholm, 2010). This potential effect was discounted on the assumption that primary production in oligotrophic waters was phosphorus (P) limited. However, within the last decade, this prevailing paradigm (e.g. Schindler, 1977) has been challenged, with much debate in the literature about the relative roles of nitrogen (N) and P (Sterner, 2008; Schindler et al., 2008; Howarth and Paerl, 2008; Schindler and Hecky, 2008). There is at least some agreement that both N and P can contribute to eutrophication and control of both nutrients will in many cases be required (Conley et al., 2009). Elevated
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Fig. 6. (a) Small Water in the English Lake District, with the drinking water supply reservoir Hawes Water in the background (Photo: C.J. Curtis). (b) Sediment core analysis shows a strong decline in the stable isotope 15 N at Small Water in recent decades suggestive of anthropogenic N inputs (data from G.L. Simpson; Curtis and Simpson, 2011).
of N deposition and climate on planktonic and benthic diatoms or chrysophytes found that both drivers appeared to contribute to changes in the sediment record, but were inconclusive in attributing their relative importance (Curtis et al., 2009; Pla et al., 2009; Simpson and Anderson, 2009). For example, the upland tarn Small Water in the English Lake District is not acid-sensitive but nitrate concentrations have been elevated substantially by N deposition (Fig. 6, Curtis and Simpson, 2011). Analysis of sediment cores from Small Water showed a strong decline in the stable isotope 15 N suggestive of anthropogenic N inputs (see above), while the winter phytoplankton was dominated by the mesotrophic diatom Asterionella formosa which is unusual for an oligotrophic upland lake (Curtis and Simpson, 2011). Bioassays of phytoplankton production at this site indicated colimitation by N and P throughout the growing season (Curtis and Simpson, 2011). Excess N availability may compromise the integrity of upland water ecosystems in several ways. First, shifts in the relative abundance of planktonic diatom species towards those favouring mesotrophic waters have been widely recorded in alpine lakes, with some evidence from UK upland waters (see above Fig. 6). Similarly, Battarbee et al. (this volume-b) describe unexpected changes in the (largely benthic) diatom ora of some AWMN lakes recovering from acidication that may be indicative of nutrient enrichment by enhanced nitrate inputs. Nutrient addition experiments have also demonstrated responses in the composition of epilithic periphyton in addition to shifts in productivity (Maberly et al., 2002). There are suggestions of a shift from N-xing epilithic cyanobacteria in low N deposition areas of Sweden to non N-xing species in high deposition areas, although the role of organic carbon and light climate cannot be ruled out (Liess et al., 2009). Some studies in upland lakes, for example the AWMN site Loch Coire Fionnaraich, have found evidence of increased relative abundance of small chrysophytes in P rather than N limited lakes where they have a competitive advantage in P uptake (Pla et al., 2009). It was further speculated that this phenomenon could be linked to the increases in small
planktonic cladocera observed by Kattel et al. (2006) at Loch Coire Fionnaraich. Effects of P limitation (induced by excess N availability) on zooplankton and macroinvertebrate grazers have been suggested. Phosphorus-limited algae are poor quality food for zooplankton, and their development may therefore induce effects throughout aquatic food webs (Elser et al., 2009b). Nitrogen:phosphorus recycling ratios in macroinvertebrates were much greater in high, relative to low, N deposition lakes of Sweden (Liess et al., 2009). Ultimately, increased N loading may reduce the biodiversity of lake phytoplankton because producer diversity tends to be low when one nutrient is present in excess relative to others (Elser et al., 2009b). Evidence for ecological changes in remote lakes has been used to support the application of nutrient N critical loads, primarily in North America (e.g. Baron et al., 2011; Saros et al., 2011; Nanus et al., 2012). These studies suggest very low critical loads for nutrient N to alpine lakes in the US, ranging from 1.4 to 6.0 kg N ha1 year1 . Changes in diatom communities may already have occurred in response to N inputs in lakes where measured late summer nitrate concentrations are <1 equiv. L1 (Arnett et al., 2012; Nanus et al., 2012). In the UK, the application of such low critical loads would imply that all upland waters are exceeded and that alterations to aquatic nutrient cycles and planktonic assemblages must be ubiquitous (Curtis et al., in press-b). However, convincing evidence remains elusive, possibly because of the confounding effects of acidication and climate change. The effects of increased nutrient inputs from N deposition on softwater macrophytes, the predominant type in upland waters in the UK, have been reviewed recently by De Wit and Lindholm (2010). They identied N deposition as a likely cause of the major 20th century decline in isoetid communities that tend to dominate oligotrophic lakes in Western Europe, adding to the negative impacts caused by high aluminium concentrations and low pH et al., 2009). Various potential mechanisms were (e.g. Ctvrtlkov proposed, including a competitive advantage for the macrophyte Juncus bulbosus under conditions of increased ammonium
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concentrations and CO2 availability due to acidication or liming, reduced light availability to isoetids due to smothering by lamentous green algae, or enhanced phytoplankton production and/or increases in oating leaved plants due to eutrophication. Empirical nutrient N critical loads for softwater macrophytes have been suggested to be in the range of 310 kg N ha1 year1 (Bobbink and Hettelingh, 2011) for EUNIS Category C1.1 (permanent oligotrophic) and C1.4 (dystrophic) lakes. The lower end of the range is recommended for alpine lakes (which are not dened) while the upper end is recommended for Atlantic softwater lakes. As with empirical critical loads based on phytoplankton change in alpine lakes, the nutrient N critical loads for oligotrophic lakes (based on softwater macrophytes) are currently exceeded across most of the UK (Curtis and Simpson, 2011). Overall, there is a growing body of evidence that N deposition to UK upland waters is likely to have induced or enhanced P limitation of primary production in both the plankton and the epilithon. Effects may include changes in algal assemblages, rates of nutrient cycling and aquatic food webs. It is also likely that pressures on softwater macrophyte communities may have caused changes in some sites but the level of threat to isoetid communities is largely unknown. 2.1.3. Trace metals and POPs Fossil fuel combustion leads to the emission and long range transport not only of acid gases but also of toxic metals such as Hg and Pb, and persistent organic pollutants (POPs). Toxic trace metals and POPs are also produced by a range of other industrial processes. Such toxic air pollutants have contaminated upland waters throughout most of the world, including remote Arctic (high latitude) and alpine (high altitude) regions where, despite often great distances from sources, volatile contaminants such as POPs may accumulate through condensation due to low temperatures. Heavy metals and POPs have been found at signicant concentrations in Svalbard in the high Arctic (Rognerud et al., 2002), Tibet (Yang et al., 2010) and high mountain lakes around Europe, especially at the AWMN site Lochnagar in Scotland (Yang et al., 2002a,b; Vives et al., 2004a; Rosseland et al., 2007), which is the most highly studied mountain lake in the UK for trace metals and POPs. The few other studies conducted in the UK have also shown evidence of contamination in remote lakes (Rose et al., 2012) reecting the intense industrialisation of the UK over the last 200 years, but the spatial extent of the relatively high levels of contamination as seen at Lochnagar remains largely unknown. A wide range of trace metals have been identied at elevated concentrations in remote lakes. For example, studies of sh gills and kidneys from nine remote European lakes found high gill concentrations of Al, Pb and Cd and the highest kidney concentrations of Pb and Cd at Lochnagar (Rosseland et al., 2007). Mercury is subject to both bioaccumulation and biomagnication, and sh muscle concentrations have been found to exceed the World Health Organisation (WHO) dietary recommendation of 0.3 g g1 in piscivorous Arctic charr in the high Arctic site Arresjen, Svalbard (Rognerud et al., 2002), despite low sediment and water concentrations, which is indicative of biomagnication. Lower muscle concentrations of Hg (still approaching WHO limits for older sh) were found in brown trout at Lochnagar, but the sampled sh were non-piscivorous (as shown by stomach content analysis) and the Hg accumulation at the site was the highest of all non-piscivorous sh studied (Rosseland et al., 2007). In the same studies of European mountain lakes, a range of POPs were also analysed, including polycyclic aromatic hydrocarbons (PAHs), polybrominated diphenyl ethers (PBDEs) and organochlorines such as polychlorinated biphenyls (PCBs). Many of these compounds are endocrine disruptors. All lakes sampled revealed concentrations of PCBs and DDTs in sh liver and muscle which
Fig. 7. Percentage reduction in decadal full basin inventory of Pb and Hg for upland lakes (redrawn from Rose et al., 2012). Note that negative values for eroded sites indicate an increase in inventories.
were comparable with sh from low altitude systems (Rosseland et al., 2007). Similarly, an earlier study by Rose et al. (2001) found levels of the pesticide toxaphene in sediment from Lochnagar to be higher than at any other site receiving only atmospheric sources and comparable to levels found in the U.S. Great Lakes where there are important riverine inputs. The highest levels of PBDEs in the European mountain lakes study were found in sh muscle and liver from Lochnagar, but concentrations in all the high mountain lakes were relatively low compared with sh from lower altitudes (Vives et al., 2004b). The burden of trace metals and POPs in the above studies of sh from remote lakes led to visible toxic effects in histological examinations of liver tissues, including the trout from Lochnagar (Rosseland et al., 2007). Hence there is clear evidence, albeit from a small number of study sites, for contamination of remote lakes with trace metals and POPs at concentrations of ecological relevance and potentially of concern for human consumption of sh caught in these remote lakes. Many of the most toxic pollutants emitted by human activity have been banned or restricted in recent decades with resultant large reductions in emissions, but some are still in current use and there remains a legacy of trace metals and POPs in soils and sediments including in upland areas (e.g. Rose et al., 2001, 2012; Yang et al., 2002a,b; Grimalt et al., 2004a,b). A future threat to upland waters is therefore posed from these legacy pollutants if environmental change leads to their remobilisation. In particular, trace metals and POPs immobilised in upland catchment soils may be remobilised by soil erosion, induced by land-use or climate change impacts (Rose et al., 2004, 2012). There may also be poorly understood changes to the toxicity of some PAHs under changing ultraviolet exposure regimes due to increasing DOC trends (Schindler, 2009). Rose et al. (2012) argue that many of the changes associated with predicted climate change, such as increased winter rainfall, prolonged droughts and more high-intensity rainfall events are likely to lead to increased soil erosion and transport of terrestrially bound pollutants into surface waters. Fig. 7 compares the reduction in full basin inventories for lake sediment Pb and Hg in catchments with thin soils, eroded soils and non-eroded soils. Catchments with thin
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soils have responded most strongly to reductions in emissions of Pb and Hg while catchments with non-eroded soils show a weaker response. Sites with eroded soils show continued increases in lake basin inventories of Pb and Hg despite emissions reductions in recent decades. In a separate study, deposition of methylmercury to the remote catchment of Lochnagar was found to peak during winter snowfall and it is therefore possible that the main deposition route for this specic pollutant will be reduced if predictions of reduced snowfall under future climate change are realised at this site (Rose et al., 2013). Furthermore, the anaerobic methylation process within catchment soils and sediments is itself a climate sensitive process affected by both precipitation and temperature (Grimalt et al., 2010). Hence climate change clearly has a role to play in affecting both the inputs of trace metals to remote sites and more significantly, the remobilisation of the legacy pollutants stored within catchment soils. 2.2. Land-use change in the uplands Although the evidence suggests that post-industrial revolution changes in the ecology of upland waters in the UK are mainly a consequence of air pollution, land-use change and changes in land management practices in the uplands have also been important and may become more important in future as new economic and policy pressures emerge. Historical pressures in the last century have included large scale afforestation with coniferous, often exotic, tree species, subsidised drainage of moorlands and changes in livestock grazing practices and stocking densities. Various future land-use change scenarios for upland regions were considered in a review paper on the future of the uplands by Reed et al. (2009), including the restoration of degraded peatland for carbon offsetting, expansion of agricultural production further upslope into the uplands with climate change, expansion of renewable hydro-power and wind farms and expansion of upland forestry driven by increasing global commodity prices. Alternative scenarios of reduced farming impacts include reduced nancial support for hill farming with resultant reductions in grazing density and re-wilding of upland landscapes. Land-use change scenarios with recognised impacts on aquatic ecosystems are outlined below. 2.2.1. Forestry One of the most marked changes in the use of the British uplands over the last century has been large scale afforestation, replacing semi-natural vegetation with (usually exotic) coniferous species (Reed et al., 2009). The role of forestry in exacerbating surface water acidication in the uplands, primarily through enhancing rates of pollutant interception, is well established (Harriman and Morrison, 1982; Ormerod et al., 1989; Fowler et al., 1989; Jenkins et al., 1990) and is further conrmed by several studies in this Special Issue. Kreiser et al. (1990) and Battarbee et al. (this volume-b) have both argued that the diatom record preserved in lake sediments shows that afforestation in some upland catchments probably exacerbated acidication. In a spatial study of streams across the North York Moors, afforested catchments showed higher concentrations of sulphate, nitrate and labile aluminium compared with adjacent moorland catchment (Evans et al., this volume). A long-term study compared stream subcatchments with varying degrees of forestry around Loch Ard in central Scotland (Malcolm et al., this volume). Results showed that streams draining large areas of mature or second phase forestry had higher concentrations of non-marine sulphate and labile aluminium, plus lower pH and ANC, than streams with small forested areas or moorland catchments, conrming earlier observations by Harriman and Morrison (1982) and Stoner and Gee (1985). While the forest streams remain more acidied, ANC
is recovering more rapidly than the moorland streams, possibly reecting larger reductions in acid deposition. In a comparison of three paired forest/non-forest catchments in the AWMN, there was also an overall tendency for sulphate to decline and ANC to increase more rapidly in the forested sites, although differences in the strength of trends between individual pairs of sites were mostly insignicant (Monteith et al., this volume). Over the last 20 years changes in forestry policy nationally have resulted in a steep decline in new planting on deep peat soils, a reduction in the planting of exotic monocultures and increased emphasis on enhancing the contribution of forestry to ecosystem services (Reed et al., 2009). For example, The Scottish Forestry Strategy sets out a range of targets for economic, social and environmental outcomes to be achieved by expansion of forestry activities (Scottish Government, 2009), including climate change mitigation and sustainable water resource management. It indicates the need for an increase in woodland cover in Scotland from the current 17% to around 25% in the second half of the century, which would involve the creation of some 650,000 ha of new woodland to add to the current resource of 1,334,000 ha, an expansion of the existing forest area in Scotland by almost 50%. A new planting requirement of at least 10,000 ha per year is required simply to maintain carbon sequestration levels (Natural Scotland, 2011). Sustainable forest management initiatives include increasing the proportion of native species, increasing the variety of conifer species, replacing even-aged stands with broader age ranges, and a greater use of broadleaf species (Helliwell et al., this volume-a). Some of these measures may have benecial impacts on aquatic ecosystems, for example through reducing the scavenging of air pollutants and associated acidication where broadleaves replace conifers. However, replacement of conifers with broadleaves could have complex effects on water chemistry, since lower C/N ratios in broadleaf litter could promote nitrate leaching and store less carbon, but reduce DOC production in the litter layer; these processes are not currently included in dynamic models. Dynamic modelling applications using new planting scenarios presented in this Special Issue predict only minor impacts on water quality (Helliwell et al., this volume-b), but small improvements in water quality are predicted for (hypothetical) total forest removal scenarios. The future role of forest in exacerbating soil and surface water acidication is likely to be much smaller in absolute terms than during the period of maximum S and N emissions in the 1970s and 1980s. However, expansion of forestry into previously unafforested areas, especially in regions with a historical record of soil and water acidication and continuing elevated levels of acid deposition, could have more serious impacts on water quality. In addition to impacts on water quality, forestry also affects hydrology and temperature. Holden et al. (2007a,b) suggest that there is likely to be an increase in planting of mixed leaf woodland in the UK uplands and riparian zones to meet various aims including increased water retention and reduced stormwater runoff, improved water quality and carbon sequestration. Riparian tree planting initiatives are already in place in some upland areas to provide shading as a form of mitigation against potential temperature increases which could adversely affect juvenile salmonids under climate change (Malcolm et al., 2008; Hannah et al., 2008). Hence there are evidently some potential benets of new planting for aquatic ecosystems as well as the threats described above. 2.2.2. Moorland management (burning, drain blocking) Another contentious issue in the uplands is whether moorland management practices have a signicant impact on water quality, in particular DOC and particulate organic carbon (POC) uxes. Peatlands are one of the dominant habitat types in the uplands of the UK. In many parts of the UK these have been extensively eroded over recent centuries (Fig. 8, Stevenson et al., 1990) and
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Worrall et al., 2007). Effects of drain blocking on aquatic biota are poorly understood (Ramchunder et al., 2009). A more recent issue has been the practice of some new wind farm construction projects to clear fell forest and use the mulch to block moorland drains, which may pose a threat to surface water quality in terms of nutrients and acidity (I. Malcolm, pers. comm.). Rotational burning of upland sites is a long established practice for creating a mosaic of different aged vegetation stands to improve habitats for grouse as well as sheep and deer (Ramchunder et al., 2009). In the 21st century there has been an increase in the extent of new burns, reecting in part new incentives such as the Moorland Regeneration Programme of 2001 (Ramchunder et al., 2009; Yallop et al., 2010). Some studies indicate that the extent of new moorland burn (<c. 4 years) on blanket peat soils increases colour and DOC concentrations in streams, but not older burns or non blanket peat soils (Yallop and Clutterbuck, 2009). In contrast, a study of a set of headwater sub-catchments of the River Nidd, Yorkshire, Chapman et al. (2010) found no relationship between water colour, or long term changes in water colour, and the spatial extent of burning. However, there are differences in results from laboratory, plot and catchment scale studies and hence the net impacts of changes in burning practice are dependent on the scale of study (Holden et al., 2012). It is also unclear whether impacts on water chemistry are a result of the burning per se or associated changes in vegetation cover (Bain et al., 2011). Burning has also been linked directly to effects on aquatic invertebrate communities through effects of temperature, ash, sediment transport and macronutrient release (Ramchunder et al., 2009).
Fig. 8. Severe gully erosion on Bleaklow, the headwaters of the River Etherow in the Peak District of north-central England. In places the depth of eroded peat is more than 2 m down to bedrock. Photos: C.J. Curtis.
increased degradation over recent decades has resulted in less than 50% remaining in a favourable condition (Bain et al., 2011). Causes of damage have included land drainage, acid deposition, overgrazing and poorly managed burning. Physical damage to peatlands has been proposed as a cause of higher concentrations of DOC in surface waters (Bain et al., 2011) although to date it has been difcult to distinguish between effects of catchment-specic factors and the regional effect of declining acid deposition in these heavily manipulated systems (see Section 3.1). There are moves towards large scale changes to land-use and habitat management for restoring peatlands which could include blocking drainage ditches, changing grazing regimes and modifying burning practices (Reed et al., 2009; Bain et al., 2011). In some areas direct interventions are being implemented, including gully blocking, removal of woodlands and revegetation of eroded peat surfaces. While no longer widely practised on peatland, the subsidised cutting of drainage ditches in the late 20th century resulted in major changes to catchment hydrology in large parts of the uplands, affecting local water tables, ood hydrographs, drying out and erosion of areas downslope of ditches (Ramchunder et al., 2009). Associated water quality impacts include increased suspended sediment loads (Holden et al., 2007a) and DOC uxes (Mitchell, 1990; Wallage et al., 2006). More recently, measures have been taken to reverse the damage and restore peatlands through blocking gullies, grips and drains. Drain-blocking in peatlands has been found in some studies to reduce suspended sediment (Holden et al., 2007b), DOC and colour in surface waters (Armstrong et al., 2010; Wilson et al., 2011), while others found little impact on stream DOC (e.g.
2.2.3. Grazing Several economic drivers of land-use change have impacted the uplands over the last few centuries, ranging from a switch from summer moorland grazing to hardier breeds of sheep able to overwinter during the 18th century, to European Common Agricultural Policy subsidies that led to a 30% increase in sheep numbers on UK moorlands from the 1970s to the 1990s (Holden et al., 2007a) and more recent post-subsidy declines in stocking. The intensity of grazing is an important factor in determining its potential impact on soils and waters. Light grazing may benet terrestrial biodiversity but overgrazing and trampling may cause vegetation changes and ultimately soil erosion and gullying in peatlands (Bain et al., 2011). In the uplands, sheep and deer grazing are most widespread although cattle grazing may also be practised, as observed at several sites in the UK Acid Waters Monitoring Network (e.g. Burnmoor Tarn, Scoat Tarn, Llyn Llagi, Round Loch of Glenhead) and at the Loch Laidon grazing experiment on Rannoch Moor (Shilland et al., 2011). For aquatic ecosystems the main impacts of grazing practices relate to possible soil erosion and transport of particulates into surface waters. Trampling has also been shown to affect catchment hydrology by compacting soils and increasing inltration-excess overland ow (Holden et al., 2007a,b). Another role played by grazers is a removal mechanism for nutrients from catchments; for example, net biomass accrued within a catchment by grazers is effectively a sink for deposited N if the animals are ultimately removed from the catchment. Some mass balance models account for grazing removal of N in this way. There have in recent decades been calls for a reduction in sheep numbers in the uplands to reduce impacts on upland soils and ecosystems (Holden et al., 2007a,b) so this particular threat to aquatic ecosystems may decline in importance. However, in the Scottish uplands deer grazing pressure does not seem to be declining, although there are local exceptions such as the Mar Lodge Estate where managed reductions have taken place (Mar Lodge Independent Review Panel, 2011).
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Table 1 Regional range of projected changes from long term mean baseline (19611990) in UK air temperature and precipitation under medium emissions scenarios for the 2080s (UKCP09) Source: http://ukclimateprojections.defra.gov.uk/21730. Accessed 25th October 2013. Probability 10% 50% 90% Mean winter temperature ( C) 0.91.6 2.23.0 3.64.7 Mean summer temperature ( C) 1.52.1 3.03.9 4.96.5 Mean winter precipitation (%) +1 to +6 +11 to +23 +24 to +54 Mean summer precipitation (%) 56 to 29 28 to 12 0 to +8
Sheep are able to crop vegetation at close to ground level, whereas cattle remove it less selectively using a ripping action that often leaves a longer, more heterogeneous sward (Armstrong, 1998) and any management changes resulting in a change in the dominant grazing animal, or especially in animal density, may have consequences for soil erosion (Albon et al., 2007). The effects of a switch from sheep to cattle grazing on upland water quality have been tested experimentally for a number of years at the Loch Laidon grazing experiment (Shilland et al., 2011). While this has resulted in substantial shifts in the composition of terrestrial vegetation, no signicant differences in water quality have been detected over the rst two decades of the experiment between cattle-grazed and control catchments (Shilland et al., 2011). Changes in diatom assemblages during the course of this experiment were similar at both cattle-grazed and control catchments, both indicating shifts towards less acid tolerant taxa suggesting that the reduction in acid deposition that has taken place has been the most important inuence on water quality. This nding supports the conclusions from a number of palaeoecological studies (e.g. Battarbee et al., 1985) that demonstrated evidence for catchment vegetation change related to land-use change but no evidence for impacts on water quality.
2.3. Climate change impacts on upland waters Climate change affects the hydrological, physical, chemical and biological characteristics of all freshwaters and is thereby a key inuence on biogeochemical cycling (nutrients, major ions, DOC, organic pollutants, metals), food webs and biodiversity (Curtis and Simpson, 2007). While pressures on upland ecosystems from sulphur and N deposition are declining, climate change is expected to have an increasing impact on upland biogeochemical cycling, the behaviour of pollutants stored in catchment soils, land use and land management, and hence water quality over the coming decades. These interactions are illustrated conceptually in Fig. 2. The potential impacts of climate change on upland waters have been investigated in a number of international research projects including EMERGE (Catalan et al., 2009; Kernan et al., 2009) and Euro-limpacs (e.g. Whitehead et al., 2009). Predictions of the direct effects of climate change on aquatic ecosystems are very complex, and once combined with other human impacts pose an even greater challenge. The latest UK climate projections, UKCP09 (Murphy et al. 2009), provide ranges of projected winter temperature increases by the 2080s for the UK, which are summarised for regionally aggregated data in Table 1. The percentiles refer to the probability of different increases: the 50th percentile is the central estimate of change, while the 10th percentile is very likely to be exceeded and the 90th percentile is very likely not to be exceeded (Murphy et al., 2009). It should be noted that UK climate has been relatively stable over the period the AWMN has been operating to date, and while data from the network are already being used to examine the effects of interannual climate variability on biogeochemistry and biota, further years of monitoring will be necessary before effects of climate change can begin to be assessed. Furthermore, there are very few long-term data series for water temperature in the UK. UKCP09 projects UK mean winter air temperature increases in all parts of the UK with 50th percentile projections of 2.23.0 C
increases and a 1090th percentile range of 0.94.7 C with the medium emissions scenario. The lowest regional increases are projected for northern and eastern Scotland and the highest for southern and eastern England. For mean summer air temperatures the 50th percentile projections are increases in the range 3.03.9 C (1090th percentile 1.56.5 C) with the smallest increases in northern Scotland and the greatest in southern and eastern England. Hence the greatest air temperature increases may be in the lowland eastern and southern regions of the UK while most MMH catchments are in regions with lower projected increases. Winter precipitation may increase in all regions, with a 50th percentile range of 1123% increase and a 1090th percentile range of 154% increases. The smallest increases are projected for eastern Scotland and Northern Ireland while the largest are projected for south-west England. Overall, decreases in summer precipitation are more likely, with a 50th percentile range of 1228% reductions. The 10th percentile of model runs project decreases in the range 2956% while the 90th percentile of runs project small increases in the range 08%. The largest changes are modelled for southern England and the smallest for northern Scotland. Here we consider two aspects of these climate change impacts on upland waters; direct changes in terms of temperature and hydrology, and biogeochemical cycles in upland catchments, specically the carbon cycle.
2.3.1. Direct impacts of climate change in the uplands: hydrology, temperature and aquatic ecology Extensive analysis of weather station data over several decades indicates that upland areas are experiencing greater climatic change than lowland areas in the UK (Burt and Holden, 2010; Holden and Rose, 2011), with stronger increases in winter minimum temperatures and increasing winter precipitation. Increasing temperature may impact upland lake ecosystems in several ways, through decreases in winter ice cover, changes to the thermal structure of lakes and their mixing behaviour, and physical changes brought about by temperature increase, which can have signicant impacts on surface water chemistry and biology. There may also be a feedback in terms of increasing DOC concentrations (see Section 3.1) which could affect absorbance of solar radiation and hence lead to earlier and more frequent onset of ice melting. Mountain lakes are very sensitive indicators of environmental change and have responded to climate change both chemically, through pH changes in response to weathering inputs from catchments (Psenner and Schmidt, 1992; Sommaruga-Wgrath et al., 1997; Koinig et al., 1998; Mast et al., 2011) and biologically, as indicated by palaeolimnological records of changes in lake phytoplankton (e.g. Battarbee et al., 2002; Catalan et al., 2002; Smol et al., 2005; Curtis et al., 2009). Similar changes have also been recorded in Arctic lakes (e.g. Prowse et al., 2006; Hobbs et al., 2010; Holmgren et al., 2010). The most alpine site in the UK AWMN, Lochnagar, lies within a maritime altitude zone deemed to be particularly sensitive to climatic inuences on ice cover duration (Thompson et al., 2007; Kettle and Thompson, 2007). While currently freezing for about 3 months of the year, it was likely to have experienced around 6 weeks extra ice cover during the Little Ice Age, while just modest warming from the present could mean very limited opportunities
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Fig. 9. Modelled effects of climate warming on lake ice-cover duration in Scotland. Source: Kettle and Thompson (2007); with kind permission from Springer Science and Business Media.
for ice formation in future (Fig. 9, Thompson et al., 2007). Previous climate change projections (UKCIP02) suggested an increase in summer maximum temperatures by 24 C by 2080 while monthly minima in winter would rarely be below 0 C (Kettle and Thompson, 2007). Even low emission scenarios predicted drastic reductions in snowfall at Lochnagar due to temperature increases, although the overall change in precipitation (as rainfall) is unknown. The growing season for primary producers within the loch was also predicted to increase, perhaps by as much as 50% under high emission scenarios (Kettle and Thompson, 2007). More recent climate projections under UKCP09 suggest even greater warming than UKCIP02 for both winter and summer (Murphy et al., 2009). The UK uplands experience high windspeeds and upland lakes tend thereby to be well mixed. Temporary stratication can occur in summer during calm periods (e.g. Curtis and Simpson, 2011; Shilland et al., 2012), so most lakes can be classied as being polymictic. Only one site in the AWMN, Loch Chon, straties consistently each year (Shilland et al., 2012). The extent to which the mixing characteristics of UK lakes will change will depend on future changes in summer air temperature and on wind strength, and the altitude and exposure of individual sites. Overall we might expect a shift to more stable systems with stratication taking place more frequently and for longer periods of time as summer wind speeds decline slightly while temperatures increase at sites like Lochnagar (Kettle and Thompson, 2007). In running waters there are concerns for the future for cold stenothermic taxa, most notably salmonid sh such as salmon (Salmo salar) and brown trout (Salmo trutta), in terms of potential changes in geographic range in response to changing water temperatures and ows (e.g. Jonssen and Jonssen, 2009; Elliott and Elliott, 2010; Moore et al., 2012). Climate warming resulted in an upward shift of thermal habitat for brown trout in Swiss alpine rivers and streams, indicating a loss of habitat due to physical barriers to upstream migration and linked to an increase in temperature dependent kidney disease (Hari et al., 2006). Predictions of lake and stream warming in northern North America indicate temperatures will increase beyond lethal levels for cold stenothermic sh taxa like lake trout and whitesh (Schindler, 2009; Cunjak
et al., 2013). In Scotland, Hrachowitz et al. (2010) demonstrated that small upland streams with no forest cover were most vulnerable to the adverse effects of projected climate change on water temperature regime suitability for salmonids. Clews et al. (2010) found signicant relationships between juvenile salmonid population densities and antecedent summer climate in the River Wye in Wales over 20 years, with hot, dry summers associated with reductions in salmon density. Aquatic macroinvertebrates are also sensitive to the effects of climate change. Local extinctions of sensitive macroinvertebrate species under 3 C warming at Llyn Brianne in Wales were predicted by Durance and Ormerod (2007), while temperature was also found to be the most important determinant of macroinvertebrate assemblages in northern streams across a gradient from Denmark to Greenland and northern Sweden under the North Watch study (Friburg et al., 2013). A key unknown is the adaptive capacity of the impacted macroinvertebrate species, but with increasing temperatures, metabolic rates will increase with likely impacts on the functioning of whole ecosystems (Tetzlaff et al., 2013a). As well as changes in water temperature, changes in hydrological regimes are also anticipated under climate change, with poorly understood impacts on the structure and function of upland water ecosystems. Hydrological modelling at one of the AWMN sites, Dargall Lane, indicates that most UKCP09 projections would result in major changes in ow seasonality with increased winter discharges and reduced summer ows (Thompson, 2012). Decreases in snowfall have been projected for Scottish upland streams including the AWMN site Allt aMharchaidh, with associated reductions in snowmelt inputs but increased winter ows (Capell et al., 2013). Such hydrological changes are likely to result in important hydrochemical and biological changes in aquatic ecosystems; indeed, Tetzlaff et al. (2013b) describe northern rivers studied under the North Watch project, which included the AWMN sites Allt aMharcaidh and Allt na Coire nan Con, as being on the cusp of major changes in ecohydrology. In the North Watch study, temperature seems to be a more important determinant of change for macroinvertebrate communities (Friburg et al., 2013) while geographic range suitability for salmon may be impacted by changes in both hydrology and temperature (Tetzlaff et al., 2005; Clews et al., 2010; Moore et al., 2012; Cunjak et al., 2013). Lower ows and higher temperatures during summer, as predicted for upland catchments, are likely to increase stress on salmonids, while large oods may wash out salmonid eggs deposited in spawning gravels (Clews et al., 2010; Thompson, 2012; Cunjak et al., 2013). Finally, catchment vegetation is also expected to respond to changes in climate, with resultant inuences on water quality and ecohydrology which are poorly understood (Tetzlaff et al., 2013b). The composition and distribution of catchment vegetation may be affected by changing temperature and precipitation regimes, with physiological responses that may include, for example, altered rooting patterns and phenology (Tetzlaff et al., 2013b). There are likely to be complex feedbacks between catchment hydrology and vegetation change (for example through stomatal control of water losses and changed evapotranspiration) which are difcult to predict, as well as impacts on biogeochemical cycling of carbon and N. 2.3.2. Climate change impacts on catchment biogeochemical cycles The macronutrient cycles of upland catchments are intimately linked to both climate and vegetation type. Since vegetation cover and type will also respond directly to changes in climate there are potential feedbacks in terms of nutrient cycles, all of which will impact on upland aquatic ecosystems.
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Considerable scientic attention is currently being devoted to the quantication of the major carbon uxes in peatlands (many of which are located in the uplands in the UK) to determine their future role as potential sinks or sources of carbon under climate change. The key question is whether peatlands will continue to sequester carbon and build peat (a net sink) or will degrade and release old carbon (net source). Climate change impacts on upland peatlands may depend on whether they are intact and healthy or degraded, since damaged peatlands are considered to be less resilient to climate change (Bain et al., 2011). Climate change may lead to increased erosion, ooding and risk of wildres (Bain et al., 2011) which could increase carbon loss both to the atmosphere and to surface waters. Increased peat erosion, transport of particulate organic carbon and sedimentation impacts on downstream sheries, e.g. for salmon, could result (House et al., 2010; Bain et al., 2011). Carbon losses from peatlands may be increased both directly by the inuence of higher temperatures on organic matter decomposition, and by droughts, whereby increased oxygen penetration results in greater phenol oxidase activity and reduced concentrations of the phenolic compounds which retard decomposition (Fenner and Freeman, 2011). Davidson and Janssens (2006) argued that the decomposition rate of more refractory organic matter such as peat is particularly sensitive to increasing temperature as a result of the relatively high activation energies of such substrates, a hypothesis supported by incubation experiments of Craine et al. (2010) on soils across North America. A modelling study of c. 1000 Norwegian headwater lakes suggested that increased terrestrial vegetation cover under climate change could lead to major increases in organic carbon in lakes (Larsen et al., 2011). Such an effect could be additive to the deacidication mediated increases in DOC described above (e.g. Evans et al., 2012). Analysis of bioclimatic envelope (niche) models suggests that more than half of UK peatlands will experience change by 2050 (Clark et al., 2010) while over the longer term, the area of actively growing blanket bogs could shrink by 84% from the present day (Gallego-Sala et al., 2010). However, other dynamic process-based model applications gave mixed results with respect to the net storage or release of carbon, showing the critical importance of monitoring the response of peatlands to climate change for reducing uncertainties in modelling and future predictions (House et al., 2010). The carbon cycle is not the only aspect of upland catchment biogeochemistry to be impacted by climate change. Whitehead et al. (2009) reviewed climate variables that can affect the acidication process, including higher temperatures, summer drought, wetter winters, reduced snow cover and changes in hydrological pathways. Climate change is likely to lead to changes in the incidence of storm events that are known to deposit large sea-salt loads onto upland catchments, leading to temporary displacement of acid cations stored in catchment soils and episodic acidication (Skjelkvle et al., 2007; Evans et al., 2008b; Monteith et al., this volume). Drought and rewetting cycles can be important mechanisms for the oxidation of reduced sulphur compounds in organic soils and release of acidity (Clark et al., 2005, 2009; Schindler, 2009) and in upland waters in parts of the USA, climate induced moderation of sulphate release from catchments soils has overtaken S deposition as the main determinant of streamwater sulphate concentrations (Mitchell and Likens, 2011). Such processes are likely to be important in areas of the UK uplands such as the North York Moors where slow release of stored S occurs (Evans et al., this volume). Climatic controls on the leaching of nitrate from upland soils have been demonstrated through synchronous patterns in standardised nitrate concentrations and the North Atlantic Oscillation
(NAO) Index (Monteith et al., 2000, this volume). This observation is consistent with independent evidence from stable isotopes that a major proportion of the nitrate in upland waters has been produced microbially (Curtis et al., 2012), since climate is an important determinant of mineralisation and nitrication processes. Hence it is clear that climate induced changes to biogeochemical cycles in the uplands will inuence surface water uxes of carbon (both particulate and dissolved), N and potentially sulphur as well. The water quality implications of all these changes have been discussed previously in the context of land-use change and recovery from acidication above. 3. Synthesis and recommendations The diverse drivers of change affecting upland water ecosystems result in complex, interacting responses in terms of upland catchment biogeochemistry, aquatic chemistry and aquatic ecology (see Fig. 2) and future directions of change are therefore difcult to predict. Outcomes will vary from site to site depending on the continuing, although declining, inuence of acid deposition and the emergence of new pressures, especially from climate change. 3.1. The legacy of acid deposition Although acid deposition has declined substantially its legacy will continue to inuence surface waters in the UK for many decades to come, as soils slowly release S and N compounds and toxic trace metals through leaching and erosion and begin to regain buffering capacity through the process of geological weathering. While partial chemical recovery of acidied watersheds is already clear, pH and labile aluminium concentrations in more impacted waters are unlikely to reach conditions required to meet good ecological status for several decades. Where soils become saturated by N as predicted by FAB modelling, recovery may be even further delayed. Chemical and biological recovery processes will both be inuenced by changes in temperature and hydrology brought about by climate change and changes in upland management. Any increase in winter precipitation, or increase in sea-salt deposition brought about by increased storminess, is likely to depress pH and raise labile aluminium concentrations in recovering waters by favouring more lateral ow paths and stimulating displacement of acid cations from soils by marine base cations. Meanwhile, any changes in the extent and species composition of upland forests will continue to inuence atmospheric acid interception rates, while changes in terrestrial N demand brought about by felling and replanting will result in pulsed releases of acidifying nitrate. Of continuing concern is the extent to which persistence of occasional highly acid episodes brought about by these interactions may serve as a ceiling for biological recovery (Lepori et al., 2003; Kowalik et al., 2007). Data from AWMN stream sites suggest that the severity of acid episodes is declining at least as rapidly as mean acidity (Monteith et al., this volume), but the magnitude of occasional spikes in labile aluminium and hydrogen ion concentrations may still act as a barrier for the return of some acid-sensitive species. A further consequence of acid deposition on upland waters has been changes in DOC concentration, although the interpretation of DOC trends has not been without controversy. Data from the UK Acid Waters Monitoring Network provided some of the rst evidence for recent large regional-scale increases in upland water DOC uxes (Freeman et al., 2001; Evans et al., 2006). Subsequent debate over drivers of increasing DOC, whether reductions in acid deposition, climate change (see Section 2.3.2), N deposition or land-use change (see Section 2.2.2), has prompted various
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national-scale investigations (e.g. Evans et al., 2006; De Wit et al., 2007; Erlandsson et al., 2010; SanClements et al., 2012), a large international study of DOC trends involving sites across the ICP Waters network (Monteith et al., 2007) and eld experiments (Evans et al., 2012). Together these provide highly persuasive evidence that soil acidity, and hence acid deposition, exerts a major temporal control on the mobility of DOC and that reduced acid deposition is the primary driver of increasing DOC trends seen in streams and lakes draining the uplands of the UK and other acidication-impacted regions. This conclusion has important implications for our understanding of the reference conditions of acidied lakes in terms of pre-industrial DOC concentrations and pH (Evans et al., 2008a; Erlandsson et al., 2011), as it suggests that organic acidity may have partially countered changes in mineral acidity from atmospheric pollutants. Consequently, the pre-industrial pH of surface waters in some areas may not have been as high, and recovery in pH may therefore not be as marked, as dynamic models such as MAGIC once predicted (Battarbee et al., 2005). The link established between rising DOC and falling acid deposition is highly pertinent to the interests of water supply industries serving most of the north and west of the UK. Rising levels of DOC are increasing water treatment costs substantially, as removal of DOC is necessary prior to chlorination. Deposition-driven increases in the solubility of soil organic matter are likely to make DOC export more vulnerable to the range of climatic factors and soil degradation described in earlier sections. Levels of DOC reaching the water supply system in these regions are therefore likely to at least be sustained, or even further increased, in future as soils continue to recover from acidication, and attempts to reduce uxes through local catchment management will need to take into account this dominant regional effect. Another factor linked to air pollution with the potential to inuence DOC levels in upland waters is the eutrophying effect of N deposition on terrestrial ecosystems. Pregitzer et al. (2004) demonstrated that nitrate additions of 30 kg N ha1 year1 to hardwood forests, comparable to high N deposition regions of the UK uplands, increased soilwater leaching uxes of nitrate, DOC and also DON. Findlay (2005) suggested a similar mechanism may be responsible for increased DOC uxes into the Hudson River, possibly due to changes in soil enzyme activity. Even if terrestrial primary productivity across the UK uplands is effectively N-saturated across the region today, the legacy of historical N deposition is likely to be affecting nutrient cycling. It is therefore possible that DOC concentrations and uxes may be approaching, or already have exceeded, pre-industrial levels as a result of the combined effects of N-enhanced NPP and an acid deposition-induced return towards pre-industrial organic matter solubility. Regardless of the driver, in upland waters where levels of planktonic algae and suspended sediment are often relatively low, DOC concentration is normally the dominant determinant of the degree of penetration of photosynthetically active radiation (PAR) through the water column. Rising DOC concentrations are therefore reducing PAR penetration of upland lakes, where in some cases benthic primary production may be limited by light rather than nutrients (Karlsson et al., 2009). Dissolved organic matter also has an important function as a sunscreen, limiting the exposure of biota to harmful ultraviolet radiation (Schindler, 2009). Furthermore, changes in water colour may affect lake freezing dynamics. The ecosystem services provided by upland waters and their catchments in terms of carbon storage, aquatic biodiversity and the provision of high quality drinking water are evidently affected by multiple interacting factors. Climate change and land-use clearly play important roles in the mobilisation of carbon from catchment
soils and as acid deposition continues to decline there is likely to be a shift in the relative importance of the key drivers of change in DOC. 3.2. Emerging pressures on upland waters It is clear that the main pressures on upland waters are shifting in relative importance and presenting new challenges to scientists, policymakers, landscape and environmental managers. As atmospheric deposition of pollutants continues to decrease, climate change is set to continue, with major changes for upland waters anticipated in temperature, seasonal ow patterns and the role of snowfall, snowmelt and lake ice-cover. Concomitantly, land use and management interventions are also expected to change. While forestry practices have moved away from large scale use of conifers in acid-sensitive areas, there is nevertheless a clear intention in some regions, as in Scotland, to signicantly increase the planted area and this may lead to conicting interests with other stakeholders in the uplands such as groups working to restore salmonid sheries in previously acidied areas. On the other hand, mitigation efforts to reduce the impacts of increasing temperatures on upland stream ecosystems may include riparian tree planting. Even as acid deposition is decreasing across Europe and North America, there is increasing evidence for nutrient-N induced changes in phytoplankton assemblages of Arctic and alpine lakes at deposition levels lower than those currently experienced anywhere in the UK. Likewise, changes in aquatic macrophyte communities in upland waters have been attributed to N deposition in various European countries, but not yet in the UK. In both cases, convincing evidence for such changes in the UK uplands is still lacking. In part this may be due to the availability of historical data, the main impacts of N deposition possibly having occurred several decades prior to the establishment of robust monitoring systems, but palaeolimnological studies of upland lakes in high N deposition areas of the UK have yet to provide conclusive evidence of signicant enrichment during this period (Curtis and Simpson, 2011). Potential changes due to changing nutrient regimes may also have been masked, or inhibited, by the confounding effects of acidication (Fig. 1b). Deposition of both N and S, as well as trace metals and toxic organic compounds, is however still declining and future issues are likely to revolve around the legacy of cumulative deposition loads over many decades in the context of a changing climate in the uplands. The possibility that climate change could lead to increasing mobilisation of such pollutants stored in upland catchment soils is a very real one. Changes in perceived pressures may sometimes result in seemingly conicting benets and disbenets. Hence the reduction in acid deposition that is promoting the return of acid-sensitive aquatic species would also appear to be resulting in increased water treatment costs due to increasing trends in DOC, and potentially, depending on the fate of the uvial carbon, a net loss of soil carbon to the atmosphere. On the other hand, there are also potential co-benets which policymakers should aim to maximise. For example, reduced exploitation and physical restoration of peatlands can improve water quality and reduce UK carbon emissions. There are complex interactions between changes in atmospheric deposition, land use, climate change and catchment hydrology with poorly understood feedbacks, as discussed in Section 3.1 above in the context of DOC trends. Nevertheless, a wide range of studies indicate ongoing, but changing, impacts on sensitive biota in aquatic ecosystems, from phytoplankton and aquatic macrophytes to macroinvertebrates and salmonid sh, leading to changes in food webs and in ecosystem functioning.
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Given the wide range of often interacting and evolving pressures that upland water ecosystems are facing, an integrated ecosystem approach will be necessary to balance the needs of conservation and the wider stakeholder community. Such an integrated approach, along with the establishment of resilient ecological networks, is indeed recommended in the UK Government White Paper on the Environment (H.M. Government, 2011). A further target in the White Paper is to identify where land can be managed to deliver multiple benets, including improved water quality, ood alleviation and biodiversity. Integrated monitoring of upland aquatic ecosystems is a prerequisite for understanding and implementing these approaches. 3.3. Shifting baselines and novel recovery endpoints While the pressure from deposition of pollutants may be decreasing, climate change over the foreseeable future will have a strong inuence on upland freshwater ecosystems limiting the extent to which restoration targets based on the reference condition concept can be attained. In the absence of such a benchmark it is difcult to dene an ecological endpoint that managers can use for guidance. The recent Government White Paper (H.M. Government, 2011, section 2.69) states that the aim for the majority of UK water bodies is to achieve good ecological status as soon as possible and for as many as possible to attain good ecological status by 2027. This target is set under a section titled Restoring nature in our rivers and water bodies. However, restoring nature is a troublesome concept. Battarbee et al. (this volume-b) and Malcolm et al. (this volume) raise in particular the issue of difculties associated with dening chemical and especially biological endpoints for upland waters recovering from acidication. Critical loads of acidity are set relative to a target ANC value which makes no direct reference to baseline values, so achievement of critical loads, as envisaged under the UN ECE Convention on Long-Range Transboundary Air Pollution, does not imply any return to baseline or reference conditions (Kernan et al., 2010). Dynamic models like MAGIC can be used to hindcast a notional pre-industrial, unimpacted (with respect to acid deposition) chemical baseline (Helliwell et al., this volume-b) and attempts have been made to link these to reference biological assemblages using modelling approaches (e.g. Juggins, 2001). Palaeolimnological records provide a more direct (if only partial) indication of pre-acidication communities. Indications from sediment trap data at AWMN sites suggest that trajectories of recovery in diatom communities are not necessarily returning towards these baseline conditions but potentially towards entirely novel assemblages (Battarbee et al., this volume-b). Dynamic modelling of recovery from acidication indicates that baseline conditions for soil base saturation (and hence resilience to acid episodes) are unlikely to be achieved by 2100 (Helliwell et al., this volume-b). Furthermore, increasing DOC trends due to recovery from acidication in upland soils suggests that many upland waters are returning towards a more highly coloured natural baseline, implying a shift in the relative roles of benthic and planktonic primary production (Monteith et al., this volume). Therefore the key question for setting recovery targets is whether these chemical or biological baselines are achievable or even appropriate in the context of a changing climate (Fig. 10). 3.4. The requirements for integrated monitoring of upland waters Many authors have highlighted the dearth of high quality, longterm monitoring sites which can provide the integrated chemical and biological datasets required to investigate the ongoing and future changes in upland water (or indeed any other) ecosystems. For example, Tetzlaff et al. (2005) point out that for studies linking
Fig. 10. Schematic diagram of potential recovery outcomes in the context of declining acidication pressures and other dynamic drivers of change indicating how palaeolimnological (dashed line) and monitoring data (solid line) can be combined to track change from pre-acidication reference conditions to the present day. Potential future change is shown by dotted lines indicating the uncertainty of moving back to the reference (a) or towards a novel endpoint (c). Modied from Battarbee et al. (2012).
hydrological events to biological responses, in this case for juvenile feeding and adult spawning of salmon, high resolution ow data are required and daily mean data may be insufcient. Ramchunder et al. (2009) identify a major lack of research in the investigation of upland stream ecological changes in response to drainage, drain-blocking and burning. The urgent need to continue with existing long-term monitoring programmes in the uplands, such as the Countryside Survey, Environmental Change Network and Acid Waters Monitoring Network, has been highlighted by Dise (2009), Bain et al. (2011) and Tetzlaff et al. (2013a). Upland waters monitoring addresses a series of very specic research needs, highlighted above, in small headwater streams and lakes which are largely neglected under the EU Water Framework Directive and by national regulators due to a historical focus on point source pollution. Nevertheless, these small upland waterbodies full a range of ecosystem services which are very disproportionate to their size. From the range of scientic ndings drawn from the AWMN and other long-term upland monitoring studies reported in this special issue (and see Battarbee et al., this volume-a, this volume-b) there can be little argument that their high quality long-term integrated datasets have made, and continue to make, a substantial contribution to UK environmental scientic capability. The ability to compare and contrast ecological and hydrochemical datasets across years and between sites delivers information and raises new environmental change hypotheses that can then be tested experimentally. The accumulated data have clearly allowed the AWMN to go beyond its original tight remit to provide precise and accurate data on upland environments more widely. Given the long-term time series and high quality of data provided by the AWMN and other important datasets presented in this special issue, these networks will be well placed to contribute to our understanding of emerging issues relating to the use of and damage to upland ecosystems. The AWMN in particular is used here as an example of the type of integrated monitoring that is required to address the research questions posed above and integrated management approaches proposed in the Government White Paper (H.M. Government, 2011). Clearly, long-term monitoring networks need to be continually modied to adapt to changing circumstances. The AWMN, for example, was designed to address questions relating to the chemical and ecological impacts of acid deposition on acidsensitive upland aquatic environments, incorporating both lakes
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and streams, gradients of deposition (including control sites) and paired moorland/afforested catchments (see Battarbee et al., this volume-a). However, the environmental threats to upland waters in the 21st century extend way beyond the problems faced by acid-sensitive systems from acid deposition. In order to develop the evidence base required to understand and track these evolving pressures the remit of the AWMN has been expanded to track changes in upland waters related to pressures from climate and land-use change as well as from air quality. Consequently it has been renamed the UK Upland Waters Monitoring Network (UK UWMN) to reect its broadening role in supporting UK environmental monitoring national capability. New higher alkalinity sites are being added, temperature loggers have been installed and lake and stream stage instrumentation is gradually being added. There are also plans to track changes in nutrients, toxic compounds and land-use enabling observed changes in water quality and biodiversity to be related to the full range of external pressures faced by upland waters. The new network remains closely afliated with the UK Environmental Change Network (ECN), contributing the bulk of its upland water monitoring capacity, while also providing valuable information for national conservation and regulatory bodies on the status of the otherwise understudied upland waters. In addition, given the fundamental links between upland waters and their catchment soils and vegetation, integrated monitoring needs also to combine aquatic ecosystem monitoring with terrestrial biogeochemical monitoring. Closer linkages between the UWMN and the upland Environmental Change Network site, Moor House for example, could provide such an opportunity. The large numbers of studies described above have made use not only of data from the AWMN, but also from other long-term studies and monitoring programmes in the UK. Much more could be done at the UK level to attempt to co-ordinate and link existing programmes to address the key concerns for upland waters. If the UK is committed to leading the way in applying the ecosystem services concept via integrated ecosystem approaches to establish resilient ecological networks, then high quality integrated monitoring is an essential prerequisite. Acknowledgements We are grateful to the various funding bodies that have supported the AWMN over more than 20 years (see Battarbee et al., this volume-a for details of funders) and to all those who have participated in sample collection and analysis. We thank our many colleagues and contributing authors from papers elsewhere in this volume for comments on this manuscript, especially Neil Rose and Iain Malcolm, and Glenn Watts for guidance on climate change scenarios. We also acknowledge with gratitude the improvements made by two reviewers. References
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