iForest - Biogeosciences and Forestry


Changes in vegetation diversity and composition following livestock removal along an upland elevational gradient

G Matt Davies (1)   , Julien Bodart (2)

iForest - Biogeosciences and Forestry, Volume 8, Issue 5, Pages 582-589 (2015)
doi: https://doi.org/10.3832/ifor1557-008
Published: Apr 22, 2015 - Copyright © 2015 SISEF

Research Articles

European heathland habitats are cultural landscapes derived from previously-forested ecosystems. Heathlands are of significant conservation interest but have experienced prolonged degradation due to a range of factors including overgrazing by domestic livestock. There is growing recognition of the need to restore upland landscapes to produce a diverse mosaic of woodlands, heathlands and forest edge ecotones. In some studies stock removal has been sufficient to promote heathland recovery, but often more intensive interventions are required. Few studies have specifically examined how abiotic gradients associated with changing elevation might relate to restoration success. We examined differences in vegetation between grazed and restored areas over a 500 m elevational gradient split across two hillsides that were part of a landscape-scale restoration project in the Scottish Southern Uplands. Species alpha and gamma diversity showed non-linear responses to elevation but the effects of grazing differed between sites. Grazing increased diversity on the lower elevation site but reduced it at higher elevations. The differing effects of grazing with elevation can be interpreted in the context of levels of competition and likely impacts on rates of colonization and extinction. Differences in community composition were assessed using PERMANOVA, NMDS and Cluster Analysis and were primarily controlled by elevation with no significant effect of grazing. The keystone heathland species Calluna vulgaris was not recorded in any of our monitoring plots but some other dwarf shrubs were common. Changes in community structure following stock removal are slow on upland sites but initial impacts interact strongly with abiotic site conditions and pre-restoration vegetation composition. During large-scale restoration it is therefore vital to consider how widely-applied treatments might differ in their effects across landscapes. Changes in diversity may provide a useful early indicator of important ecological processes and likely directions of change.


Calluna vulgaris, Community Change, Grazing, Heathland, Moorland, Restoration, Scotland

Authors’ address

G Matt Davies
School of Environment and Natural Resources, The Ohio State University, Kottman Hall, 2021 Coffey Road, 43210 Columbus, OH (USA)
Julien Bodart
Solway Centre for Environment and Culture, University of Glasgow, Rutherford/McCowan Building, Crichton University Campus, Dumfries DG1 4ZL (Scotland)

Corresponding author

G Matt Davies


Davies GM, Bodart J (2015). Changes in vegetation diversity and composition following livestock removal along an upland elevational gradient. iForest 8: 582-589. - doi: 10.3832/ifor1557-008

Academic Editor

Arthur Gessler

Paper history

Received: Jan 12, 2015
Accepted: Mar 18, 2015

First online: Apr 22, 2015
Publication Date: Oct 01, 2015
Publication Time: 1.17 months

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List of the papers citing this article based on CrossRef Cited-by.

Adair S (2009)
Corehead National Vegetation Classification Survey with comments on management proposals. Full Report, Borders Forest Trust, Jedburgh, Scotland, pp. 38.
Alonso I, Hartley SE, Thurlow M (2009)
Competition between heather and grasses on Scottish moorlands: Interacting effects of nutrient enrichment and grazing regime. Journal of Vegetation Science 12: 249-260.
CrossRef | Gscholar
Anderson P, Radford E (1994)
Changes in vegetation following reduction in grazing pressure on the National Trust’s Kinder Estate, Peak District, Derbyshire, England. Biological Conservation 69: 55-63.
CrossRef | Gscholar
Anderson MJ (2001)
A new method for non-parametric multivariate analysis of variance. Austral Ecology 26: 32-46.
CrossRef | Gscholar
Anderson MJ, Ellingsen KE, McArdle BH (2006)
Multivariate dispersion as a measure of beta diversity. Ecology Letters 9: 683-693.
CrossRef | Gscholar
Ascoli D, Lonati M, Marzano R, Bovio G, Cavallero A, Lombardi G (2013)
Prescribed burning and browsing to control tree encroachment in southern European heathlands. Forest Ecology and Management 289: 69-77.
CrossRef | Gscholar
Bakker J, Berendse F (1999)
Constraints in the restoration of ecological diversity in grassland and heathland communities. Trends in Ecology and Evolution 14: 63-68.
CrossRef | Gscholar
Bennion H, Carvalho L, Sayer CD, Simpson GL, Wischnewski J (2012)
Identifying from recent sediment records the effects of nutrients and climate on diatom dynamics in Loch Leven. Freshwater Biology 57: 2015-2029.
CrossRef | Gscholar
Bobbink R, Hettelingh JS (2011)
Review and revision of empirical critical loads and dose-response relationships. In: Proceedings of an expert workshop. Noordwijkerhout (Netherlands), 23-25 Jun 2010. Coordination Centre for Effects, National Institute for Public Health and the Environment (RIVM), Bilthoven, The Netherlands, pp. 246.
Bokdam J, Gleichman JM (2000)
Effects of grazing by free-ranging cattle on vegetation dynamics in a continental north-west European heathland. Journal of Applied Ecology 37: 415-431.
CrossRef | Gscholar
Britton AJ, Fisher JM (2007)
Interactive effects of nitrogen deposition, fire and grazing on diversity and composition of low-alpine prostrate Calluna vulgaris heathland. Journal of Applied Ecology 44: 125-135.
CrossRef | Gscholar
Britton AJ, Pearce ISK, Jones B (2005)
Impacts of grazing on montane heath vegetation in Wales and implications for the restoration of montane areas. Biological Conservation 125: 515-524.
CrossRef | Gscholar
Bullock JM, Pakeman RJ (1997)
Grazing of lowland heath in England: management methods and their effects on healthland vegetation. Biological Conservation 79: 1-13.
CrossRef | Gscholar
Colwell RK, Gotelli NJ, Rahbek C, Entsminger GL, Farrell C, Graves GR (2009)
Peaks, plateaus, canyons, and craters: the complex geometry of simple mid-domain effect models. Evolutionary Ecology Research 11: 355-370.
Online | Gscholar
Common TG, Wright IA, Grant SA (1998)
The effect of grazing by cattle on animal performance and floristic composition in Nardus-dominated swards. Grass and Forage Science 53: 260-269.
CrossRef | Gscholar
Critchley CNR, Adamson HF, Mclean BML, Davies OD (2008)
Vegetation dynamics and livestock performance in system-scale studies of sheep and cattle grazing on degraded upland wet heath. Agriculture, Ecosystems and Environment 128: 59-67.
CrossRef | Gscholar
Critchley CNR, Mitchell RJ, Rose RJ, Griffiths JB, Jackson E, Scott H, Davies OD (2013)
Re-establishment of Calluna vulgaris (L.) Hull in an eight-year grazing experiment on upland acid grassland. Journal for Nature Conservation 21: 22-30.
CrossRef | Gscholar
D’Odorico P, Okin GS, Bestelmeyer BT (2012)
A synthetic review of feedbacks and drivers of shrub encroachment in arid grasslands. Ecohydrology 5: 520-530.
CrossRef | Gscholar
Fagúndez J (2012)
Heathlands confronting global change: drivers of biodiversity loss from past to future scenarios. Annals of Botany 111 (2): 151-172.
CrossRef | Gscholar
Fernandes PM, Davies GM, Ascoli D, Fernández C, Moreira F, Rigolot E, Stoof CR, Vega JA, Molina D (2013)
Prescribed burning in southern Europe: developing fire management in a dynamic landscape. Frontiers in Ecology and the Environment 11: e4-e14.
CrossRef | Gscholar
Fottner S, Härdtle W, Niemeyer M, Niemeyer T, Von Oheimb G, Meyer H, Mockenhaupt M (2007)
Impact of sheep grazing on nutrient budgets of dry heathlands. Applied Vegetation Science 10: 391-398.
CrossRef | Gscholar
Fraser MD, Theobald VJ, Dhanoa MS, Davies OD (2011)
Impact on sward composition and stock performance of grazing Molinia-dominant grassland. Agriculture, Ecosystems and Environment 144: 102-106.
CrossRef | Gscholar
Friedrich U, Von Oheimb G, Dziedek C, Kriebitzsch W, Selbmann K, Härdtle W (2011)
Mechanisms of purple moor-grass (Molinia caerulea) encroachment in dry heathland ecosystems with chronic nitrogen inputs. Environmental Pollution 159: 3553-3559.
CrossRef | Gscholar
García RR, Fraser MD, Celaya R, Mendes Ferreira LM, García U, Garcia KO (2013)
Grazing land management and biodiversity in the Atlantic European heathlands: a review. Agroforestry Systems 87: 19-43.
CrossRef | Gscholar
Grant SA, Torvell L, Sim EM, Small JL, Armstrong RH (1996)
Controlled grazing studies on Nardus grassland: effects of between-tussock sward height and species of grazer on Nardus utilization and floristic composition in two fields in Scotland. Journal of Applied Ecology 33: 1053-1064.
CrossRef | Gscholar
Grytnes JA, Heegaard E, Romdal TS (2008)
Can the mass effect explain the mid-altitudinal peak in vascular plant species richness? Basic and Applied Ecology 9: 373-382.
CrossRef | Gscholar
Hargrove WW, Pickering J (1992)
Pseudoreplication: a sine qua non for regional ecology. Landscape Ecology 6: 251-258.
CrossRef | Gscholar
Hester AJ, Miller DR, Towers W (1996)
Landscape-scale vegetation change in the Cairngorms, Scotland, 1946-1988: implications for land management. Biological Conservation 77: 41-51.
CrossRef | Gscholar
Hobbs R (2009)
Woodland restoration in Scotland: ecology, history, culture, economics, politics and change. Journal of Environmental Management 90: 2857-2865.
CrossRef | Gscholar
Hulme PD, Pakeman RJ, Torvell L, Fisher JM, Gordon IJ (1999)
The effects of controlled sheep grazing on the dynamics of upland Agrostis-Festuca grassland. Journal of Applied Ecology 36: 886-900.
CrossRef | Gscholar
Hulme PD, Merrell BG, Torvell L, Fisher JM, Small JL, Pakeman RJ (2002)
Rehabilitation of degraded Calluna vulgaris (L.) Hull-dominated wet heath by controlled sheep grazing. Biological Conservation 107: 351-363.
CrossRef | Gscholar
Humphrey JW, Ferris F, Quine CP (2003)
Biodiversity in Britain’s planted forests. Forestry Commission, Edinburgh, UK, pp. 117.
Online | Gscholar
Hurlbert SH (1984)
Pseudoreplication and the design of ecological field experiments. Ecological Monographs 54 (2): 187-211.
CrossRef | Gscholar
Kozlov MV, Hurlbert SH (2006)
Pseudoreplication, chatter, and the international nature of science: a response to DV Tatarnikov. Journal of Fundamental Biology (Moscow) 67: 145-152.
Online | Gscholar
Körner C (2007)
The use of “altitude” in ecological research. Trends in Ecology and Evolution 22: 569-574.
CrossRef | Gscholar
Littlewood NA, Pakeman RJ, Woodin SJ (2006)
A field assessment of the success of moorland restoration in the rehabilitation of whole plant assemblages. Applied Vegetation Science 9: 295-306.
CrossRef | Gscholar
Littlewood NA, Greenwood S, Quin SLO, Pakeman RJ, Woodin SJ (2014)
Long-term trends in restored moorland vegetation assemblages. Community Ecology 15: 104-112.
CrossRef | Gscholar
Lomolino MV (2001)
Elevation gradients of species density: historical and prospective views. Global Ecology and Biogeography 10: 3-13.
CrossRef | Gscholar
Maddock A (2011)
UK biodiversity action plan priority habitat descriptions. JNCC, Peterborough, UK, pp. 100.
Online | Gscholar
Marrs RH, Phillips JDP, Todd PA, Ghorbani J, Le Duc MG (2004)
Control of Molinia caerulea on upland moors. Journal of Applied Ecology 41: 398-411.
CrossRef | Gscholar
Mitchell RJ, Rose RJ, Palmer SCF (2008)
Restoration of Calluna vulgaris on grass-dominated moorlands: the importance of disturbance, grazing and seeding. Biological Conservation 141: 2100-2111.
CrossRef | Gscholar
Moleele NM, Perkins JS (1998)
Encroaching woody plant species and boreholes: is cattle density the main driving factor in the Olifants Drift communal grazing lands, south-eastern Botswana? Journal of Arid Environments 40: 245-253.
CrossRef | Gscholar
Newton AC, Stewart GB, Myers G, Diaz A, Lake S, Bullock JM, Pullin AS (2009)
Impacts of grazing on lowland heathland in north-west Europe. Biological Conservation 142: 935-947.
CrossRef | Gscholar
Oksanen L (2001)
Logic of experiments in ecology: is pseudoreplication a pseudoissue? Oikos 94: 27-38.
CrossRef | Gscholar
Oksanen J, Blanchet FG, Kindt R, Legendre P, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Wagner H (2014)
vegan: community Ecology Package. R package version 2.0-3, web resource.
Online | Gscholar
Olff H, Ritchie ME (1998)
Effects of herbivores on grassland plant diversity. Trends in Ecology and Evolution 13: 261-265.
CrossRef | Gscholar
Pakeman RJ (2004)
Consistency of plant species, trait responses to grazing along a productivity gradient: a multi-site analysis. Journal of Ecology 92: 893-905.
CrossRef | Gscholar
Pakeman R, Hulme PD, Torvell L, Fisher JM (2003)
Rehabilitation of degraded dry heather (Calluna vulgaris (L.) Hull) moorland by controlled sheep grazing. Biological Conservation 114: 389-400.
CrossRef | Gscholar
R Development Core Team (2012)
R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
Online | Gscholar
Rawes M (1981)
Further results of excluding sheep from high-level grasslands in the North Pennines. Journal of Ecology 69: 651-669.
CrossRef | Gscholar
Rodwell JS (1992)
British plant communities: volume 3. Grasslands and montane communities. Cambridge University Press, Cambridge, UK, pp. 552.
Soil Survey of Scotland Staff (1981)
Soil maps of Scotland at a scale of 1:250 000. Macaulay Institute for Soil Research, Aberdeen, Scotland, UK.
Online | Gscholar
Stace C (2010)
New flora of the British isles. Cambridge University Press, Cambridge, UK, pp. 1236.
Online | Gscholar
Stevenson AC, Thompson DBA (1993)
Long-term changes in the extent of heather moorland in upland Britain and Ireland: palaeoecological evidence for the importance of grazing. The Holocene 3: 70-76.
CrossRef | Gscholar
Suding KN, Gross KL, Houseman GR (2004)
Alternative states and positive feedbacks in restoration ecology. Trends in Ecology and Evolution 19: 46-53.
CrossRef | Gscholar
Thompson DBA, MacDonald AJ, Marsden JH, Galbraith CA (1995)
Upland heather moorland in Great Britain: a review of international importance, vegetation change and some objectives for nature conservation. Biological Conservation 71: 163-178.
CrossRef | Gscholar
Vandvik V, Heegaard E, Måren IE, Aarrestad PA (2005)
Managing heterogeneity: the importance of grazing and environmental variation on post-fire succession in heathlands. Journal of Applied Ecology 42: 139-149.
CrossRef | Gscholar
Verhagen R, Klooker J, Bakker JP, Van Diggelen R (2001)
Restoration success of low-production plant communities on former agricultural soils after top-soil removal. Applied Vegetation Science 4: 75-82.
CrossRef | Gscholar
Watt AS, Jones EW (1948)
The ecology of the Cairngorms: part 1. The environment and the altitudinal zonation of the vegetation. Journal of Ecology 36: 283-304.
CrossRef | Gscholar
Webb NR (1998)
The traditional management of European heathlands. Journal of Applied Ecology 35: 987-990.
CrossRef | Gscholar
Welch D, Scott D (1995)
Studies in the grazing of heather moorland in northeast Scotland. VI. 20-year trends in botanical composition. Journal of Applied Ecology 32: 596-611.
CrossRef | Gscholar

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