Impact of herbivory by red deer (Cervus elaphus L.) on patterns and processes in subalpine grasslands in the Swiss National Park

The spatial grazing patterns of red deer (Cervus elaphus L.) in a subalpine grassland ecosystem of the Swiss National Park were analysed with reference to the phosphorus content in the topsoil and to the former agricultural management system. Changes in the composition of the vegetation and in plant species richness were studied on permanent plots set up between 1917 and 1945 in areas which today are heavily grazed by red deer, and were related to the development of the red deer population. The spatial grazing patterns of red deer are, 80 years after the Park’s foundation, still determined by former agricultural use. Red deer prefer the most phosphorus-rich, formerly irrigated parts for grazing. Grazing by red deer was found to be the driving force behind temporal changes in the vegetation observed on the subalpine grasslands. Plant succession has proceeded faster in the preferred grazing areas and the vegetation has adapted to the grazing pressure with a decrease in tall-growing plant species. At the same time, the abundance of unpalatable species has increased due to (i) morphological defences, (ii) spatial avoidance, i.e. by growing close to the ground and (iii) temporal avoidance because they have a short life-span (annuals). Changes in vegetation composition have coincided with a sharp increase in species richness. Between 1917 and 1999, the number of plant species has doubled on average. We found a highly significant correlation between the changes in the numbers of plant species and red deer, with species richness increasing with increasing deer density. # 2003 Elsevier B.V. All rights reserved.

[1]  G. Gauthier,et al.  Effect of grazing by greater snow geese on the production of graminoids at an arctic site (Bylot Island, NWT, Canada) , 1995 .

[2]  J. P. Grime,et al.  Competitive Exclusion in Herbaceous Vegetation , 1973, Nature.

[3]  H. Zoller Vegetationskarte des Schweizerischen Nationalparks : Erläuterungen , 1995 .

[4]  J. Connell Diversity in tropical rain forests and coral reefs. , 1978, Science.

[5]  G. Shaver,et al.  Vertebrate Herbivores and Northern Plant Communities: Reciprocal Influences and Responses , 1994 .

[6]  J. Rosenthal,et al.  Terrestrial plant tolerance to herbivory. , 1994, Trends in ecology & evolution.

[7]  J. P. Grime,et al.  A QUANTITATIVE ANALYSIS OF SHOOT PHENOLOGY AND DOMINANCE IN HERBACEOUS VEGETATION , 1977 .

[8]  J. Harper Population Biology of Plants , 1979 .

[9]  I. Gordon Vegetation community selection by ungulates on the Isle of Rhum. II: Vegetation community selection , 1989 .

[10]  G. Spatz Succession patterns on mountain pastures , 1980, Vegetatio.

[11]  David Tilman,et al.  Secondary Succession and the Pattern of Plant Dominance Along Experimental Nitrogen Gradients , 1987 .

[12]  H. Hess,et al.  Flora der Schweiz und Angrenzender Gebiete , 1972 .

[13]  A. Agrawal,et al.  The ecology and evolution of plant tolerance to herbivory. , 1999, Trends in ecology & evolution.

[14]  V. Nams,et al.  Nitrogen fertilization stimulates herbivory by snowshoe hares in the boreal forest , 1996 .

[15]  Browsing by Moose on Scots Pine in Relation to Plant Resource Availability , 1993 .

[16]  Dolph Schluter,et al.  Species diversity in ecological communities: historical and geographical perspectives. , 1993 .

[17]  David J. Augustine,et al.  Ungulate effects on the functional species composition of plant communities; herbivore selectivity and plant tolerance , 1998 .

[18]  J. P. Grime,et al.  Plant Strategies and Vegetation Processes. , 1980 .

[19]  R. Palo,et al.  Plant Defenses Against Mammalian Herbivory , 1991 .

[20]  M. Rosenzweig,et al.  How are diversity and productivity related , 1993 .

[21]  Michael L. Rosenzweig,et al.  Species Diversity in Space and Time , 1997 .

[22]  M. Hill,et al.  Data analysis in community and landscape ecology , 1987 .

[23]  J. P. Bakker,et al.  Micro-patterns in grassland vegetation created and sustained by sheep-grazing , 1984, Vegetatio.

[24]  P. Groffman,et al.  Ungulate stimulation of nitrogen cycling and retention in Yellowstone Park grasslands , 2000, Oecologia.

[25]  TABORD, a program for structuring phytosociological tables , 1978 .

[26]  I. Gordon,et al.  The influence of vegetation pattern on the grazing of heather moorland by red deer and sheep. I: The location of animals on grass/heather mosaics , 1995 .

[27]  H. Gleason,et al.  On the Relation Between Species and Area , 1922 .

[28]  M. Crawley,et al.  Red deer management and vegetation on the Isle of Rum , 2002 .

[29]  M. Schütz,et al.  Reconstruction of a long-term recovery process from pasture to forest , 2000 .

[30]  W. Carson,et al.  PLANT DENSITY DETERMINES SPECIES RICHNESS ALONG AN EXPERIMENTAL FERTILITY GRADIENT , 1999 .

[31]  T. Clutton‐Brock,et al.  Grazing and reproductive success of red deer: the effect of local enrichment by gull colonies , 1986 .

[32]  P. White,et al.  Scale Dependence and the Species-Area Relationship , 1994, The American Naturalist.

[33]  M. Crawley Herbivory: the Dynamics of Animal-plant Interactions , 1984 .

[34]  Michael J. Crawley,et al.  Colonization, Succession, and Stability , 1988 .

[35]  Matt A. Sanderson,et al.  Patterns of plant species richness in pasture lands of the northeast United States , 2000, Plant Ecology.

[36]  A. Magurran,et al.  Biological diversity : the coexistence of species on changing landscapes , 1994 .

[37]  M. Rico,et al.  Variation in structure and diversity in Mediterranean grasslands related to trophic status and grazing intensity , 1990 .

[38]  J. Altmann,et al.  Observational study of behavior: sampling methods. , 1974, Behaviour.

[39]  A. D. Fox,et al.  The effects of simulated spring goose grazing on the growth rate and protein content of Phleum pratense leaves , 1998, Oecologia.

[40]  J. P. Grime Control of species density in herbaceous vegetation , 1973 .

[41]  J. Silvertown,et al.  An experimental study of the effects of sheep grazing on vegetation change in a species-poor grassland and the role of seedling recruitment into gaps , 1994 .

[42]  D. Mccowan,et al.  SELECTION OF UPLAND SWARDS BY RED DEER (CERVUS ELAPHUS L.) ON RHUM , 1977 .

[43]  J. Jackson Feeding habits of deer , 1974 .

[44]  S. McNaughton,et al.  Promotion of the cycling of diet-enhancing nutrients by african grazers , 1997, Science.

[45]  A. Grootjans,et al.  Mechanisms of vegetation succession: a review of concepts and perspectives , 1993 .

[46]  J. Blair,et al.  Modulation of diversity by grazing and mowing in native tallgrass prairie , 1998, Science.

[47]  M. F. W. D. Vries,et al.  Foraging strategy of cattle in patchy grassland , 1994, Oecologia.

[48]  T. Clutton‐Brock,et al.  Sexual segregation and density-related changes in habitat use in male and female Red deer (Cerrus elaphus) , 1987 .

[49]  J. Oksanen Is the humped relationship between species richness and biomass an artefact due to plot size , 1996 .

[50]  J. Braun-Blanquet,et al.  Pflanzensoziologie: Grundzuge der Vegetationskunde. , 1967 .

[51]  S. Gardner,et al.  Consumer pressure, seed versus safe-site limitation, and plant population dynamics , 2000, Oecologia.

[52]  R. Jefferies,et al.  The Effects of Grazing by Lesser Snow Geese on the Vegetation of a Sub- Arctic Salt Marsh , 1984 .

[53]  P. Rothery,et al.  Effects of Grazing by Mountain Hares and Red Deer on the Production and Chemical Composition of Heather , 1981 .

[54]  V. Brown,et al.  Grazing and Vegetation Change: Deflected or Modified Succession? , 1992 .

[55]  James B. Grace,et al.  Perspectives on Plant Competition , 1991 .

[56]  P. J. Edwards,et al.  Vegetational and faunal changes in an area of heavily grazed woodland following relief of grazing , 1989 .

[57]  Berger‐Landefeldt Braun-Blanquet, J.: Pflanzensoziologie. Grundzüge der Vegetationskunde. 2. Aufl. Wien 1951 (Springer-Verlag). 2. Aufl. 631 S. mit 350 Textabb. DM 63.—, Ganzleinen DM 67.20 , 1952 .