A plant trait analysis of responses to grazing in a long-term experiment

Summary 1There are few long-term experimental studies of plant community responses to changes in grazing intensity. Here we report species’ changes in a mesotrophic grassland after 12 years of a grazing experiment and relate these changes to species’ life-history traits. 2The experiment was set up in 1986 on an extensified species-poor grassland in lowland UK. Treatments comprised sheep grazing vs. no grazing in winter, grazing vs. no grazing in spring, and two grazing intensities in summer, in a 2 × 2 × 2 factorial design with two replicate blocks. 3Point quadrat surveys in 1998 showed responses to grazing treatments by 17 of the 22 most common species. Species showed different responses, many of which were specific to a grazing season. Community changes were similar under spring and winter grazing, but the heavier summer grazing had different consequences. Species richness was increased by spring grazing, decreased by heavier summer grazing and unaffected by winter grazing. 4More species responded to treatments in the 1998 survey compared with a survey in 1990. Furthermore, the whole experimental grassland had changed between the surveys, probably as a result of falling soil fertility. The two dominant grasses had declined drastically and most other species had increased in abundance. Five new species were found in 1998. 5Intensive surveys of dicotyledonous species in 1998 showed five of the 12 most common species had responded to grazing treatments. In most cases dicotyledonous species had increased in abundance under heavier grazing in one or more season, and species richness was increased by spring and winter grazing. Compared with a 1991 survey, the number of species responding to treatments had increased by 1998 and seven new species were found. 6We tested whether species’ responses to grazing were linked to life-history traits according to three hypotheses: that heavier grazing would disadvantage (i) species grazed preferentially, (ii) species less able to colonize gaps or (iii) more competitive species. Mechanisms differed among seasonal treatments. Responses to heavier summer grazing were linked strongly to gap colonization ability. Responses to spring and winter grazing were positively related to grazer selectivity, a surprising result that might be explained if selectivity was positively related to plant regrowth ability. 7This study shows the need for long-term experimental analyses of community responses to grazing as vegetation responses may develop over a long time. The traits analysis suggests it may be possible to predict changes in species composition under grazing through an understanding of the mechanisms of plant responses. Grassland managers require such information in order to manipulate grazing regimes to achieve, for example, diversification or weed control.

[1]  D. Elston,et al.  Seasonal Pattern of Leaf Growth and Senescence of Nardus stricta and Responses of Tussocks to Differing Severity, Timing and Frequency of Defoliation , 1996 .

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

[3]  L. K. Ward,et al.  Succession of disturbed and undisturbed chalk grassland at Aston Rowant National Nature Reserve: dynamics of species changes. , 1990 .

[4]  A. Diaz Can plant palatability trials be used to predict the effect of rabbit grazing on the flora of ex-arable land? , 2000 .

[5]  Sandra Lavorel,et al.  Plant response to disturbance in a Mediterranean grassland: How many functional groups? , 1999 .

[6]  S. Lavorel,et al.  Plant functional classifications: from general groups to specific groups based on response to disturbance. , 1997, Trends in ecology & evolution.

[7]  Wolfgang Cramer,et al.  Plant functional types and disturbance dynamics – Introduction , 1999 .

[8]  M. Bullard,et al.  The conservation management of mesotrophic (meadow) grassland in northern England. 1. Effects of grazing, cutting date and fertilizer on the vegetation of a traditionally managed sward , 1996 .

[9]  M. Hartley Effect of scotch thistles on sheep growth rates. , 1983 .

[10]  Reversion of vegetation following the cessation of fertilizer application , 1996, Journal of vegetation science : official organ of the International Association for Vegetation Science.

[11]  T. A. Watt,et al.  The use of sheep grazing to recreate species-rich grassland from abandoned arable land , 1987 .

[12]  M. J. Liddle,et al.  Resistance, survival and recovery of trampled corals on the Great Barrier Reef , 1987 .

[13]  R. Clarke,et al.  Individual variation and the effects of weather, age and flowering history on survival and flowering of the long-lived perennial Gentiana pneumonanthe , 1998 .

[14]  J. P. Grime,et al.  Comparative Plant Ecology , 1988, Springer Netherlands.

[15]  E. C. Diekmahns,et al.  Grasses and legumes in British agriculture , 1973 .

[16]  Robin J. Pakeman,et al.  Grazing of lowland heath in England: Management methods and their effects on healthland vegetation , 1997 .

[17]  A. Magurran Ecological Diversity and Its Measurement , 1988, Springer Netherlands.

[18]  N. Aranwela,et al.  Methods of assessing leaf‐fracture properties , 1999 .

[19]  J. Treweek,et al.  An experimental study of the impact of seasonal sheep grazing on formerly fertilized grassland , 1996 .

[20]  J. Silvertown,et al.  Mapping the Microenvironment for Seed Germination in the Field , 1989 .

[21]  J. Kigel,et al.  Vegetation response to grazing management in a Mediterranean herbaceous community: a functional group approach , 2000 .

[22]  Stephens,et al.  Consequences of the Allee effect for behaviour, ecology and conservation. , 1999, Trends in ecology & evolution.

[23]  D. Tilman,et al.  The Park Grass Experiment: insights from the most long-term ecological study , 1994 .

[24]  V. Brown,et al.  The effects of grazing on local colonisation and extinction during early succession , 1991 .

[25]  J. Humphrey,et al.  Effects of late summer cattle grazing on the diversity of riparian pasture vegetation in an upland conifer forest , 2000 .

[26]  J. Rodwell,et al.  British Plant Communities. Vol. 3. Grasslands and Montane Communities. , 1994 .

[27]  J. Lepš,et al.  Use of paired plots and multivariate analysis for the determination of goat grazing preference , 1995 .

[28]  I. Gordon,et al.  The effects of controlled sheep grazing on the dynamics of upland Agrostis–Festuca grassland , 1999 .

[29]  J. Treweek,et al.  Integration of Sheep Production and Nature Conservation: Experimental Management , 1997 .

[30]  Sandra Lavorel,et al.  Grazing response groups among understorey plants in arid rangelands , 1999 .

[31]  T. Stohlgren,et al.  How grazing and soil quality affect native and exotic plant diversity in Rocky Mountain grasslands , 1999 .

[32]  R. H. Armstrong,et al.  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 , 1996 .

[33]  Scott E. Maxwell,et al.  Designing Experiments and Analyzing Data , 1991 .

[34]  Long-Term Dynamics of Standing Crop and Species Composition after the Cessation of Fertilizer Application to Mown Grassland , 1991 .

[35]  Kannan Soundararajan,et al.  Demography of Cirsium vulgare in a grazing experiment. , 1994 .

[36]  P. Kenney,et al.  Factors affecting diet selection by sheep. 2. Height and density of pasture , 1984 .

[37]  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 .

[38]  M. Gutman,et al.  Responses of mediterranean grassland plants to grazing and protection. , 1989 .

[39]  R. Tremont Life-History Attributes of Plants in Grazed and Ungrazed Grasslands on the Northern Tablelands of New South Wales , 1994 .

[40]  P. Keddy,et al.  Competitive Performance and Species Distribution in Shortline Plant Communities: A Comparative Approach , 1995 .

[41]  J. Silvertown,et al.  Tiller dynamics of two grasses: responses to grazing, density and weather , 1994 .

[42]  Elizabeth A. Kellogg,et al.  An ordinal classification for the families of flowering plants , 1998 .

[43]  Scott E. Maxwell,et al.  Designing Experiments and Analyzing Data , 1992 .

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

[45]  J. Bullock,et al.  Plant responses to grazing, and opportunities for manipulation. , 2000 .

[46]  J. F. Dormaar,et al.  EFFECT OF FORTY-FOUR YEARS OF GRAZING ON FESCUE GRASSLAND SOILS , 1998 .

[47]  M. Hill,et al.  Long-term effects of excluding sheep from hill pastures in North Wales , 1992 .

[48]  C. Loehle Problems with the triangular model for representing plant strategies , 1988 .

[49]  S. Rushton,et al.  The effects of grazing management on the vegetation of mesotrophic (meadow) grassland in Northern England , 1994 .

[50]  Ian F. Spellerberg,et al.  The Scientific Management of Temperate Communities for Conservation. , 1992 .

[51]  J. Silvertown,et al.  A phylogenetic approach to community assembly from a local species pool , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[52]  James M. Bullock,et al.  Gap colonization as a source of grassland community change: effects of gap size and grazing on the rate and mode of colonization by different species , 1995 .

[53]  H. Olff,et al.  Effects of herbivores on grassland plant diversity. , 1998, Trends in ecology & evolution.

[54]  J. S. Rodwell,et al.  British Plant Communities: British Plant Communities , 2000 .

[55]  D. Millward,et al.  The interactive effects of management on the productivity and plant community structure of an upland meadow: an 8-year field trial , 2000 .

[56]  Jonathan Silvertown,et al.  Gaps in the canopy: the missing dimension in vegetation dynamics , 1988 .

[57]  J. Ivins THE RELATIVE PALATABILITY OF HERBAGE PLANTS , 1952 .

[58]  R. Marrs Soil Fertility and Nature Conservation in Europe: Theoretical Considerations and Practical Management Solutions , 1993 .

[59]  H. K. Smith,et al.  COMPARATIVE STUDIES OF DIET SELECTION BY SHEEP AND CATTLE: THE HILL GRASSLANDS , 1985 .

[60]  H. Olff,et al.  Changes in nitrogen mineralization, tissue nutrient concentrations and biomass compartmentation after cessation of fertilizer application to mown grassland , 1994 .

[61]  K. McConway,et al.  community stability - a 60-year record of trends and outbreaks in the occurrence of species in the park grass experiment , 1995 .

[62]  S. Lavorel,et al.  Plant life-history attributes: their relationship to disturbance response in herbaceous vegetation. , 1995 .