Abstract
A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock.
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References
Bartolome J.W. 1979. Germination and seedling establishment in California annual grassland. Journal of Ecology 67: 273–281.
Boeken B., Lipchin C., Gutterman Y. and van Rooyen N. 1998.Annual plant community responses to density of small-scale soil disturbances in the Negev Desert of Israel. Oecologia 114: 106–117.
Boeken B. and Shachak M. 1994. Desert plant communities in human-made patches. Implications for management. Ecological Applications 4: 702–716.
Bullock J.M., Clear Hill B., Dale M.P. and Silvertown J. 1994. An experimental study of vegetation change due to grazing in a species-poor grassland and the role of seedling recruitment into gaps. Journal of Applied Ecology 31: 493–507.
Canals R.M. and Sebastià M.T. 2000. Soil nutrient fluxes and vegetation changes on molehills. Journal of Vegetation Science 11: 23–30.
Collins S.L. and Barber S.C. 1985. Effects of disturbance on diversity in mixed-grass prairie. Vegetatio 64: 87–94.
Fenner M. 2000. Seeds: The Ecology of Regeneration in Plant Communities. CABI Publishing, New York.
Fenner M. and Kitajima K. 1999. Seed and Seedling Ecology. In: Pugnaire F.I. and Valladares F. (eds), Handbook of Functional Plant Ecology. Marcel Dekker Inc., New York, pp. 589–621.
Fernández Alés R., Laffarga J.M. and Ortega F. 1993. Strategies in Mediterranean grassland annuals in relation to stress and disturbance.Journal of Vegetation Science 4: 313–322.
Fields M.J., Coffin D.P. and Gosz J.R. 1999. Burrowing activities of kangaroo rats and patterns in plant species dominance at a shortgrass steppe-desert grassland ecotone. Journal of Vegetation Science 10: 123–130.
Goldberg D. and Estabrook G.F. 1998. Separating the effects of number of individuals sampled and competition on species diversity: an experimental and analytic approach. Journal of Ecology 86: 983–988.
Goldberg D.E. and Gross K.L. 1988. Disturbance regimes of midsuccessional old fields. Ecology 00: 1677–1688.
Gómez-García D., Borghi C.E. and Giannoni S.M. 1995. Vegetation differences caused by pine vole mound building in subalpine plant communities in the Spanish Pyrenees. Vegetatio 117: 61–67.
Gutterman Y. 1994. Strategies of seed dispersal and germination in plant inhabiting deserts. Botanical Review 60: 373–425.
Hobbs R.J. and Huenneke L.F. 1992. Disturbance, diversity, and invasion: implications for conservation. Conservation Biology 6: 324–337.
Hobbs R.J. and Mooney H.A. 1985. Community and population dynamics of serpentine grassland annuals in relation to gopher disturbance. Oecologia (Berlin) 67: 342–351.
Hobbs R.J. and Mooney H.A. 1991. Effects of rainfall variability and gopher disturbance on serpentine annual grassland dynamics.Ecology 72: 59–68.
Hobbs R.J. and Mooney H.A. 1995. Spatial and temporal variability in California annual grassland: results from a long-term study. Journal of Vegetation Science 6: 43–56.
Huntly N. and Reichman O.J. 1994. Effects of subterranean mammalian herbivores on vegetation. Journal of Mammalogy 75: 852–859.
Hurlbert S.H. 1984. Pseudoreplication and the design of ecological field experiments. Ecological Monograph 54: 187–211.
Inouye R.S., Huntly N.J., Tilman D. and Tester J.R. 1987. Pocket gophers (Geomys bursarius), vegetation, and soil nitrogen along a successional sere in east central Minesota. Oecologia (Berlin) 72: 178–184.
Jones C.G., Lawton J.H. and Shachak M. 1994. Organisms as ecosystem engineers. Oikos 69: 373–386.
Lavorel S., Lepart J., Debussche M., Lebreton J.D. and Beffy J.L. 1994. Small scale disturbances and the maintenance of species diversity in Mediterranean old fields. Oikos 70: 455–473.
Leck M.A. and Graveline K.J. 1979. The seed bank of a freshwater tidal marsh. American Journal of Botany 66: 1006–1015.
Levassor C., Ortega M. and Peco B. 1990. Seed bank dynamics of Mediterranean pastures subjected to mechanical disturbance.Journal of Vegetation Science 1: 339–344.
Martinsen G.D., Cushman J.H. and Whitham T.G. 1990. Impact of pocket gopher disturbance on plant species diversity in a shortgrass prairie community. Oecologia 83: 132–138.
Milchunas D.G., Sala O.E. and Lauenroth W.K. 1988. A generalized model of the effects of grazing by large herbivores on grasslands community structure. The American Naturalist 132: 87–106.
Milton S.J., Dean W.R.J. and Klotz S. 1997. Effects of small-scale animal disturbances on plant assemblages of set-aside land in Central Germany. Journal of Vegetation Science 8: 45–54.
Noy-Meir I. 1988. Dominant grasses replaced by ruderal forbs in a vole year in undergrazed Mediterranean grasslands in Israel.Journal of Biogeography 15: 579–587.
Noy-Meir I., Gutman M. and Kaplan Y. 1989. Responses of Mediterranean grassland plants to grazing and protection. Journal of Ecology 77: 290–310.
Noy-Meir I. 1998. Effects of grazing on Mediterranean grasslands: the community level. In: Papanastasis V.P. and Peter D. (eds), Ecological Basis of Livestock Grazing in Mediterranean Ecosystems.European Commission, Brussels, Belgium, pp. 27–39.
Olff H. and Ritchie M.E. 1998. Effects of herbivores on grassland plant diversity. Trends in Ecology & Evolution 13: 261–265.
Peet R.K. 1974. The measurement of species diversity. Annual Review of Ecology and Systematics 5: 285–307.
Pickett S.T.A. and White P.S. 1985. The Ecology of Natural Disturbances and Patch Analysis. Academic Press, Orlando.
Pielou E.C. 1966. The measurement of diversity in different types of biological collections. Journal of Theoretical Biology 13: 131–144.
Pineda F.D., Casado M.A., Peco B., Olmeda C. and Levassor C. 1987. Temporal changes in therophytic communities across the boundary of a disturbed-intact ecosystems. Vegetatio 69: 109– 114.
Platt W.J. and Weis J.M. 1977. Resource partitioning and competition within a guild of fugitive prairie plants. The American Naturalist 111: 479–513.
Platt W.J. 1975. The colonization and formation of equilibrium plant species associations on badger disturbances in a tall-grass prairie. Ecological Monograph 45: 285–305.
Rebollo S., Pérez-Camacho L., García de Juan M.T., Rey-Benayas J.M. and Gómez-Sal A. 2001. Seedling emergence and survival in an annual plant community: relationships with germinable seeds, litter and density of con-and non conspecifics. Oikos 95: 485–495.
Reichman O.J. and Smith S.C. 1985. Impact of pocket gopher burrows on overlying vegetation. Journal of Mammalogy 66: 720– 725.
Ritchie M.E. and Olff H. 1999. Herbivory diversity and plant dynamics: compensatory and additive effects. In: Olff H., Brown V.K. and Drent R.H. (eds), Herbivores: Between Plants and Predators. Blackwell Science, Oxford, pp. 175–204.
Shannon C.E. and Weaver W. 1949. The Mathematical Theory of Communication. University of Illinois Press, Urbana, Illinois, USA.
Sirotnak J.M. and Huntly N.J. 2000. Direct and indirect effects of herbivores on nitrogen dynamics: voles in riparian areas. Ecology 81: 78–87.
Soriguer R.C. and Amat J.A. 1980. On the structure and function of the burrows of the Mediterranean vole (Pitymys duodecimcostatus).Acta Theriologica 25: 268–270.
Sousa W.P. 1984. The role of disturbance in natural communities.Annual Review of Ecology and Systematics 15: 353–391.
Thompson K. and Grime J.P. 1979. Seasonal variation in the seed banks of herbaceous species in ten contrasting habitats. Journal of Ecology 67: 893–921.
Tilman D. 1983. Plant succession and gopher disturbance along an experimental gradient. Oecologia (Berlin) 60: 285–292.
Tutin T.G. et al. 1964–1981. Flora Europaea. Cambridge University Press, Cambridge.
Umbanhowar E.C. Jr. 1995. Revegetation of earthen mounds along a topographic-productivity gradient in a northern mixed prairie.Journal of Vegetation Science 6: 637–646.
Whittaker R.H. 1952. A study of summer foliage insect communities in the Great Smoky Mountains. Ecological Monograph 22: 1–44.
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Rebollo, S., Pérez-Camacho, L., Valencia, J. et al. Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes. Plant Ecology 169, 227–243 (2002). https://doi.org/10.1023/A:1026059006184
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DOI: https://doi.org/10.1023/A:1026059006184