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Testing seed-size predictions in Mediterranean annual grasslands

Published online by Cambridge University Press:  07 June 2010

Francisco M. Azcárate*
Affiliation:
Terrestrial Ecology Group (TEG), Departamento de Ecología, Universidad Autónoma de Madrid, Cantoblanco, E- 28049Madrid, Spain
Pablo Manzano
Affiliation:
Terrestrial Ecology Group (TEG), Departamento de Ecología, Universidad Autónoma de Madrid, Cantoblanco, E- 28049Madrid, Spain
Begoña Peco
Affiliation:
Terrestrial Ecology Group (TEG), Departamento de Ecología, Universidad Autónoma de Madrid, Cantoblanco, E- 28049Madrid, Spain
*
*Correspondence Fax: +34 91 49 78 001 Email: fm.azcarate@uam.es

Abstract

On the basis of previous research, we predict that Mediterranean grasslands should show larger-seeded annuals in: (1) more-arid grasslands; (2) more-fertile soils; (3) less-grazed grasslands; and (4) grasslands with lower intensities of seed predation by ants. To test these predictions, we set 29 sampling units of 50 m × 50 m in a 1000 km2 grassland area in Central Spain, and characterized them according to the former factors. We then recorded annual vegetation using ten quadrats of 20 cm × 20 cm in each sampling unit. Seed size at the community level was described using six variables: (1) mean seed mass; (2) standard deviation of seed mass; (3) weighted mean seed mass (by species frequencies); (4) proportion of small-seeded annuals; (5) proportion of medium-seeded annuals; and (6) proportion of large-seeded annuals. Most climate variables (mean annual temperature, length of the summer drought, water balance and mean annual precipitation) correlated with seed-size descriptors, showing that large-seeded annuals increase in warmer and more-arid communities. Mean seed size was modelled as a function of mean annual temperature and grazing pressure. According to this model, warmer and less-grazed communities tend to show a smaller mean seed size. These results confirm the importance of seed-size descriptors at the community level in Mediterranean grasslands, and the role of climate and grazing as major drivers in these communities. Conversely, hypotheses about soil fertility and seed predation by ants were not supported by our results.

Type
Research Article
Copyright
Copyright © Cambridge University Press 2010

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References

Aarssen, L.W. and Jordan, C.Y. (2001) Between-species patterns of covariation in plant size, seed size and fecundity in monocarpic herbs. Ecoscience 8, 471477.CrossRefGoogle Scholar
Ackerly, D.D., Knight, C.A., Weiss, S.B., Barton, K. and Starmer, K.P. (2002) Leaf size, specific leaf area and microhabitat distribution of chaparral woody plants, contrasting patterns in species level and community level analyses. Oecologia 130, 449457.CrossRefGoogle ScholarPubMed
Acosta, B., Sánchez-Jardon, L., Del Pozo, A., Garcia-Ibañez, E., Casado, M.A., Montalvo, J. and Pineda, F.D. (2008) Grassland species composition and morpho-functional traits along an altitudinal gradient in a Mediterranean environment, relationship with soil water availability and evaporative dynamic. Acta Oecologica 34, 2637.CrossRefGoogle Scholar
Azcárate, F.M. and Peco, B. (2003) Spatial patterns of seed predation by harvester ants (Messor Forel) in Mediterranean grassland and scrubland. Insectes Sociaux 50, 120126.CrossRefGoogle Scholar
Azcárate, F.M. and Peco, B. (2004) Seed predation by ants (Messor spp.) in Iberian Dehesa grassland and scrubland. Proceedings 10th MEDECOS Conference, April–May 2004, Rhodes, Greece.Google Scholar
Azcárate, F.M. and Peco, B. (2006) Effects of seed predation by ants on Mediterranean grassland related to seed size. Journal of Vegetation Science 17, 353360.CrossRefGoogle Scholar
Azcárate, F.M. and Peco, B. (2007) Harvester ants (Messor barbarus) as disturbance agents in Mediterranean grasslands. Journal of Vegetation Science 18, 103110.CrossRefGoogle Scholar
Azcárate, F.M., Sánchez, A.M., Arqueros, L. and Peco, B. (2002) Abundance and habitat segregation in Mediterranean grassland species: the importance of seed weight. Journal of Vegetation Science 13, 159166.CrossRefGoogle Scholar
Azcárate, F.M., Arqueros, L., Sánchez, A.M. and Peco, B. (2005) Seed and fruit selection by harvester ants, Messor barbarus, in Mediterranean grassland and scrubland. Functional Ecology 19, 273283.CrossRefGoogle Scholar
Bekker, R.M., Bakker, J.P., Grandin, U., Kalamees, R., Milberg, P., Poschlod, P., Thompson, K. and Willems, J.H. (1998) Seed size, shape and vertical distribution in the soil: indicators of seed longevity. Functional Ecology 12, 834842.CrossRefGoogle Scholar
Belsky, J.A. (1992) Effects of grazing, competition, disturbance and fire on species composition and diversity in grassland communities. Journal of Vegetation Science 3, 187200.CrossRefGoogle Scholar
Benjamini, Y.Hochberg, Y. (1995) Controlling the false discovery rate: a practical and powerful approach to multiple testing. Journal of the Royal Statistical Society, Series B 57, 289300.Google Scholar
Bruun, H.H.andPoschlod, P. (2006) Why are small seeds dispersed through animal guts: large numbers or seed size per se? Oikos 113, 402411.CrossRefGoogle Scholar
Crist, T.O. and MacMahon, J.A. (1992) Harvester ant foraging and shrub steppe seeds interactions of seed resources and seed use. Ecology 73, 17681779.CrossRefGoogle Scholar
Davidson, D.W., Inouye, R.S. and Brown, J.H. (1984) Granivory in a desert ecosystem–experimental evidence for indirect facilitation of ants by rodents. Ecology 65, 17801786.CrossRefGoogle Scholar
de Pablo, C.L., Peco, B., Galiano, E.F., Nicolás, J.P. and Pineda, F.D. (1982) Space–time variability in Mediterranean pastures analyzed with diversity parameters. Vegetatio 50, 113125.CrossRefGoogle Scholar
de Pablos, I. and Peco, B. (2007) Diaspore morphology and the potential for attachment to animal coats in Mediterranean species: an experiment with sheep and cattle coats. Seed Science Research 17, 109114.CrossRefGoogle Scholar
Dean, W.R.J. and Yeaton, R.I. (1992) The importance of harvester ant Messor capensis nest-mounds as germination sites in the southern Karoo, South-Africa. African Journal of Ecology 30, 335345.CrossRefGoogle Scholar
Detrain, C. and Pasteels, J.M. (2000) Seed preferences of the harvester ant Messor barbarus in a Mediterranean mosaic grassland (Hymenoptera: Formicidae). Sociobiology 35, 3548.Google Scholar
Diaz, S., Lavorel, S., Mcintyre, S., Falczuk, V., Casanoves, F., Milchunas, D.G., Skarpe, C., Rusch, G., Sternberg, M., Noy-Meir, I., Landsberg, J., Zhang, W., Clark, H. and Campbell, B.D. (2007) Plant trait responses to grazing. A global synthesis. Global Change Biology 13, 313341.CrossRefGoogle Scholar
Fenner, M. and Thompson, K. (2005) The ecology of seeds. Cambridge, UK, Cambridge University Press.CrossRefGoogle Scholar
Figueroa, M.E. and Davy, A.J. (1991) Response of Mediterranean grassland species to changing rainfall. Journal of Ecology 79, 925941.CrossRefGoogle Scholar
García, L.V. (2003) Controlling the false discovery rate in ecological research. Trends in Ecology and Evolution 18, 553554.CrossRefGoogle Scholar
Gaussen, H. (1955) Expression des milieux par des formules écologiques; leur représentation cartographique. Colloques Internationaux du Centre Naturel de Recherche Scientifique 59, 257269.Google Scholar
Hanley, M.E., Cordier, P.K., May, O. and Kelly, C.K. (2007) Seed size and seedling growth: differential response of Australian and British Fabaceae to nutrient limitation. New Phytologist 174, 381388.CrossRefGoogle ScholarPubMed
Jurado, E. and Westoby, M. (1992) Seedling growth in relation to seed size among species of arid Australia. Journal of Ecology 80, 407416.CrossRefGoogle Scholar
Jurjavcic, N.L., Harrison, S. and Wolf, A.T. (2002) Abiotic stress, competition, and the distribution of the native annual grass Vulpia microstachys in a mosaic environment. Oecologia 130, 555562.CrossRefGoogle Scholar
Kahmen, S.andPoschlod, P. (2008a) Does germination success differ with respect to seed mass and germination season? Experimental testing of plant functional trait responses to grassland management. Annals of Botany 101, 541548.CrossRefGoogle ScholarPubMed
Kahmen, S. and Poschlod, P. (2008b) Effects of grassland management on plant functional trait composition. Agriculture, Ecosystems and Environment 128, 137145.CrossRefGoogle Scholar
Kos, M. and Poschlod, P. (2008) Correlates of inter-specific variation in germination response to water stress in a semi-arid savannah. Basic and Applied Ecology 9, 645652.CrossRefGoogle Scholar
Lavorel, S., Rochette, C. and Lebreton, J.D. (1999) Functional groups for response to disturbance in Mediterranean old fields. Oikos 84, 480498.CrossRefGoogle Scholar
Leishman, M.R. and Westoby, M. (1994) The role of seed size in seedling establishment in dry soil conditions. Experimental evidence from semiarid species. Journal of Ecology 82, 249258.CrossRefGoogle Scholar
López, F., Acosta, F.J. and Serrano, J.M. (2000) Asymmetric interactions between plants and seed-harvesting ants in a Mediterranean pasture. Ecological Research 15, 449452.CrossRefGoogle Scholar
Marañón, T. and Grubb, P.J. (1993) Physiological basis and ecological significance of the seed size and relative growth-rate relationship in Mediterranean annuals. Functional Ecology 7, 591599.CrossRefGoogle Scholar
McIntyre, S. and Lavorel, S. (2001) Livestock grazing in subtropical pastures: steps in the analysis of attribute response and plant functional types. Journal of Ecology 89, 209226.CrossRefGoogle Scholar
Milberg, P. and Lamont, B.B. (1997) Seed/cotyledon size and nutrient content play a major role in early performance of species on nutrient-poor soils. New Phytologist 137, 665672.CrossRefGoogle Scholar
Milberg, P., Perez-Fernandez, M.A. and Lamont, B.B. (1998) Seedling growth response to added nutrients depends on seed size in three woody genera. Journal of Ecology 86, 624632.CrossRefGoogle Scholar
Moles, A.T. and Westoby, M. (2004) Seedling survival and seed size: a synthesis of the literature. Journal of Ecology 92, 372383.CrossRefGoogle Scholar
Montalvo, J., Casado, M.A., Levassor, C. and Pineda, F.D. (1993) Species-diversity patterns in Mediterranean grasslands. Journal of Vegetation Science 4, 213222.CrossRefGoogle Scholar
Monturiol, F. and Alcalá del Olmo, L. (1990) Mapa de asociaciones de suelos de la Comunidad de Madrid escala 1:200.000. Madrid, Spain, Consejo Superior de Investigaciones Científicas.Google Scholar
Moran, M.D. (2003) Arguments for rejecting the sequential Bonferroni in ecological studies. Oikos 100, 403405.CrossRefGoogle Scholar
Moro, M.J., Pugnaire, F.I., Haase, P. and Puigdefabregas, J. (1997) Effect of the canopy of Retama sphaerocarpa on its understorey in a semiarid environment. Functional Ecology 11, 425431.CrossRefGoogle Scholar
Mouissie, A.M., Van Der Veen, C.E.J., Veen, G.F. and Van Diggelen, R. (2005) Ecological correlates of seed survival after ingestion by fallow deer. Functional Ecology 19, 284290.CrossRefGoogle Scholar
Murray, B.R., Brown, A.H.D., Dickman, C.R. and Crowther, M.S. (2004) Geographical gradients in seed mass in relation to climate. Journal of Biogeography 31, 379388.CrossRefGoogle Scholar
Ninyerola, M., Pons, X. and Roure, J.M. (2005) Atlas Climático Digital de la Península Ibérica. Metodología y aplicaciones en bioclimatología y geobotánica. Bellaterra, Spain, Universidad Autónoma de Barcelona.Google Scholar
Noy-Meir, I., Gutman, M. and Kaplan, Y. (1989) Responses of Mediterranean grassland plants to grazing and protection. Journal of Ecology 77, 290310.CrossRefGoogle Scholar
Osem, Y., Perevolotsky, A. and Kigel, J. (2006) Size traits and site conditions determine changes in seed bank structure caused by grazing exclusion in semiarid annual plant communities. Ecography 29, 1120.CrossRefGoogle Scholar
Pakeman, R.J., Digneffe, G. and Small, J.L. (2002) Ecological correlates of endozoochory by herbivores. Functional Ecology 16, 296304.CrossRefGoogle Scholar
Pakeman, R.J., Garnier, E., Lavorel, S., Ansquer, P., Castro, H., Cruz, P., Dolezal, J., Eriksson, O., Freitas, H., Golodets, C., Kigel, J., Kleyer, M., Leps, J., Meier, T., Papadimitriou, M., Papanastasis, V.P., Quested, H., Quetier, F., Rusch, G., Sternberg, M., Theau, J.P., Thebault, A. and Vile, D. (2008) Impact of abundance on the response of seed traits to climate and land use. Journal of Ecology 96, 355366.CrossRefGoogle Scholar
Peco, B. and Espigares, T. (1994) Floristic fluctuations in annual pastures. The role of competition at the regeneration stage. Journal of Vegetation Science 5, 457462.CrossRefGoogle Scholar
Peco, B., Traba, J., Levassor, C., Sánchez, A.M. and Azcárate, F.M. (2003) Seed size, shape and persistence in dry Mediterranean grass and scrublands. Seed Science Research 13, 8795.CrossRefGoogle Scholar
Peco, B., de Pablos, I., Traba, J. and Levassor, C. (2005) The effect of grazing abandonment on species composition and functional traits: the case of dehesa grasslands. Basic and Applied Ecology 6, 175183.CrossRefGoogle Scholar
Peco, B., Lopez-Merino, L. and Alvir, M. (2006) Survival and germination of Mediterranean grassland species after simulated sheep ingestion: ecological correlates with seed traits. Acta Oecologica 30, 269275.CrossRefGoogle Scholar
Peco, B., Rico, L. and Azcárate, F.M. (2009) Seed size and response to rainfall patterns in annual grasslands: 16 years of permanent plots. Journal of Vegetation Science 20, 816.CrossRefGoogle Scholar
Retana, J., Pico, F.X. and Rodrigo, A. (2004) Dual role of harvesting ants as seed predators and dispersers of a non-myrmechorous Mediterranean perennial herb. Oikos 105, 377385.CrossRefGoogle Scholar
Rice, W.R. (1989) Analyzing tables of statistical tests. Evolution 43, 223225.CrossRefGoogle ScholarPubMed
Rivas-Martínez, S. (1987) Memoria del mapa de series de vegetación de Espańa 1:400.000. Madrid, Spain, ICONA.Google Scholar
Römermann, C., Tackenberg, O. and Poschlod, P. (2005) How to predict attachment potential of seeds to sheep and cattle coat from simple morphological seed traits. Oikos 110, 219230.CrossRefGoogle Scholar
Russi, L., Cocks, P.S. and Roberts, E.H. (1992) The fate of legume seeds eaten by sheep from a Mediterranean grassland. Journal of Applied Ecology 29, 772778.CrossRefGoogle Scholar
Sánchez, A.M., Azcárate, F., Arqueros, L. and Peco, B. (2002) Volumen y dimensiones como predictores del peso de semilla en especies herbáceas del Centro de la Península Ibérica. Anales del Jardín Botánico de Madrid 59, 249262.Google Scholar
Sánchez, A.M., Azcárate, F.M. and Peco, B. (2006) Effects of harvester ants on seed availability and dispersal of Lavandula stoechas subsp. Pedunculata in a Mediterranean grassland–scrubland mosaic. Plant Ecology 185, 4956.CrossRefGoogle Scholar
Seligman, N.G. and Vankeulen, H. (1989) Herbage production of a Mediterranean grassland in relation to soil depth, rainfall and nitrogen nutrition – a simulation study. Ecological Modelling 47, 303311.CrossRefGoogle Scholar
Sterling, A., Peco, B., Casado, M.A., Galiano, E.F. and Pineda, F.D. (1983) Influence of microtopography on floristic variation in the ecological succession in grassland. Oikos 42, 334342.CrossRefGoogle Scholar
Sternberg, M., Gutman, M., Perevolotsky, A., Ungar, E.D. and Kigel, J. (2000) Vegetation response to grazing management in a Mediterranean herbaceous community: a functional group approach. Journal of Applied Ecology 37, 224237.CrossRefGoogle Scholar
Thompson, K., Band, S.R. and Hodgson, J.G. (1993) Seed size and shape predict persistence in soil. Functional Ecology 7, 236241.CrossRefGoogle Scholar
Thornthwaite, C.W. (1948) An approach toward a rational classification of climate. Geographical Review 38, 5594.CrossRefGoogle Scholar
Veenendaal, E.M., Ernst, W.H.O. and Modise, G.S. (1996) Effect of seasonal rainfall pattern on seedling emergence and establishment of grasses in a savannah in South-Eastern Botswana. Journal of Arid Environments 32, 305317.CrossRefGoogle Scholar
Westoby, M. (1998) A leaf-height-seed (LHS) plant ecology strategy scheme. Plant and Soil 199, 213227.CrossRefGoogle Scholar
Westoby, M., Leishman, M. and Lord, J. (1996) Comparative ecology of seed size and dispersal. Philosophical Transactions: Biological Sciences 351, 13091317.Google Scholar
Willott, S.J., Compton, S.G. and Incoll, L.D. (2000a) Foraging, food selection and worker size in the seed harvesting ant Messor bouvieri. Oecologia 125, 3544.CrossRefGoogle ScholarPubMed
Willott, S.J., Miller, A.J., Incoll, L.D. and Compton, S.G. (2000b) The contribution of rabbits (Oryctolagus cuniculus L.) to soil fertility in semi-arid Spain. Biology and Fertility of Soils 31, 379384.CrossRefGoogle Scholar
Wolff, A. and Debussche, M. (1999) Ants as seed dispersers in a Mediterranean old-field succession. Oikos 84, 443452.CrossRefGoogle Scholar
Zeiter, M., Stampfli, A. and Newbery, D.M. (2006) Recruitment limitation constrains local species richness and productivity in dry grassland. Ecology 87, 942951.CrossRefGoogle ScholarPubMed