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Functional groups are groupings of species based on physiology, morphology, life history, or other traits relevant to controls on an ecosystem process; classification will depend on the process under investigation. Although functional groups are not always clearcut, they help to elucidate mechanisms by which species influence ecosystem processes, help to generalize such mechanisms across species, and help to simplify studies in systems with a large diversity of species [P. M. Vitousek and D. U. Hooper, in Biodiversity and Ecosystem Function, E.-D. Schulze and H. A. Mooney, Eds. (Springer-Verlag, Berlin, 1993), pp. 3-14; S. E. Hobbie, D. B. Jensen, F. S. Chapin III, ibid., pp. 385-408; C. Körner, ibid., pp. 117-140].
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Functional groups are groupings of species based on physiology, morphology, life history, or other traits relevant to controls on an ecosystem process; classification will depend on the process under investigation. Although functional groups are not always clearcut, they help to elucidate mechanisms by which species influence ecosystem processes, help to generalize such mechanisms across species, and help to simplify studies in systems with a large diversity of species [P. M. Vitousek and D. U. Hooper, in Biodiversity and Ecosystem Function, E.-D. Schulze and H. A. Mooney, Eds. (Springer-Verlag, Berlin, 1993), pp. 3-14; S. E. Hobbie, D. B. Jensen, F. S. Chapin III, ibid., pp. 385-408; C. Körner, ibid., pp. 117-140].
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note
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Early season annuals: Lasthenia californica DC. ex Lindley, Microseris douglasii (DC.) Schultz-Bip., and Plantago erecta (Morris); Late season annuals: Hemizonia luzulaefolia (DC.) ssp. rudis and Lessingia micradenia E. Greene var. glabrata (Keck) Ferris; Perennial bunchgrasses: Stipa pulchra Hitchc. and Sitanion jubatum Smith; N-fixers: Lotus subpinnatus Lag. and Astragalus gambellianus Sheldon.
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2 in 1-group treatments, based on previous measurements of aboveground biomass in undisturbed serpentine grassland [S. J. McNaughton, Ecology 49, 962 (1968); S. N. Turitzin, Am. Midl. Nat. 107, 95 (1982)]. In mixture treatments, 1-group planting densities were cut to one-half, one-third, or one-fourth to maintain constant overall planting density. Because of the small stature of most serpentine species, they attained self-supporting populations within relatively small plots (9). Plots were located near the Kirby Canyon Landfill (Waste Management Inc.), on an area that was originally bare of topsoil. Serpentine topsoil was graded over the subsoil to a depth of approximately 30 cm. Treatments were planted in 1.5 m by 1.5 m plots (0.5 to 1 m buffer zone between plots) in a randomized complete block design, with 10 treatments per block and six replicates of each treatment. Treatments were planted in the winter of 1991-92, and measurements were made in the following growing season (winter/spring 1993).
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2 in 1-group treatments, based on previous measurements of aboveground biomass in undisturbed serpentine grassland [S. J. McNaughton, Ecology 49, 962 (1968); S. N. Turitzin, Am. Midl. Nat. 107, 95 (1982)]. In mixture treatments, 1-group planting densities were cut to one-half, one-third, or one-fourth to maintain constant overall planting density. Because of the small stature of most serpentine species, they attained self-supporting populations within relatively small plots (9). Plots were located near the Kirby Canyon Landfill (Waste Management Inc.), on an area that was originally bare of topsoil. Serpentine topsoil was graded over the subsoil to a depth of approximately 30 cm. Treatments were planted in 1.5 m by 1.5 m plots (0.5 to 1 m buffer zone between plots) in a randomized complete block design, with 10 treatments per block and six replicates of each treatment. Treatments were planted in the winter of 1991-92, and measurements were made in the following growing season (winter/spring 1993).
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in press
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We measured aboveground biomass by clipping annual species at the soil surface in five randomly placed 10 cm by 10 cm quadrats in each plot. For P's, we measured diameter and height of random individuals within each plot, then determined regressions for these measurements, with aboveground biomass on randomly harvested individuals [D. U. Hooper, Ecology, in press]. Clippings were sorted by species, dried at 65°C, and weighed. Total aboveground biomass is the sum of measurements made at peak biomass for each functional group: April for E's and N's, May for P's, and September for L's.
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15N, where values for the bare plot are meaningless, we used a means model coding for the ANOVA to avoid confounding the results. In these cases, ANOVA main effects and interactions were tested with balanced comparisons among the remaining treatments [For details, see D. U. Hooper and P. M. Vitousek, Ecol. Monogr., in press].
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15N, where values for the bare plot are meaningless, we used a means model coding for the ANOVA to avoid confounding the results. In these cases, ANOVA main effects and interactions were tested with balanced comparisons among the remaining treatments [For details, see D. U. Hooper and P. M. Vitousek, Ecol. Monogr., in press].
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4, filtered after 36 hours in refrigeration, and the filtrate was measured colorimetrically for ammonium and nitrate on an Alpkem RFA/2 autoanalyzer (Perstorp Analytical, Silver Spring, MD).
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thesis, Stanford University, Stanford, CA
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15N as above. See [D. U. Hooper, thesis, Stanford University, Stanford, CA (1996)] for more details.
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We thank Waste Management Inc. and the Center for Conservation Biology, Stanford University, for access to field sites. D. Turner, D. Herman, C. Chu, P. Brookes, M. Hanes, L. Jackson, M. A. Read, N. M. Holbrook, C. Benton, L. Chu, A. Cottrell, H. Farrington, M. Jones, D. Mallery, B. Tibble, M. Vandermarck, and E. Vela all provided valuable field and laboratory assistance. T. Chapin and J. Neff gave useful comments on earlier drafts of this manuscript. Financial support was provided by grants to D.U.H. from NSF (Predoctoral Fellowship and Doctoral Dissertation Improvement Grant DEB-9212995), from the Morrison Institute for Population and Resource Studies, and from the NASA/Stanford Program for Global Change. Additional support came from the Pew Scholars Program in Conservation and Environment and from the A. W. Mellon Foundation.
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