The soil seed bank of a montane meadow: consequences of conifer encroachment and implications for restoration

We examined changes in the soil seed bank associated with conifer encroachment of montane meadows in the western Cascade Range of Oregon. We asked whether, and over what period of time, meadow species maintain viable seeds in the soil, and by implication, whether the seed bank can contribute to restoration if conifers are removed. Seed bank composition, ground vegetation, and forest age structure were quantified for 209 samples representing a chronose- quence of open meadow, young forest ( 200 years). The seed bank was substantial (44 taxa and 2332 germinants/m 2 ), but dominated by native ruderals (16 species comprising 71% of germinants). Greater than 70% of meadow species were absent from the seed bank. Thirteen meadow species accounted for 21% of all germinants, but most of these were the dominant sedge, Carex pensylvanica Lam.. Seed density, richness, and composition showed weak relationships to forest age, and little resemblance to the ground vegetation, which changed markedly with forest de- velopment. Our results suggest that there is limited potential for recovery of most meadow species via the seed bank. Nat- ural reestablishment of these species will require seed dispersal or gradual vegetative spread from existing openings, but competitive interactions with ruderal or forest species may limit recruitment or recovery.

[1]  M. Zobel,et al.  Soil seed bank composition in different successional stages of a species rich wooded meadow in Laelatu, western Estonia , 1998 .

[2]  P. Milberg Soil Seed Bank after Eighteen Years of Succession from Grassland to Forest , 1995 .

[3]  J. Bakker,et al.  Soil seed banks in European grasslands: does land use affect regeneration perspectives? , 1997 .

[4]  B. Osborne,et al.  Irradiance and nitrate-dependent variation in growth and biomass allocation of Mycelis muralis. An analysis of its significance for a functional categorization of ‘sun’ and ‘shade’ plants , 1997 .

[5]  H. J. Oosting,et al.  Buried viable seeds in a successional series of old field and forest soils. , 1940 .

[6]  P. Legendre,et al.  SPECIES ASSEMBLAGES AND INDICATOR SPECIES:THE NEED FOR A FLEXIBLE ASYMMETRICAL APPROACH , 1997 .

[7]  R. Bekker,et al.  Restoration of ditch bank plant species richness: The potential of the soil seed bank , 2003 .

[8]  P. Milberg,et al.  Soil seed bank and species turnover in a limestone grassland , 1994 .

[9]  Jan Lepš,et al.  Multivariate Analysis of Ecological Data , 2006 .

[10]  O. W. Archibold CHAPTER 7 – Seed Banks and Vegetation Processes in Coniferous Forests , 1989 .

[11]  James H. Brown,et al.  Using constraint lines to characterize plant performance , 1998 .

[12]  G. Mahy,et al.  Vegetation and seed bank in a calcareous grassland restored from a Pinus forest , 2005 .

[13]  Hj Norussis,et al.  SPSS for Windows , 1993 .

[14]  Charles B. Halpern,et al.  Soil seed banks in young, closed-canopy forests of the Olympic Peninsula, Washington: potential contributions to understory reinitiation , 1999 .

[15]  A. Knapp,et al.  Woody Plant Encroachment and Removal in Mesic Grassland: Production and Composition Responses of Herbaceous Vegetation , 2005 .

[16]  A. M. Olson,et al.  SPECIES REPLACEMENT DURING EARLY SECONDARY SUCCESSION: THE ABRUPT DECLINE OF A WINTER ANNUAL , 1997 .

[17]  C. L. Knight,et al.  Expansion of gallery forest on Konza Prairie Research Natural Area, Kansas, USA , 1994, Landscape Ecology.

[18]  Constance J. Burke Historic fires in the central Western Cascades, Oregon , 1979 .

[19]  The seed rain and seed bank of an adjacent native tallgrass prairie and old field , 1997 .

[20]  A. Swanson,et al.  Conifer invasion of forest meadows transforms soil characteristics in the Pacific Northwest , 2005 .

[21]  P. Petraitis,et al.  The importance of scale in testing the origins of alternative community states , 1999 .

[22]  P. Morgan,et al.  Seed-bank contributions to regeneration of shrub species after clear-cutting and burning , 1988 .

[23]  P. Kemp CHAPTER 12 – Seed Banks and Vegetation Processes in Deserts , 1989 .

[24]  T. Dutoit,et al.  Permanent seed banks in chalk grassland under various management regimes: their role in the restoration of species-rich plant communities , 1995, Biodiversity & Conservation.

[25]  R. Boyd Indians, Fire, and the Land in the Pacific Northwest , 1999 .

[26]  J. C. Hickman Non-Forest Vegetation of the Central Western Cascade Mountains of Oregon , 1976 .

[27]  T. Jones The Restoration Gene Pool Concept: Beyond the Native Versus Non‐Native Debate , 2003 .

[28]  A. McDonald The role of seedbank and sown seeds in the restoration of an English flood‐meadow , 1993 .

[29]  M. Feller,et al.  Seed banks of forested and disturbed soils in southwestern British Columbia , 1993 .

[30]  S. Waite,et al.  The persistence of calcareous grassland species in the soil seed bank under developing and established scrub , 1998, Plant Ecology.

[31]  Julia A. Jones,et al.  Role of Light Availability and Dispersal in Exotic Plant Invasion along Roads and Streams in the H. J. Andrews Experimental Forest, Oregon , 2000 .

[32]  Mark R. T. Dale,et al.  Spatial Pattern Analysis in Plant Ecology: Spatial Pattern Analysis in Plant Ecology , 1999 .

[33]  M. Kellman The viable seed content of some forest soil in coastal British Columbia , 1970 .

[34]  K. Thompson,et al.  Distribution of buried viable seeds along a successional series , 1980 .

[35]  A. McKee,et al.  Climatic summaries and documentation for the primary meteorological station, H.J. Andrews Experimental Forest, 1972 To 1984. , 1989 .

[36]  David M. Schaepe,et al.  Historical Meadow Dynamics in Southwest British Columbia: a Multidisciplinary Analysis , 2003 .

[37]  N. Mantel The detection of disease clustering and a generalized regression approach. , 1967, Cancer research.

[38]  O. V. Auken Shrub Invasions of North American Semiarid Grasslands , 2000 .

[39]  M. Willson,et al.  Patterns of seed rain at the edge of a tropical Queensland rain forest , 1989, Journal of Tropical Ecology.

[40]  Paul A. Keddy,et al.  North American Terrestrial Vegetation , 1988 .

[41]  J. Heltshe,et al.  Estimating species richness using the jackknife procedure. , 1983, Biometrics.

[42]  B. Cade,et al.  Estimating effects of limiting factors with regression quantiles , 1999 .

[43]  V. T. Parker,et al.  Ecology of Soil Seed Banks , 1989 .

[44]  R. Marrs,et al.  A comparative study of the seedbanks of heathland and successional habitats in Dorset, Southern England , 1998 .

[45]  Henry Howe,et al.  Ecology of Seed Dispersal , 1982 .

[46]  M. Kellman Preliminary Seed Budgets for Two Plant Communities in Coastal British Columbia , 1974 .

[47]  G. Strickler,et al.  Emergent Seedlings from Coniferous Litter and Soil in Eastern Oregon , 1976 .

[48]  Mark V. H. Wilson,et al.  Fire and seedling population dynamics in western Oregon prairies , 2000 .

[49]  K. Gross A comparison of methods for estimating seed numbers in the soil , 1990 .

[50]  Charles B. Halpern,et al.  Plant Succession on Gopher Mounds in Western Cascade Meadows: Consequences for Species Diversity and Heterogeneity , 2008 .

[51]  Michael Fenner Seeds: The Ecology of Regeneration in Plant Communities , 1992 .

[52]  Charles B. Halpern,et al.  Vegetation responses to conifer encroachment in a western Cascade meadow: a chronosequence approach. , 2007 .

[53]  A. G. Valk,et al.  CHAPTER 15 – Seed Banks and the Management and Restoration of Natural Vegetation , 1989 .

[54]  C. Hitchcock,et al.  Vascular plants of the Pacific Northwest , 1962 .

[55]  R. Greenberg Biometry , 1969, The Yale Journal of Biology and Medicine.

[56]  Charles B. Halpern,et al.  Early Successional Patterns of Forest Species: Interactions of Life History Traits and Disturbance , 1989 .

[57]  J. Bakker,et al.  Soil seed bank composition along a gradient from dry alvar grassland to Juniperus shrubland , 1996 .

[58]  C. Halpern,et al.  The seed ecology of Iliamna logisepala (Torr.) Wiggins, an east Cascade endemic. , 2005 .

[59]  J. Antos,et al.  Tree invasion into a mountain‐top meadow in the Oregon Coast Range, USA , 1992 .

[60]  M. Hutchings,et al.  Estimation of the seed bank of a chalk grassland ley established on former arable land , 1988 .

[61]  Arthur Getis,et al.  Spatial Pattern Analysis , 2005 .

[62]  H. G. Baker CHAPTER 2 – Some Aspects of the Natural History of Seed Banks , 1989 .

[63]  C. T. Dyrness,et al.  Natural Vegetation of Oregon and Washington , 1988 .

[64]  P. Bierzychudek LIFE HISTORIES AND DEMOGRAPHY OF SHADE‐TOLERANT TEMPERATE FOREST HERBS: A REVIEW , 1982 .

[65]  B. McCune,et al.  Analysis of Ecological Communities , 2002 .

[66]  J. Major,et al.  Buried, viable seeds in two California bunchgrass sites and their bearing on the definition of a flora , 1966, Vegetatio.

[67]  J. Antos,et al.  Ecological Implications of Belowground Morphology of Nine Coniferous Forest Herbs , 1984, Botanical Gazette.

[68]  J. Bakker,et al.  Constraints in the restoration of ecological diversity in grassland and heathland communities. , 1999, Trends in ecology & evolution.

[69]  T. Spies,et al.  Growing‐Season Microclimatic Gradients from Clearcut Edges into Old‐Growth Douglas‐Fir Forests , 1995 .

[70]  R. Wein,et al.  Viable seed populations by soil depth and potential site recolonization after disturbance , 1977 .

[71]  J. Franklin,et al.  Seed rain and seed bank of third- and fifth-order streams on the western slope of the cascade range. Forest Service research paper , 1995 .

[72]  M. Palmer,et al.  Estimating Species Richness: The Second‐Order Jackknife Reconsidered , 1991 .

[73]  R. Livingston,et al.  Buried viable seed in successional field and forest stands, Harvard Forest, Massachusetts. , 1968 .

[74]  S. Arno,et al.  Douglas-fir encroachment into mountain grasslands in southwestern Montana. , 1986 .

[75]  P. Poschlod,et al.  Plant species richness in calcareous grasslands as affected by dispersability in space and time , 1998 .

[76]  D. H. Knight,et al.  Aims and Methods of Vegetation Ecology , 1974 .

[77]  D. W. Pratt,et al.  Buried viable seed in a ponderosa pine community , 1984 .

[78]  K. Rice CHAPTER 10 – Impacts of Seed Banks on Grassland Community Structure and Population Dynamics , 1989 .

[79]  D. Inouye,et al.  Spatial Pattern Analysis of Seed Banks: An Improved Method and Optimized Sampling , 1988 .

[80]  M. M. Moore,et al.  Soil seed banks in Pinus ponderosa forests in Arizona: Clues to site history and restoration potential , 2005 .

[81]  Charles B. Halpern,et al.  Effects of environment and grazing disturbance on tree establishment in meadows of the central Cascade Range, Oregon, USA , 1998 .

[82]  C. Braak Canonical Correspondence Analysis: A New Eigenvector Technique for Multivariate Direct Gradient Analysis , 1986 .

[83]  C. Baskin,et al.  Seeds: Ecology, Biogeography, and, Evolution of Dormancy and Germination , 1998 .

[84]  J. Miller,et al.  Distribution and Functional Roles of Rare and Uncommon Moths (Lepidoptera: Noctuidae: Plusiinae) Across a Coniferous Forest Landscape , 2003 .

[85]  M. Hermy,et al.  THE POTENTIAL OF SOIL SEEDBANKS IN THE ECOLOGICAL RESTORATION OF GRASSLAND AND HEATHLAND COMMUNITIES , 2003 .

[86]  R. H. Økland Are ordination and constrained ordination alternative or complementary strategies in general ecological studies , 1996 .

[87]  Restoration of the Cirsio dissecti-Molinietum in The Netherlands: Can we rely on soil seed banks? , 2003 .

[88]  M. Hermy,et al.  Influence of land use history on seed banks in European temperate forest ecosystems: a review , 2001 .

[89]  F. D. Johnson,et al.  Mature forest seed banks of three habitat types in central Idaho , 1987 .

[90]  J. Bakker,et al.  The Soil Seed Banks of North West Europe: Methodology, Density and Longevity , 1996 .

[91]  Douglass C. Brown Estimating the composition of a forest seed bank: a comparison of the seed extraction and seedling emergence methods , 1992 .

[92]  A. McKee,et al.  Species Composition and Diversity During Secondary Succession of Coniferous Forests in the Western Cascade Mountains of Oregon , 1988, Forest Science.

[93]  Peter Lesica,et al.  Ecological Genetics and the Restoration of Plant Communities: Mix or Match? , 1999 .

[94]  A. Grootjans,et al.  Restoration of the Cirsio dissecti-Molinietum in The Netherlands: Can we rely on soil seed banks? , 2003 .

[95]  M. Fischer,et al.  Plants with longer-lived seeds have lower local extinction rates in grassland remnants 1950–1985 , 1999, Oecologia.

[96]  Arthur Cronquist,et al.  Flora of the Pacific Northwest , 1974 .

[97]  Kenneth J. Berry,et al.  Data-dependent permutation techniques for the analysis of ecological data , 1988, Vegetatio.

[98]  M. Hill,et al.  Detrended correspondence analysis: an improved ordination technique , 1980 .

[99]  D. Peterson,et al.  Temporal and Spatial Distribution of Trees in Subalpine Meadows of Mount Rainier National Park, Washington, U.S.A , 2007 .

[100]  T. Vale Tree Invasion of Montane Meadows in Oregon , 1981 .

[101]  Mark V. H. Wilson,et al.  Buried propagules in an old-growth forest and their response to experimental disturbances , 1990 .