Folia Geobotanica

, Volume 40, Issue 1, pp 13–34 | Cite as

Diaspore and gap availability are limiting species richness in wet meadows



Species pool theory claims that diaspore and microsite availability limit species richness in plant communities. Wet meadows (Calthion) and litter meadows (Molinion, Caricion davallianae) belonging to the most species-rich meadows in the foothills of the Alps have suffered from a strong decrease since the 1970s. Restoration efforts including nutrient impoverishment and rewetting management after drainage and fertilization did not result in the re-establishment of the former species richness although the abiotic filter would have been appropriate for the re-colonization of many locally extinct species. In our experiment at four sites in the largest fen of Southwest-Germany we tested if the restoration success was seed- and gap-limited. We applied sowing and hay spreading (hay seed) as treatments to make seeds available and harrowing to increase gap availability. Sowing seeds or hay seed of species of the former meadow types increased species richness immediately. The proportion of re-established species was higher when additional harrowing was applied. Species richness could be increased not only in vascular plants but also in bryophytes when hay spreading was applied. The strongest re-development towards the target communities (defined through the abiotic filter and the species richness before drainage and fertilization) took place on those sites where hay spreading and harrowing were applied. Sowing seeds and hay seed were traditional techniques to establish e.g. litter meadows, both techniques have been applied for centuries. Even harrowing was described as early as the 19th century to increase the chance of establishing certain species. Therefore, the “application of the knowledge coming from the species pool theory” (although not named during this time) has been common practice since at least the 19th century.


Bryophytes Diaspore availability Dispersal Gap Microsite availability Species pool concept Vascular plants 


Frahm & Frey (1988) Oberdorfer (1983) 

Electronic appendix (

Appendix 1 Appendix 2 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Abt K. (1991): Landschaftsökologische Auswirkungen des Agrarstrukturwandels im Württembergischen Allgäu.Schriftenreihe Stud. Agrarökol. 1: 1–151.Google Scholar
  2. Bakker J.P., Poschlod P., Strykstra R.J., Bekker R.M. &Thompson K. (1996): Seed banks and seed dispersal: important topics in restoration ecology.Acta Bot. Neerl. 45: 461–490.Google Scholar
  3. Bazzaz F.A. (1996):Plants in changing environments. Linking physiological, population, and community ecology. Cambridge Univ. Press, Cambridge.Google Scholar
  4. Biewer H. (1997): Regeneration artenreicher Feuchtwiesen. In:Biewer H. & Poschlod P. (eds.), Regeneration artenreicher Feuchtwiesen im Federseeried,Veröff. Projekt Angew. Ökol. 24: 3–323.Google Scholar
  5. Bonn S. &Poschlod P. (1998):Ausbreitungsbiologie der Pflanzen Mitteleuropas. Quelle & Meyer, Wiesbaden.Google Scholar
  6. Brown R.L. &Fridley J.D. (2003): Control of plant species diversity and community invasibility by species immigration: seed richness versus seed density.Oikos 102: 15–24.CrossRefGoogle Scholar
  7. Bullock J.M. (2000): Gaps and seedling colonization. In:Fenner M. (ed.),Seeds, The ecology of regeneration in plant communities, CABI Publishing, Oxon, New York, pp. 375–395.Google Scholar
  8. Burke M.J.W. &Grime J.P. (1996): A experimental study on plant community invasibility.Ecology 77: 776–790.CrossRefGoogle Scholar
  9. Cain M.L., Milligan B.G. &Strand A.E. (2000): Long-distance seed dispersal in plant populations.Amer. J. Bot. 87: 1217–1227.CrossRefGoogle Scholar
  10. Clark J.S., Fastie C., Hurtt G., Jackson S.T., Johnson C., King G.A., Lewis M., Lynch J., Pacala S., Prentice C., Schupp E.W., Webb J. III &Wyckoff P. (1998): Reid’s paradox of rapid plant migration.BioScience 48: 13–24.CrossRefGoogle Scholar
  11. Eriksson A. (1998): Regional distribution ofThymus serpyllum: management history and dispersal limitation.Ecography 21: 35–43.CrossRefGoogle Scholar
  12. Eriksson O. (1993): The species-pool hypothesis and plant community diversity.Oikos 68: 371–374.CrossRefGoogle Scholar
  13. Eriksson O. (1997): Colonization dynamics and relative abundance of three plant species (Antennaria dioica, Hieracium pilosella andHypochoeris maculata) in dry semi-natural grasslands.Ecography 20: 559–568.CrossRefGoogle Scholar
  14. Foster B.L. (2001): Constraints on colonization and species richness along a productivity gradient: the role of propagule availability.Ecol. Letters 4: 530–535.CrossRefGoogle Scholar
  15. Fowler N. (1988): What is a safe site? Neighbour, litter, germination date, and patch effects.Ecology 69: 947–961.CrossRefGoogle Scholar
  16. Frahm J.-P. &Frey W. (1988):Moosflora. Ulmer, Stuttgart.Google Scholar
  17. Gibson C.W.D., Watt T.A. &Brown V.K. (1987): The use of sheep grazing to recreate species-rich grasslands from abandoned arable land.Biol. Conservation 42: 165–183.CrossRefGoogle Scholar
  18. Grime J.P., Hodgson J.G. &Hunt R. (1988):Comparative plant ecology. A functional approach to British species. Unwin Hyman, London.Google Scholar
  19. Häfener F. (1847):Der Wiesenbau in seinem ganzen Umfange nebst Anleitung zum Nivellieren, zur Erbauung von Schleußen, Wehren, Brücken usw. Reutlingen, Leipzig.Google Scholar
  20. Harvey H.J. &Meredith T.C. (1981): Ecological studies ofPeucedanum palustre and their implications for conservation management at Wicken Fen, Chambridgeshire. In:Synge H. (ed.),The biological aspects of rare plant conservation, Wiley, Chichester, pp. 365–378.Google Scholar
  21. Hitchmough J.D. (2003): Effects of sward height, gap size, and slug grazing on emergence and establishment ofTrollius europaeus (globeflower).Restoration Ecol. 11: 20–28.CrossRefGoogle Scholar
  22. Hulme P.E. (1994): Seedling herbivory in grassland: relative impact of vertebrate and invertebrate herbivores.J. Ecol. 82: 873–880.CrossRefGoogle Scholar
  23. Huston M.A. (1999): Local processes and regional patterns: appropriate scales for understanding variation in the diversity of plants and animals.Oikos 86: 393–401.CrossRefGoogle Scholar
  24. Jacquemyn H., Butaye J., Dumortier M., Hermy M. &Lust N. (2001a): Effects of age and distance on plant species composition of mixed deciduous forest fragments in an agricultural landscape.J. Veg. Sci. 12: 635–642.CrossRefGoogle Scholar
  25. Jacquemyn H., Butaye J. &Hermy M. (2001b): Forest plant species richness in small, fragmented mixed deciduous forest patches: the role of area, time and dispersal limitation.J. Biogeogr. 28: 801–812.CrossRefGoogle Scholar
  26. Jensen K. &Schrautzer J. (1999): Consequences of abandonment for a regional fen flora and mechanisms of successional change.Appl. Veg. Sci. 2: 79–88.CrossRefGoogle Scholar
  27. Kahmen S., Poschlod P. &Schreiber K.-F. (2002): Conservation management of calcareous grasslands. Changes in plant species composition and response of functional traits during 25 years.Biol. Conservation 104: 319–328.CrossRefGoogle Scholar
  28. Kahmen S. &Poschlod P. (2004): Plant functional trait responses to grassland succession over 25 years.J. Veg. Sci. 15: 21–32.CrossRefGoogle Scholar
  29. Kalamees R. &Zobel M. (2002): The role of the seed bank in gap regeneration in a calcareous grassland community.Ecology 83: 1017–1025.Google Scholar
  30. Kapfer A. (1988): Versuche zur Renaturierung gedüngten Feuchtgrünlandes — Aushagerung und Vegetationsentwicklung.Diss. Bot. 120: 1–144.Google Scholar
  31. Kapfer A. (1994): Erfolgskontrolle bei Renaturierungsmaßnahmen im Feuchtgrünland.Schriftenreihe Landschaftspflege Naturschutz 40: 125–142.Google Scholar
  32. Keddy P.A. (1992) Assembly and response rules: two goals for predictive community ecology.J. Veg. Sci. 3: 157–167.CrossRefGoogle Scholar
  33. Konold W. (1991): Wasser, Wiesen und Wiesenwässerung im Allgäu. Ein Beitrag zur Agrar- und Stadtgeschichte.Schriftenreihe Verein Gesch. Bodensees & Umgebung 109: 161–213.Google Scholar
  34. Konold W. &Popp S. (1994): Zur Geschichte der Wiesenwässerung im Bereich der Württembergischen Donau. In:Konold W. (Hrsg.),Historische Wasserwirtschaft im Alpenraum und an der Donau, Wittwer, Stuttgart, pp. 377–398.Google Scholar
  35. Korneck D., Schnittler M., Klingenstein F., Gerhard L., Takla M., Bohn U. &May R. (1998): Warum verarmt unsere Flora? Auswertung der Roten Liste der Farn- und Blütenpflanzen Deutschlands.Schriftenreihe Vegetationsk. 29: 299–359.Google Scholar
  36. Kotorová I. &Lepš J. (1999): Comparative ecology of seedling recruitment in an oligotrophic wet meadow.J. Veg. Sci. 10: 175–186.CrossRefGoogle Scholar
  37. Kupferschmid A.D., Stampfli A. &Newbery D.M. (2000): Dispersal and microsite limitation in an abandoned calcareous grassland of the southern Prealps.Folia Geobot. 35: 125–141.CrossRefGoogle Scholar
  38. Mouquet N., Leadley P., Meriguet J. &Loreau M. (2004): Immigration and local competition in herbaceous plant communities: a three year seed-sowing experiment.Oikos 104: 77–90.CrossRefGoogle Scholar
  39. Müller M. the older called Hohentwieler (1752):Gründlicher Bericht, wie aus des Erdbodens Beschaffenheit vorlängstens unweit Ulm, zwischen Grimmelfingen und Gögglingen, in dem sogenannten Tauben Riedt, dass unfehlbar Turf oder Torf vorhanden seyn müssen, beurtheilet .... Ulm.Google Scholar
  40. Oberdorfer E. (1983):Pflanzensoziologische Exkursionsflora. Ulmer, Stuttgart.Google Scholar
  41. Overbeck G., Kiehl K. &Abs C. (2003): Seedling recruitment ofSuccisella inflexa in fen meadows: Importance of seed and microsite availability.Appl. Veg. Sci. 6: 97–104.CrossRefGoogle Scholar
  42. Pärtel M., Zobel M., Liira J. &Zobel K. (2000): Species richness limitations in productive and oligotrophic communities.Oikos 90: 191–193.CrossRefGoogle Scholar
  43. Pärtel M., Zobel M., Zobel K. &van der Maarel E. (1996): The species pool and its relation to species richness: evidence from Estonian plant communities.Oikos 75: 111–117.CrossRefGoogle Scholar
  44. Patzelt A. &Pfadenhauer J. (1998): Keimungsbiologie und Etablierung von Niedermoor-Arten bei Ansaat durch Mähgutübertragung.Z. Ökol. Naturschutz 7: 1–13.Google Scholar
  45. Pfadenhauer J. &Maas D. (1987): Samenpotential in Niedermoorböden des Alpenvorlandes bei Grünlandnutzung unterschiedlicher Intensität.Flora 179: 85–97.Google Scholar
  46. Pfadenhauer J. &Grootjans A.P. (1999): Wetland restoration in Central Europe: aims and methods.Appl. Veg. Sci. 2: 95–102.CrossRefGoogle Scholar
  47. Poschlod P. &Bloch D. (1998): Verbreitung, Nutzungsgeschichte und heutige Situation der Moore in Baden-Württemberg. In:Weisser H. &Renner F. (eds.),Zehn Jahre Projekt Wurzacher Ried, Margraf, Weikersheim, pp. 173–188.Google Scholar
  48. Poschlod P. &Bonn S. (1998): Changing dispersal processes in the central European landscape since the last ice age — an explanation for the actual decrease of plant species richness in different habitats.Acta Bot. Neerl. 47: 27–44.Google Scholar
  49. Poschlod P., Kleyer M., Jackel A.-K., Dannemann A. &Tackenberg O. (2003): BIOPOP — A database of plant traits and internet application for nature conservation.Folia Geobot. 38: 263–271.CrossRefGoogle Scholar
  50. Poschlod P., Kiefer S., Tränkle U., Fischer S. &Bonn S. (1998): Plant species richness in calcareous grasslands as affected by dispersability in space and time.Appl. Veg. Sci. 1: 75–90.CrossRefGoogle Scholar
  51. Poschlod P., Tackenberg O. &Bonn S. (2004): Plant dispersal potential and its relation to species frequency and coexistence In:van der Maarel E. (ed.),Vegetation ecology, Oxford, Blackwells, pp. 147–171.Google Scholar
  52. Reynolds H.L., Packer A., Bever J.D. &Clay K. (2003): Grassroots ecology: plant-microbe-soil interactions as drivers of plant community structure and dynamics.Ecology 84: 2281–2291.CrossRefGoogle Scholar
  53. Ryser P. (1993): Influence of neighbouring plants on seedling establishment in limestone grassland.J. Veg. Sci. 4: 195–202.CrossRefGoogle Scholar
  54. Schmidt W. (1974): Bericht über die Arbeitsgruppe für Sukzessionsforschung auf Dauerflächen der Internationalen Vereinigung für Vegetationskunde.Vegetatio 29: 69–73.CrossRefGoogle Scholar
  55. Schütz W. (2000): The importance of seed regeneration strategies for the persistence of species in the changing landscape of central Europe.Z. Ökol. Naturschutz 9: 73–83.Google Scholar
  56. Silvertown J.W. &Lovett Doust J. (1993):Introduction to plant population biology. Blackwell, Oxford.Google Scholar
  57. SPSSInc. (1991):SPSS statistical algorithms. Ed. 2. SPSS Inc., Chicago.Google Scholar
  58. Stampfli A. &Zeitler M. (1999): Plant species decline due to abandonment of meadows cannot easily reversed by mowing. A case study from the southern Alps.J. Veg. Sci. 10: 151–164.CrossRefGoogle Scholar
  59. Stebler F.G. (1898):Die besten Streuepflanzen. IV. Teil des schweizerischen Wiesenpflanzenwerkes. Wyß, Bern.Google Scholar
  60. Stender S., Poschlod P., Vauk-Henzelt E. &Dernedde T. (1997): Die Ausbreitung von Pflanzen durch Galloway-Rinder.Verh. Ges. Ökol. 27: 173–180.Google Scholar
  61. Strykstra R.J., Bekker R.M &Verweij G.L. (1996): Establishment ofRhinanthus angustifolius in a successional hayfield after seed dispersal by mowing machinery.Acta Bot. Neerl. 45: 557–562.Google Scholar
  62. Ter Heerdt G.J.N., Verweij G.L., Bekker R.M. &Bakker J.P. (1996): An improved method for seed-bank analysis: seedling emergence after removing the soil by sieving.Funct. Ecol. 10: 144–151.CrossRefGoogle Scholar
  63. Thompson K. &Grime J.P. (1983): A comparative study of germination responses to diurnally-fluctuating temperatures.J. Appl. Ecol. 20: 141–156.CrossRefGoogle Scholar
  64. Tilman D. (1982):Resource competition and community structure. Monographs in Population Biology 17. Princeton University Press, Princeton, NY.Google Scholar
  65. Tilman D. (1988):Dynamics and structure of plant communities. Monographs in Population Biology 26. Princeton University Press, Princeton, NY.Google Scholar
  66. Tilman D. (1997): Community invasibility, recruitment limitation, and grassland biodiversity.Ecology 78: 81–92.CrossRefGoogle Scholar
  67. Titus J.H. &Lepš J. (2000): The response of arbuscular mycorrhizae to fertilization, mowing and removal of dominant species in a diverse oligotrophic wet meadow.Amer. J. Bot. 87: 392–401.CrossRefGoogle Scholar
  68. van der Maarel E. &Sykes M. T. (1993): Small-scale plant species turnover in a limestone grassland: the carousel model and some comments on the niche concept.J. Veg. Sci. 4: 179–188.CrossRefGoogle Scholar
  69. Veit R. (1849):Anleitung zum Futterbau, mit einem Anhang über Streugewinnung. München.Google Scholar
  70. Vogt K., Rasran L. &Jensen K. (2004): Water-borne seed transport and seed deposition during flooding in a small river-valley in Northern Germany.Flora 199: 377–388.Google Scholar
  71. Wilby A. &Brown V.K. (2001): Herbivory, litter and soil disturbance as determinants of vegetation dynamics during early old-field succession under set-aside.Oecologia 127: 259–265.CrossRefGoogle Scholar
  72. Wildi O. (1986): Analyse vegetationskundlicher Daten. Theorie und Einsatz statistischer Methoden.Veröff. Geobot. Inst. ETH Stiftung Rübel, Zürich 90: 1–226.Google Scholar
  73. Willems J. (2001): Problems, approaches and results in restoration of Dutch calcareous grasslands during the last 30 years.Restoration Ecol. 9: 147–154.CrossRefGoogle Scholar
  74. Zobel M. (1997): The relative role of species pools in determining plant species richness: an alternative explanation of species coexistence.Trends Ecol. Evol. 12: 266–269.CrossRefGoogle Scholar
  75. Zobel M. &Kalamees R. (2005): Diversity and dispersal — can the link be approached experimentally?Folia Geobot. 40: 3–11.CrossRefGoogle Scholar
  76. Zobel M., Moora M. &Haukioja E. (1997): Plant coexistence in the interactive environment: arbuscular mycorrhizae should not be out of mind.Oikos 78: 202–206.CrossRefGoogle Scholar
  77. Zobel M., Otsus M., Liira J., Moora M. &Möls T. (2000): Is small-scale species richness limited by seed availability or microsite availability?Ecology 81: 3274–3282.Google Scholar
  78. Zobel M., van der Maarel E. &Dupré C. (1998): Species pool: the concept, its determination and significance for community restoration.Appl. Veg. Sci. 1: 55–66.CrossRefGoogle Scholar

Copyright information

© Institute of Botany 2005

Authors and Affiliations

  1. 1.Institute of Botany, Faculty of BiologyUniversity of RegensburgRegensburgGermany

Personalised recommendations