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The Role of Dispersal in the Assembly of Plant Communities 151 Chapter 10 PDF hosted at the Radboud Repository of the Radboud University Nijmegen The following full text is a publisher's version. For additional information about this publication click this link. http://hdl.handle.net/2066/71997 Please be advised that this information was generated on 2021-10-02 and may be subject to change. Assembly of plant communities in fragmented landscapes: The role of dispersal Wim A. Ozinga Assembly of plant communities in fragmented landscapes: The role of dispersal Een wetenschappelijke proeve op het gebied van de Natuurwetenschappen, Wiskunde en Informatica PROEFSCHRIFT ter verkrijging van de graad van doctor aan de Radboud Universiteit Nijmegen op gezag van de Rector Magnificus prof. mr. S.C.J.J. Kortmann, volgens besluit van het College van Decanen in het openbaar te verdedigen op dinsdag15 januari 2008 om 15.30 uur precies door Willem Adriaan Ozinga geboren op 16 augustus 1971 te Utrecht Promotores: Prof. dr. J.M. van Groenendael Prof. dr. J.P. Bakker (Rijksuniversiteit Groningen) Prof. dr. J.H.J. Schaminée Copromotor: Dr. ir. R.M. Bekker (Rijksuniversiteit Groningen) Manuscriptcommissie: Prof. dr. J.G.M. Roelofs Prof. dr. J.P. Grime (University of Sheffield) Prof. dr. P.F.M. Opdam (Wageningen Universiteit en Research Centre) Paranimfen: Nina A.C. Smits H.(Rik)P.J. Huiskes ISBN: 978-90-9022584-5 Lay-out and figures: Dick Visser Printed by: Van Denderen BV, Groningen © 2008 W.A. Ozinga; all rights reserved. Ozinga, W.A. (2008) Assembly of plant communities in fragmented landscapes: The role of dispersal. PhD thesis, Radboud University Nijmegen. The research presented in this PhD thesis was financially supported by the Netherlands Organisation for Scientific Research (NWO), project nr. 014.22.072. Contents Abstract 7 Introduction Chapter 1. General introduction 9 Box 1: A new tool in the search for community assembly rules: ecoinformatics 24 Box 2: Plant community types in the Netherlands within a habitat template 26 Chapter 2. Classification of dispersal traits of vascular plants 31 Dispersal limitation versus environmental constraints Chapter 3. Predictability of plant community composition from environmental conditions 53 is constrained by dispersal limitation. (Oikos 108: 555–561, 2005) Chapter 4. Local aboveground persistence of vascular plants: Life-history trade-offs and 65 environmental constraints. (Journal of Vegetation Science 18: 489–497, 2007) Chapter 5. Dispersal potential in plant communities depends on environmental conditions. 83 (Journal of Ecology 92: 767–777, 2004) Dispersal traits versus stochastic dispersal Chapter 6. Assessing the relative importance of dispersal in plant communities using an 101 ecoinformatics approach. (Folia Geobotanica 40: 53–68, 2005) Dispersal problems due to human interference Chapter 7. How important is long-distance seed dispersal by wind for regional survival of 117 plant species? (Diversity and Distributions 11: 165–172, 2005) Chapter 8. Availability of dispersal vectors as a key to plant losses in NW Europe. (submitted) 131 Box 3: Overview of changes in dispersal infrastructure in the Netherlands. 145 Synthesis and applied perspectives Chapter 9. The role of dispersal in the assembly of plant communities 151 Chapter 10. Restoration of dispersal processes in fragmented landscapes 163 References 187 Summary 213 Samenvatting 219 Dankwoord / Acknowledgment 233 Curriculum Vitae 237 Abstract Ozinga, W.A. (2008) Assembly of plant communities in fragmented landscapes: The role of dispersal. PhD thesis, Radboud University Nijmegen. The present research showed that dispersal problems in fragmented landscapes are as important for the understanding of plant diversity losses as the traditional expla- nation of deteriorating habitat quality. Efficient conservation and restoration of plant diversity requires a predictive ecol- ogy based on general principles of the assembly of plant communities (so-called ‘as- sembly rules’). Theories on the processes that shape local plant communities can be grouped into three broad views according to the main processes involved: niche- based processes, dispersal-based processes and trait-neutral, abundance-driven processes. The present research combined large databases with information on community composition and functional traits to compare the predictive power of these three views of community assembly. Although species were found to clearly sort along environmental gradients (niche-based processes), our results indicate that for most species the availability of seeds is a major limiting factor. This ‘seed limitation’ has two components: the limit- ed availability of seed sources (trait-neutral, abundance-driven processes) and the limited transport of the available seeds (dispersal-based processes). This thesis doc- uments the impact of ‘seed limitation’ across several levels of organization (species, community, landscape). We showed that differences between plant species in terms of adaptations to various modes of transport are a key factor in understanding losses of plant diversi- ty in Northwest Europe in the 20th century. Species with water- or fur-assisted dis- persal are over-represented among declining species, while species with wind- or bird-assisted dispersal are under-represented. This implies that past changes in the ‘dispersal infrastructure’ are at least as important in explaining diversity losses as the conventional explanation of environmental change. Traditional niche-based con- servation measures, although useful, are thus not enough to halt diversity losses. We propose measures to restore the ‘dispersal infrastructure’ across entire regions. Keywords: Coexistence; Community assembly; Dispersal limitation; Ecoinformatics; Habitat fragmentation; Functional traits; Land-use changes; Metacommunity; Neutral theory; Niche; Seed limitation; Species pool; Trait–environment linkages Correspondence: Wim Ozinga, Alterra, P.O. Box 47, NL-6700 AA Wageningen, The Netherlands; email: [email protected] Chapter1 General introduction Species-rich dune grassland with Early Marsh-orchid (Dactylorhiza incarnata). 10 Chapter 1 “The development of ecology as a rigorous predictive science now depends upon our success in recognising principles of wide generality and avoiding submergence in the rising flotsam of case studies, specialist observations, and untested theories.” Grime 2001 Halting the biodiversity crisis: the need for a predictive ecology The biodiversity crisis All over the world, biodiversity is declining at an alarming rate, which is unprece- dented except for a few periods of mass extinction (Lawton & May 1995, Pimm et al. 1995, Millennium Ecosystem Assessment 2005). This probably also applies to vascular plants, which are the organisms this thesis focuses on. In Europe, for ex- ample, approximately 20% of the vascular plant species are classified as globally threatened according to IUCN criteria, and this is a conservative estimate, since for many rare endemic species the threat status has not been estimated at all (Walter & Gillett 1998, Ozinga & Schaminée 2005, IUCN 2006). The main drivers of biodiversity loss are habitat loss and reduction of the quality and spatial coher- ence of the remaining habitat patches (Millennium Ecosystem Assessment 2005). Growing concern about the ongoing loss of biodiversity has resulted in in- creased efforts for the protection of endangered species and for the conservation and restoration of endangered ecosystems throughout the world (Schemske et al. 1994, United Nations 2002, Delbaere 2002, Balmford et al. 2005). Whereas na- ture conservation measures are being implemented all over the world to maintain existing biodiversity, ecological restoration is practised mainly in highly industrial- ized countries in Europe and North America (Hobbs & Norton 1996, Bakker 2005, Van Andel & Aronson 2006). Ecological restoration aims to develop impoverished human-dominated ecosystems into semi-natural systems in which natural process- es play a more prominent role (Hobbs & Norton 1996, Bakker & Berendse 1999, Van Andel & Aronson 2006). In practice, ecological restoration aims at directing ecosystems along desired successional trajectories by measures that accelerate succession or short-cut successional stages (Lockwood & Pimm 1999, Bakker et al. 2000). Ecological restoration projects in Europe and North America have mainly fo- cused on restoring abiotic conditions. Although there have been various local suc- cesses, the resulting vegetation developments have in many cases been disap- pointing in terms of plant diversity, especially as regards endangered species (Dobson et al. 1997, Bakker & Berendse 1999, Bekker & Lammerts 2002, Jansen et al. 2004). Introduction 11 Disappointing results are partly the consequence of changes in chemical, phys- ical and biological properties of the organic topsoil after long-term drainage, pol- lution or intensive fertilization, which are more difficult to reverse than was once hoped (Grootjans et al. 2002, Roelofs et al. 2002, Lucassen et al. 2005). In recent years, ecologists have therefore begun to question whether ecological restoration is actually effective for biodiversity conservation purposes, or whether it merely means replacing one degraded system by another (Dobson et al. 1997, Suding et al. 2004, Hodgson et al. 2005). Accumulating evidence suggests that even if abiotic conditions can be suffi- ciently restored, the degree to which endangered plant species re-colonize the re- stored area is often small, due to habitat fragmentation (e.g. Hutchings
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