Do Local Interactions or the Landscape Determine Spatial Selforganization

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1 Do Local Interactions or the Landscape Determine Spatial Selforganization in Wetland Ecosystems? Johan van de Koppel Tjeerd Bouma Peter Herman Royal Netherlands Institute for Sea Research (NIOZ) Yerseke, The Netherlands.

2

3 Spatial self-organization Spatial self-organization is the process where small-scale interactions between individuals generate structure or pattern at large spatial scales without a central authority or external element imposing it. Is it?

4 This talk Self-organization in mussel beds Diatom biofilms Salt marshes

5 What governs mussel growth?

6 Safety in numbers

7 A case study on mussel beds Mussel beds in the Wadden Sea Mussels close by Competition Facilitation Scale: ± 200 m wide Scale-dependent feedback

8 A spatial model Upper water layer Lower water layer Input Algae Mussels Uptake Model assumptions: Two water layers Lower layer can be depleted by mussels Algae limit mussel growth Lower mussel losses at higher densities

9 Self-organization in mussel banks Van de Koppel et al, AmNat 2005

10 August, 2009 Spatial ecology Norbert Dankers

11

12 September, 2009 Spatial ecology

13

14

15 Who needs biomechanics? Mussels close by Competition Facilitation Scale-dependent feedback

16 Effects of the landscape Stagnant conditions Flowing conditions

17 Self-organization and the landscape Ecological mechanism underlies pattern Scale-dependence is imposed by tides Water flow determines the shape of the pattern => Landscape determines pattern shape

18 A framework Large-scale characteristics (dynamics/patterns) interaction between physical and biological constraints Small-scale processes (mechanisms)

19 ±1 m

20

21 ±1 m

22 Diatom-sediment feedbacks Diatoms + + Silt

23

24 Modelling diatom pattern formation Diatoms increase Sediment increases Water level decreases Work of Ellen Weerman Water level increases Diatoms decrease Sediment decreases

25 Model structure & assumptions Water flows downwards: shallow water equations Low erosion when water layer is thin Scale difference: water flows on much faster time scales than diatom growth

26 Diatom biomass Sediment/water level Spatial ecology Mudflat pattern development Cross-section through the mudflat (m) Weerman et al, AmNat 2010

27 Bed level increase (cm) Spatial ecology Measuring bed elevation change Patterns Flat

28 Differences in patterns

29 The landscape again? Diatoms determine shape the interaction between water and sediment Physics is more important, even for basic understanding Landscape is again important in shaping the final pattern.

30

31 Tussock effects

32 Scale-dependent feedback Work of Bregje van Wesenbeeck Van Wesenbeeck et al. 2008

33 Explaining the landscape? 1. Small-scale plant-flowsediment feedbacks 2. Large-scale landscape patterns

34 Self-organization of salt marsh Delft3D Models: Read marsh development Work of Stijn Temmerman Temmerman et al. Geology 2007

35

36 Internal gradients Work of Qinghua YE Deltares

37 Spatial self-organization The process where local interactions generate large scale patterns Ideally! Van de Koppel et al, Science 2008

38 In the real world Local feedback processes interact with landscape forcing to determine the shape of self-organized spatial patterns.

39 Pattern theory Regular patterns found in lots of systems Local facilitation and large-scale inhibition: Scale-dependent feedback Biological-physical interactions in nearly all examples Landscape features co-shape the pattern in nearly all case studies (Van de Koppel, Bouma & Herman, J. Exp. Bio., 2012)

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