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Colonization dynamics of a clonal pioneer plant on a glacier foreland inferred from spatially explicit and size-structured matrix models

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Abstract

The regional distribution of a plant species is a result of the dynamics of extinctions and colonizations in suitable habitats, especially in strongly fragmented landscapes. Here, we studied the role of spatial dynamics of the long-lived, clonal pioneer plant Geum reptans occurring on glacier forelands in the European Alps. We used demographic data from several years and sites in the Swiss Alps in combination with dispersal data to parametrize a matrix model for G. reptans to simulate extinctions, colonizations and spatial spread of established populations on glacial forelands. We used different scenarios with varying germination rates, wind and animal dispersal capabilities, and modes of spatial spread (seed-only vs clonal spread), resulting in population growth rates (λ) ranging from 1.04 to 1.20. Our results suggest that due to the low germination rate (~1%) and the very limited wind dispersal distances (99.8% of seeds are dispersed < 5 m), G. reptans has a low probability of establishing new populations and a very low spatial spread by seed dispersal alone. In contrast to the low rate of establishment, the persistence of established populations is high and even populations of only a few individuals have an extinction probability of less than 25% within 100 years. This high persistency is partly due to clonal reproduction via aboveground stolons. Clonal reproduction increases the population size and contributes considerably to the spatial spread of established populations. Our simulation results together with the known pattern of molecular diversity of G. reptans indicate that the occurrence of populations of this species in the Alps is unlikely to be a result of recent colonizations by long-distance dispersal, but rather a result of post-glacial colonizations by large migrating populations that were fragmented when glaciers retreated. Additionally, our simulations suggest that the currently observed high rates of glacial retreat might be too fast for pioneer plants, such as G. reptans, to keep up with the retreating ice and therefore might threaten existing populations.

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Acknowledgements

We like to thank Marc Vuffray for the help with the mathematical proofs of the average dispersal distances. The computations were performed at the Vital-IT (www.vital-it.ch) Center for high-performance computing of the SIB Swiss Institute of Bioinformatics.

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Authors and Affiliations

  1. Institute of Botany, University of Basel, Schönbeinstrasse 6, 4056, Basel, Switzerland

    Daniel Scherrer & Jürg Stöcklin

  2. Department of Ecology and Evolution, University of Lausanne, Biophore, CH-1015, Lausanne, Switzerland

    Daniel Scherrer

  3. Department of Enviromental Sciences, Section of Conservation Biology, University of Basel, St. Johanns-Vorstadt 10, 4056, Basel, Switzerland

    Peter Stoll

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  1. Daniel Scherrer
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Correspondence to Jürg Stöcklin.

Electronic supplementary material

Fig. S1

Proportion of surviving (meta-)populations in a simulated valley over time under different scenarios (see Table 2 for details on scenarios). Solid lines represent the scenarios with seed-only dispersal and dashed lines the same scenarios with clonal spread included. Panels a, b and c have a glacial foreland (suitable habitat strip) of 1 km and a glacial retraction speed of 25 m, 50 m and 100 m per decade resulting in a maximum time of patch suitability of 400, 200 and 100 years, respectively. (PNG 91 kb)

Fig. S2

Median population size of (meta-)populations in a valley over time under different scenarios (see Table 2 for details on scenarios). Solid line represent the scenarios with seed-only dispersal and dashed lines the same scenarios with clonal spread included. Panels a, b and c have glacial foreland (suitable habitat strip) of 2 km and a glacial retraction speed of 50 m, 100 m and 150 m per decade, respectively. Panel (d), (e) and (f) have a glacial foreland of 1 km and a glacial retraction speed of 25 m, 50 m and 100 m per decade, respectively. The numbers in the upper right corner indicate the maximum time a habitat patch can stay suitable. (PNG 207 kb)

Fig. S3

The proportional distribution of wind dispersal, animal dispersal and clonal spread in accessing new habitat patches (grid cells) under different scenarios (see Table 2 for details on scenarios). The length of the bars indicates the median survival time of the (meta-)population in a valley. The black error lines indicate the 95% confidence interval of the survival time estimated on 1,000 simulations. Panels a, b and c have a glacial foreland (suitable habitat) of 1 km and a glacial retraction speed of 25 m, 50 m and 100 m per decade resulting in a maximum time of patch suitability of 400, 200 and 100 years, respectively. (PNG 63 kb)

Appendices

Appendix 1

Seed dispersal threshold selection

In this study we used 3.25 m as the threshold for seed dispersal outside of its cell of origin. We assumed that our plants are randomly distributed within our grid cells. Therefore, a seed starts its flight from a random position within the 5 × 5 m grid cell flying in a random direction. Depending on its starting position and direction of dispersal, the seed will need to fly between 0 and 7.07 m to leave its cell of origin. As it would take too much time/resources to model the flight of each seed individually, we decided to take the mean distance a seed needs to travel to leave its cell of origin, assuming a random distribution of plants within a grid cell and random direction of seed dispersal.

This problem can be solved mathematically by integrating

$$ {\displaystyle \begin{array}{c}\hfill 2+\sqrt{2}+5\hfill \\ {}\hfill \left(1+\sqrt{2}\right)\hfill \\ {}\hfill \left(\sqrt{x^2+{\left(y-z\right)}^2}\right) dxdydz=\frac{1}{12}+ EQ1:\underset{0}{\overset{1}{\iiint \limits }}\hfill \end{array}} $$

in the direction of x, y, and z all varying between 0 and 1. Solving this equation (EQ1) results in 0.65 × 5 m (grid size) = 3.25 m.

Appendix 2

Clonal dispersal threshold selection

To decide if a ramet produced by an above ground stolon is leaving its cell of origin, we followed a similar approach as for the seed dispersal (Appendix 1). We assumed the mother rosette to be at a random spot within the grid cell and a random direction for the growth of the stolon defining the position of the first daughter rosette. Based on these assumptions, we simulated 1,000,000 times the reproduction by clonal growth and counted the proportion of daughter rosette that were established outside of the cell of the mother rosette. Doing this for the stolon length of 0.3 m resulted in proportions of daughter rosettes leaving its mother cell of 0.022.

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Scherrer, D., Stoll, P. & Stöcklin, J. Colonization dynamics of a clonal pioneer plant on a glacier foreland inferred from spatially explicit and size-structured matrix models. Folia Geobot 52, 353–366 (2017). https://doi.org/10.1007/s12224-017-9294-z

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