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Climate change and range expansion

Wim van der Putten It’s not only carbon that matters…. - Soil biota + Soil biota

Soil biota can have profound impacts on community composition Most climate change studies are focused on how may increasing temperature/drought alter local ecosystem processes Climate envelope approach predicts loss of diversity due to climate warming Lactuca serriola

Quite some move successfully to higher latitudes and altitudes, but not necessarily at the same rate Hooftman et al. Basic and Applied Ecology 2006 What about us? Lourens Baas Becking (1930’s): "Everything is everywhere, but the environment selects” Jaccard similarity index 1

Nematodes

Bacteria 0 0 100 200 Distance (km)

Chalk grasslands in southern UK: nematode and bacteria similarity decline with distance

Monroy et al. under review Primary producers Biomass, Symbionts Secondary Primary chemistry, consumers consumers structures Pollinators predators, parasitoids eaters Seed dispersers

Active dispersal

Shoot feeders Endophytes

Chewers Suckers Miners Pathogens

Passive dispersal Detritus

Root feeders Symbionts Predators, AM fungi Chemistry parasites of N-fixers root feeders Chewers Decomposers Ecto- parasites nutrients

Active dispersal Engineers Endo-parasites Soil structure pathogens

De Deyn and van der Putten (2005) TREE Multitrophic C perspective Only move: invasive potential on climate warming

Carnivore Carnivore

Herbivore Herbivore A B All move: no invasive potential

D Herbivore Plants and herbivores move: no invasive potential

Before warming After warming Van der Putten et al. Phil. Trans. 2010 Roy van Grunsven

Tragopogon dubius (range expander; until the 1950’s to north of Luxemburg) Tragopogon pratensis (native in entire range) Testing plant range expansion effects on plant-soil interactions

Van Grunsven et al. Global Change Biology 2010 Plant-soil feedback to test enemy release

Plant species A +

Good guys > Bad guys

Soil community X Soil community A - Bad guys > Good guys

drawing by Elly Morriën Original range T. dubius (●) and T. pratensis (○) New range T. dubius, original of T. pratensis

Expansion range of T. dubius

0.6 0.6

0.4 0.4

0.2 0.2

0.0 0.0

-0.2 -0.2

Effect of inoculation Effect -0.4

Effect of soil soil of biota Effect -0.4

-0.6 -0.6 43 44 45 46 47 48 49 50 52.0 52.1 52.2 Latitude Latitude

The range expander Tragopogon dubius has positive, neutral and negative soil effects in its original range, but positive soil effects in the new range

Van Grunsven et al. Global Change Biology 2010 Phylogenetically controlled experiment with exotic range expanders and related natives that co-occur in the invaded habitat Millingerwaard: exotic and related native plant species Polyphagous insects

Soil feedback

Millingerwaard: exotic and related native plant species Plant status

Exotic Origin Native

Bidens frondosa non-Eurasian Bidens cernua Bidens tripartita

Senecio inaequidens non-Eurasian viscosus

Solidago gigantea non-Eurasian virgaurea

Angelica archangelica Eurasian Angelica sylvestris

Artemisia biennis Eurasian Artemisia vulgaris

Centaurea stoebe Eurasian Centaurea cyanus Centaurea jacea

0

-0.1

Schistocerca gregaria -0.2

-0.3 herbivory effect herbivory

-0.4 *

Myzus persicae

0.1

0.0

-0.1

-0.2 Soil invaded range -0.3

plant feedback soil plant *

-0.4 Exotic plants Native plants

Elly Morriën Tim Engelkes Engelkes, Morriën et al. 2008 Nature Herbivores Herbivores Above ground Native plant Exotic plant

Below ground Herbivores, pathogens Herbivores, pathogens Decomposers, symbionts Decomposers, symbionts

Conclusion: New plants from warm climate regions have potential for invasive dominance

(N.B.: Negative and Positive effects using arrow width as strength indicator)

Testing exposure natives and exotic plants to aboveground herbivores in the field with mixed communities of native and exotic plants

3

2.5

2

1.5

1 Non-native Native 0.5

0 -2 -1 0 1 2 3 4 -0.5

-1

-1.5

Combined effect AG (g/pot) + soil AG feedbackherbivores Combined effect Sum of individual effects aboveground herbivores and soil feedback (g/pot)

In the greenhouse, belowground and aboveground effects are additive,

but 3.5 their effect strengths did3 not correlate with each other (not shown). Open question: how does this work out in the field at longer term? 2.5 2 Morriën et al. (2011) Ecology 1.5 Non-native 1 Native 0.5 0 -3 -2 -1 0 1 2 3 4 -0.5 -1

Difference of combined effects (g/pot) of effects combined Difference -1.5 -2 Sum of individual effects (g/pot) 0.9

0.7 Resistance Tolerance

0.5

0.3 Angelica sylvestris * Bidens cernua 0.1 Angelica archangelica *

-0.1 * Bidens frondosa* Artemisia biennis* -0.3 Senecio vulgaris Centaurea stoebe* Artemisia vulgaris

-0.5 Bidens tripartita

Soil feedbackfeedback(ln(own/control)) (ln(own/control)) Soil Soil

-0.7 Centaurea jacea Centaurea cyanus ?? Susceptibility -0.9 0 1000 2000 3000 4000 5000 6000 7000 Root-feeding nematodes (N/g plant root)

Nematode multiplication, plant-soil feedback and plant strategies

(* indicates exotics) Morriën et al. (2011) Oikos 0.9 Resistance Tolerance

0.7 Resistance Tolerance

0.5

0.3 Angelica sylvestris Solidago gigantea* Bidens cernua 0.1 Angelica archangelica *

-0.1 Senecio inaequidens* Bidens frondosa* Senecio viscosus Artemisia biennis* -0.3 Senecio vulgaris Centaurea stoebe* Artemisia vulgaris

-0.5 Bidens tripartita

Soil feedbackfeedback(ln(own/control)) (ln(own/control)) Soil Soil

-0.7 Centaurea jacea Centaurea cyanus Solidago virgaurea ???? SusceptibilitySusceptibility -0.9 0 1000 2000 3000 4000 5000 6000 7000 Root-feeding nematodes (N/g plant root)

Nematode multiplication, plant-soil feedback and plant strategies

(* indicates exotics) Morriën et al. (2011) Oikos 0.9 Resistance Tolerance

0.7 Resistance Tolerance

0.5

0.3 Angelica sylvestris Solidago gigantea* Bidens cernua cernua 0.1 Angelica archangelica *

-0.1 Senecio inaequidens* Bidens frondosa* frondosa* Senecio viscosus Artemisia biennis* -0.3 Senecio vulgaris Centaurea stoebe* Artemisia vulgaris

-0.5 Bidens tripartita tripartita

Soil feedbackfeedback(ln(own/control)) (ln(own/control)) Soil Soil

-0.7 Centaurea jacea Centaurea cyanus Solidago virgaurea ???? SusceptibilitySusceptibility -0.9 0 1000 2000 3000 4000 5000 6000 7000 Root-feeding nematodes (N/g plant root)

Same nematode multiplication, plant-soil feedback differs

Morriën et al. (2011) Oikos 0.9 Resistance Tolerance

0.7 Resistance Tolerance

0.5

0.3 Angelica sylvestris Solidago gigantea* Bidens cernua 0.1 Angelica archangelica *

-0.1 Senecio inaequidens* Bidens frondosa* Senecio viscosus Artemisia biennis* -0.3 Senecio vulgaris Centaurea stoebe*stoebe * Artemisia vulgaris

-0.5 Bidens tripartita

Soil feedbackfeedback(ln(own/control)) (ln(own/control)) Soil Soil

-0.7 Centaurea jaceajacea Centaurea cyanuscyanus Solidago virgaurea ???? SusceptibilitySusceptibility -0.9 0 1000 2000 3000 4000 5000 6000 7000 Root-feeding nematodes (N/g plant root)

Nematode multiplication varies within natives, plant-soil feedback

Between natives and exotic; (* indicates exotics) Morriën et al. (2011) Oikos Consequences of range expansion for ecosystem processes • General differences between exotic plants and their native congener on nutrient cycle?

• May these result in altered nutrient Annelein Meisner concentration in the plants?

Nutrient cycling

(In)organic nutrients Shoot shoot * 1.6 native * exotic * 1.2

*

0.8

0.4

0 1.6 • Shoot: higher % N for exotic

root • Root: No uniform pattern 1.2 * * * Nitrogen concentration (% N) (% concentration Nitrogen * Root 0.8

0.4

0 Bidens Centaurea Senecio Artemisia Tragopogon Native Exotic Nitrogen concentration in plant tissue Overall effects

Meisner et al. (2011) J. Ecology • Exotic plants over all had higher N and P content in their shoots, but not in their roots • When litter enhanced exotic plants, it also enhanced the performance of the phylogenetically related native • Effect on nutrient cycling was not a general phenomenon of exotics only and there was no selective advantage to exotics only

Nutrient cycling

(In)organic

nutrients Meisner et al. (2011) J. Ecology, and submitted Conclusions

• Within-continent range expanders can have invasive properties similar to cross-continental exotics • These exotic features show up aboveground, as well as belowground • Aboveground and belowground effect sizes are not necessarily correlated • Effects on ecosystem processes are not leading to a general exotic-related pattern • Increased stress incidence may enhance exotic performance, but this depends on timing and accumulation of stresses.