Post-invasion evolution of an invasive plant : altitudinal differenciation in germination, phenology and growth
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1 Arnaud Monty Grégory Mahy Post-invasion evolution of an invasive plant : altitudinal differenciation in germination, phenology and growth Laboratory of Ecology, Gembloux Agricultural University, Passage des Déportés 2, 5030 Gembloux, Belgium
2 1. Introduction
3 Studied taxon: Senecio inaequidens DC. Perrenial herbaceous shrub from South Africa and Lesotho Numerous yellow capitulae; Plumed wind-dispersed achenes Rapid germination, no dormancy Early flowering (1st Year), long flowering period Pioneer species, invading mainly roadsides and railways
4 Studied taxon: Senecio inaequidens DC. In the indigeneous range: diploid and tetraploid populations In the introduction range: tetraploids only Introduction history: Wool alien at the end of the 19th century Verviers (Belgium) : 1892 Bremen (Germany) : 1896 Mazamet (France) : 1936 Verona (Italy) : >1940 Lag time: populations only found in wool industry areas : Rapid spread through western Europe
5 Senecio inaequidens DC.: an appropriate model for evolutionary studies! Invasion history precisely known Independant introduction spots (link between all populations of a country) Spread, from these spots, through climate contrasted areas Climate = potentially the main selection agent : Few natural ennemies altering fitness Little variability in invaded soils Anemochorous dispersion Little competition (pioneer species) Generalist pollination system Appropriate model for evolutionary studies considering climate as a selection agent
6 Invasion process: two main evolutionary phases Primary evolution phase: (More studied) Introduction Naturalisation Primary invasion Secondary evolution phase: (Less studied) Primary invasion Secondary invasion (contrasted climatic conditions)
7 Objectives of the study: Using common garden experiment to assess the species differenciation : 1) Between cytotypes 2) Between continents (Primary evolution phase) 3) Along climatic gradients in Europe (Secondary evolution phase) considering 1) Germination 2) Flowering phenology 3) Plant size
8 2. Material and methods
9 Seed collection: South Africa: Tetraploid «control» zone 2 populations x 10 individuals Diploid zone 2 populations x 10 individuals [Lafuma, 2003]
10 Seed collection: South Africa: Tetraploid «control» zone 2 populations x 10 individuals Diploid zone 2 populations x 10 individuals Europe: 2 altitudinal/climatic transects
11 Seed collection: South Africa: Tetraploid «control» zone 2 populations x 10 individuals Topography: m Diploid zone 2 populations x 10 individuals Europe: 2 altitudinal/climatic transects France: 5 zones x 2 populations x 10 individuals 100 km
12 Seed collection: South Africa: Tetraploid «control» zone 2 populations x 10 individuals Diploid zone Topography: m 2 populations x 10 individuals Europe: 2 altitudinal/climatic transects France: 5 zones x 2 populations x 10 individuals Belgium: 4 zones x 2 populations x 10 individuals
13 Seed collection: South Africa: Tetraploid «control» zone 2 populations x 10 individuals Diploid zone 2 populations x 10 individuals Europe: 2 altitudinal/climatic transects France: 5 zones x 2 populations x 10 individuals Belgium: 4 zones x 2 populations x 10 individuals Total : 240 parent individuals
14 Elimination of maternal effects: 10 biggest achenes without anomaly Parent individual Capitulae collected Sorting of achenes
15 Sowing and measurements: 10 biggest achenes without anomaly Sown in one pot 1 plant was kept (6 months) One plant per parent individual
16 Sowing and measurements: 10 bigger achenes without anomaly Sown in one pot 1 plant was kept (6 months) One plant per parent individual Germination study (n = 2400) Phenology and growth study (n = 240)
17 Sowing and measurements: 10 bigger achenes without anomaly Sown in one pot 1 plant was kept (6 months) One plant per parent individual Germination delay (2 days) Germination window (2 days) Germination rate (2 days) Flowering delay (since germination) (everyday) Flowering delay (since sowing) (everyday) Plant height (every 25 days) Plant volume (every 25 days) 4 measurements
18 Common garden experiment: randomized bloc design
19 Statistical analysis: Comparison: 3 way - ANOVAs
20 Statistical analysis: Comparison: Between cytotypes 3 way - ANOVAs
21 Statistical analysis: Comparison: Between cytotypes 3 way - ANOVAs Between African control and introduction areas (Verviers & Mazamet) (Primary evolution phase)
22 Statistical analysis: Comparison: Between cytotypes 3 way - ANOVAs Between African control and introduction areas (Verviers & Mazamet) (Primary evolution phase) Between zones of each gradient (Secondary evolution phase)
23 Statistical analysis: Comparison: Between cytotypes 3 way - ANOVAs Between African control and introduction areas (Verviers & Mazamet) (Primary evolution phase) Between zones of each gradient (Secondary evolution phase) Correlation analysis: (Secondary evolution phase) Pearson s r (linear), Spearman s R (non-parametric) For each gradients: Population means vs Altitude Population means vs gradient zones
24 3. Results
25 Results presentation: For each trait: Comparison (ANOVA) N p value Cytotypes 60 p Primary evolution phase 80 p Zones of French transect 100 p Zones of Belgian transect 80 p Correlation with altitude ( p value) r Pearson R Spearman p p p p Correlation with transect zones ( p value) r Pearson R Spearman p p p p Significant result
26 Germination Germination delay Comparison (ANOVA) N p value Cytotypes 60 0,913 Primary evolution phase 80 0,445 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,313 0,165 0,046 0,059 0,086 Zones of Belgian transect 80 0,179 0,169 0,091 0,166 0,073 Germination window Comparison (ANOVA) N p value Cytotypes 60 0,913 Germination rate Correlation with altitude Correlation with transect zones Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,889 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,897 0,591 0,476 0,5 0,561 Zones of Belgian transect 80 0,758 0,486 0,736 0,82 0,818 Comparison (ANOVA) N p value Cytotypes 60 0,348 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,611 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,213 0,532 0,7 0,683 0,683 Zones of Belgian transect 80 0,65 0,148 0,126 0,177 0,145
27 Germination Délai de première germination (j) Germination delay Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,913 Primary evolution phase 80 0,445 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,313 0,165 0,046 0,059 0,086 Zones of Belgian transect 80 0,179 0,169 0,091 0,166 0, French transect Non-linear increase with altitude Altitude (m)
28 Germination Germination delay Comparison (ANOVA) N p value Cytotypes 60 0,913 Primary evolution phase 80 0,445 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,313 0,165 0,046 0,059 0,086 Zones of Belgian transect 80 0,179 0,169 0,091 0,166 0,073 Germination window Comparison (ANOVA) N p value Cytotypes 60 0,913 Germination rate Correlation with altitude Correlation with transect zones Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,889 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,897 0,591 0,476 0,5 0,561 Zones of Belgian transect 80 0,758 0,486 0,736 0,82 0,818 Comparison (ANOVA) N p value Cytotypes 60 0,348 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,611 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,213 0,532 0,7 0,683 0,683 Zones of Belgian transect 80 0,65 0,148 0,126 0,177 0,145
29 Flowering phenology Flowering delay (since germination) Comparison (ANOVA) N p value Cytotypes 60 0,15 ( p value) ( p value) Primary evolution phase 80 0,662 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,226 0,04 0,08 0,019 0,051 Zones of Belgian transect 80 0,716 0,17 0,57 0,218 0,643 Flowering delay (since sowing) Correlation with altitude Correlation with transect zones Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,061 Primary evolution phase 80 0,657 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,589 0,123 0,192 0,116 0,125 Zones of Belgian transect 80 0,285 0,227 0,217 0,156 0,217
30 Flowering phenology Délai de floraison depuis germination (j) Flowering delay (since germination) Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,15 ( p value) ( p value) Primary evolution phase 80 0,662 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,226 0,04 0,08 0,019 0,051 Zones of Belgian transect 80 0,716 0,17 0,57 0,218 0, French transect Linear decrease with altitude and zones Altitude (m)
31 Flowering phenology Flowering delay (since germination) Comparison (ANOVA) N p value Cytotypes 60 0,15 ( p value) ( p value) Primary evolution phase 80 0,662 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,226 0,04 0,08 0,019 0,051 Zones of Belgian transect 80 0,716 0,17 0,57 0,218 0,643 Flowering delay (since sowing) Correlation with altitude Correlation with transect zones Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,061 Primary evolution phase 80 0,657 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,589 0,123 0,192 0,116 0,125 Zones of Belgian transect 80 0,285 0,227 0,217 0,156 0,217
32 Plant height Height (date 1) Comparison (ANOVA) N p value Cytotypes 60 0,527 Primary evolution phase 80 0,46 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,128 0,13 0,008 0,028 0,01 Zones of Belgian transect 80 0,171 0,231 0,289 0,333 0,482 Height (date 2) Height (date 3) Height (date 4) Correlation with altitude Correlation with transect zones Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,439 Primary evolution phase 80 0,077 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,172 0,321 0,041 0,084 0,044 Zones of Belgian transect 80 0,04 0,096 0,07 0,078 0,062 Comparison (ANOVA) N p value Cytotypes 60 0,493 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,622 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,01 0,006 0,003 0,003 0,003 Zones of Belgian transect 80 0,562 0, ,897 1 Comparison (ANOVA) N p value Cytotypes 60 0,443 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,901 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,058 0,027 0,039 0,004 0,017 Zones of Belgian transect 80 0,394 0,097 0,071 0,076 0,062
33 Plant height Height (date 1) Comparison (ANOVA) N p value Cytotypes 60 0,527 Primary evolution phase 80 0,46 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,128 0,13 0,008 0,028 0,01 Zones of Belgian transect 80 0,171 0,231 0,289 0,333 0,482 Height (date 2) Height (date 3) Height (date 4) Correlation with altitude Correlation with transect zones Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,439 Primary evolution phase 80 0,077 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,172 0,321 0,041 0,084 0,044 Zones of Belgian transect 80 0,04 0,096 0,07 0,078 0,062 Comparison (ANOVA) N p value Cytotypes 60 0,493 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,622 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,01 0,006 0,003 0,003 0,003 Zones of Belgian transect 80 0,562 0, ,897 1 Comparison (ANOVA) N p value Cytotypes 60 0,443 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,901 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,058 0,027 0,039 0,004 0,017 Zones of Belgian transect 80 0,394 0,097 0,071 0,076 0,062
34 Plant height Height (date 1) Comparison (ANOVA) N p value Cytotypes 60 0,527 Primary evolution phase 80 0,46 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,128 0,13 0,008 0,028 0,01 Zones of Belgian transect 80 0,171 0,231 0,289 0,333 0,482 Height (date 2) Height (date 3) Height (date 4) Correlation with altitude Correlation with transect zones Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,439 Primary evolution phase 80 0,077 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,172 0,321 0,041 0,084 0,044 Zones of Belgian transect 80 0,04 0,096 0,07 0,078 0,062 Comparison (ANOVA) N p value Cytotypes 60 0,493 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,622 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,01 0,006 0,003 0,003 0,003 Zones of Belgian transect 80 0,562 0, ,897 1 Comparison (ANOVA) N p value Cytotypes 60 0,443 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,901 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,058 0,027 0,039 0,004 0,017 Zones of Belgian transect 80 0,394 0,097 0,071 0,076 0,062
35 Plant height (cm) Hauteur du plant (cm) Plant height 80 French transect Decrease with altitude and zones for each measurement date mai 06-juin 03-juil 26-juil Transect Zone climatique zones
36 Plant height (cm) Hauteur du plant (cm) Plant height 80 French transect Decrease with altitude and zones for each measurement date mai 06-juin 03-juil 26-juil Transect Zone climatique zones
37 Plant volume Volume (date 1) Comparison (ANOVA) N p value Cytotypes 60 0,345 ( p value) Primary evolution phase 80 0,579 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,055 0,249 0,098 0,05 0,106 Zones of Belgian transect 80 0,648 0,343 0,736 0,755 1 Volume (date 2) Volume (date 3) Volume (date 4) Correlation with altitude ( p value) Correlation with transect zones Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,008 Primary evolution phase 80 0,024 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,242 0,205 0,074 0,083 0,106 Zones of Belgian transect 80 0,058 0,648 0,61 0,676 0,729 Comparison (ANOVA) N p value Cytotypes 60 0,084 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,178 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,039 0,014 0,001 0,001 0,001 Zones of Belgian transect 80 0,836 0,614 0,867 0,82 0,909 Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,379 Primary evolution phase 80 0,233 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,001 0,016 0, Zones of Belgian transect 80 0,537 0,131 0,233 0,157 0,22
38 Plant volume Volume (date 1) Comparison (ANOVA) N p value Cytotypes 60 0,345 ( p value) Primary evolution phase 80 0,579 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,055 0,249 0,098 0,05 0,106 Zones of Belgian transect 80 0,648 0,343 0,736 0,755 1 Volume (date 2) Volume (date 3) Volume (date 4) Correlation with altitude ( p value) Correlation with transect zones Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,008 Primary evolution phase 80 0,024 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,242 0,205 0,074 0,083 0,106 Zones of Belgian transect 80 0,058 0,648 0,61 0,676 0,729 Comparison (ANOVA) N p value Cytotypes 60 0,084 Correlation with altitude Correlation with transect zones Primary evolution phase 80 0,178 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,039 0,014 0,001 0,001 0,001 Zones of Belgian transect 80 0,836 0,614 0,867 0,82 0,909 Comparison (ANOVA) N p value Correlation with altitude Correlation with transect zones Cytotypes 60 0,379 Primary evolution phase 80 0,233 r Pearson R Spearman r Pearson R Spearman Zones of French transect 100 0,001 0,016 0, Zones of Belgian transect 80 0,537 0,131 0,233 0,157 0,22
39 Plant Volume volume du plant (cm³) Plant volume French transect Linear decrease with altitude and zones mai 06-juin 03-juil 26-juil Zone climatique Transect zones
40 Plant Volume volume du plant (cm³) Plant volume French transect Linear decrease with altitude and zones mai 06-juin 03-juil 26-juil Zone climatique Transect zones
41 4. Discussion
42 Discussion: differences between cytotypes Almost no differences between cytotypes These results cannot explain the absence of diploids in Europe
43 Discussion: primary evolution phase Almost no differences between (a) African tetraploid control populations and (b) Introduction areas populations These results don t support the hypothesis of a primary evolution phase Phenotypic plasticity? further data is needed (about reproductive success, winter survival, biomass, ) to fully characterise the primary invasion. NB: More African control populations could be suitable
44 Discussion: secondary evolution phase Almost no differences along belgian transect But clear differenciation along the French transect (more contrasted)! Secondary evolution phase (rapid evolution) correlated with altitude In France, plants originating from high elevations: - Germinate later : already shown for non-alien species - Bloom at a lower age : adaptation to shorter vegetation period? - Remain smaller : harsher conditions, difference in trade-off between reproduction and growth?
45 5. Conclusion and perspectives
46 Conclusion and perspectives Difference between cytotypes Primary evolution phase Further data needed! Clinal evolution along altitudinal and climatic transect: Rapid evolution phenomenon in a secondary evolution phase In the future: Finer analysis: correlation with direct meteorological data (real selection agent) Reciprocal transplants along the french gradient Understanding the adaptative benefit of the evolution Collaboration needed.?!
47 Thank you for your attention!
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