A general, community-level test of the Janzen-Connell hypothesis C. - - PowerPoint PPT Presentation

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A general, community-level test of the Janzen-Connell hypothesis C. - - PowerPoint PPT Presentation

A general, community-level test of the Janzen-Connell hypothesis C. E. Timothy Paine 1 , Natalia Norden 2 , Christopher Baraloto 3 , Jrme Chave 4 , Pierre-Michel Forget 5 1 ! IEU, Universitt Zrich, Switzerland 2 ! Departamento de Ciencias


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SLIDE 1

A general, community-level test

  • f the Janzen-Connell hypothesis
  • C. E. Timothy Paine1, Natalia Norden2, Christopher

Baraloto3, Jérôme Chave4, Pierre-Michel Forget5

1! IEU, Universität Zürich, Switzerland 2! Departamento de Ciencias Biológicas, Universidad de los Andes, Colombia 3! INRA, UMR "Ecologie des Forêts de Guyane", French Guiana 4! Université Paul Sabatier, CNRS, France 5! Muséum National d’Histoire Naturelle, France

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SLIDE 2

Crax alector eating seed of Lecythis persistens

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SLIDE 3

Seed/seedling survival Seed rain Seedling Recruitment

A form of negative density dependent performance, mediated by (relatively) host-specific natural enemies

Janzen-Connell (JC): Increased diversity by increasing seedling-adult distances

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SLIDE 4

Lots of evidence for JC

Seedling survival is reduced... ! Near conspecific adults ! In crowds of conspecific seedlings ! ...Often because of host-specific insects or pathogens

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* : Tree 3 Distance south of parent tree (m) FIG.
  • 3. Percentage of seedlings of Platypodium elegans on Barro Colorado Island, Panama,
dying from damping-off during the first 3 months as a function of distance along transects south
  • f four parent trees. The lines show the percentage damping-off predicted by the regression if
seedling density and light environment at each distance were at the average value for the transect. Points are the observed levels of damping-off, corrected for variation due to seedling density and light environment, i.e. "~(corr) = yi(obs) + bz('denslty
  • X ~ d e n s i t y ) )
+ b3('Â¥Lsa -x~(sa.p))> where 6, and b3 are the partial regression coefficients for X , + X , respectively, from the multiple regression presented in Table 2). Tree I

I

Tree 2 Distance south of parent t r e e ( rn FIG.
  • 4. Percentage of seedlings of Platypodium elegans on Barro Colorado Island, Panama,
dying from damping-off during the first 3 months at selected distances from Trees 1 and 2. An equal number of seedlings were observed at each distance interval. Shaded bars represent gaps. The initial mean density of seedlings (m-l) occurs above each bar.

Seedling escape from fungalpathogens

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Tree I

LTree

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A 0

1

4 Tree 4 7 Distance south of parent tree (m)

FIG.

  • 1. Seedling density in relation to distance south of four parent trees of Pla[vpodium elegans
  • n Barro Colorado Island, Panama.

TABLE

  • 1. Differences among the four trees of Platjyodium elegans on Barro

Colorado Island, Panama, in distance and density of seedlings south of the parent, and incidence of death of seedlings by damping-off during first 3 months after germination.

Tree 1 2 3 4 Number of seedlings observed 1884 926 298 115 Median distance of seedlings 22 I3 16 12 from parent (m) Seedling density at median 25.0 12.0 2.7 1.5 distance (m ') Seedlings dying (%I) 85.2 80.0 83.9 46.1 Seedlings dying due to 81.2 63.0 54.0 34.8 damping-off (96) Proportion of seedling deaths due 95.2 78.7 64.4 75.5 to damping-off ('XI)

first 3 months was higher within 20 m of the pankt than at any greater distance (Fig. 2). For seedlings of Trees 1 and 2, mortality was higher at intermediate than at far distances (Fig. 2). For Tree 1, seedling mortality was lowest in the large gap at 35-55 m; for Tree 2, mortality at far distances was slightly lower than in the small gap at 5-10 m (Fig. 2). The four trees differed in the proportion of seedlings surviving the first 3 months (Table 1); mortality increased with seedling density.

Mortalify factop

Causes of seedling mortality were determined only in the first 3 months when damping-off was the predominant cause. It accounted for 64% (Tree 3) to 95% (Tree 1) of seedling deaths (Table 1). The remaining mortality was unevenly divided among six other factors: (i) the fall of branches and leaves onto the seedling; (ii) poor root establishment;

Augspurger, 1983, J. Ecology

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SLIDE 5

Why return to JC when so much has already been done?

Most studies performed on few species Studies of recruitment beneath con-/hetero- specifics are limited to species common enough to provide decent sample size What can we add? ! A community-wide, taxonomy-free analysis ! Using functional traits and evolutionary history

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SLIDE 6

Three ways natural enemies could find seedlings

Strictly host-specific Similar evolutionary history Similar functional traits Unlikely Novel adaptations may make closely-related species ecologically divergent Which are relevant to measure? Also, environmental filtering shapes trait distributions Caveat Possibility

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SLIDE 7

Objective: test generality of JC across entire tree community

Why? Lots of evidence for JC for single species, ! generality is little known How? Functional and phylogenetic similarity !

  • f seedlings and adults

Who? Fabulous collaborators!

Chris Baraloto: Traits Jérôme Chave: Phylogeny Natalia Norden: Seedlings Pierre-Michel Forget: Seeds

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SLIDE 8

Sampling of adults (3626) and seedlings (5247)

98 seed collectors, with 202 seedling plots

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SLIDE 9

Sampling functional traits: 7 traits

  • n each of 4672 trees (668 spp)

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Leaf traits: ! Chlorophyll content ! Leaf toughness ! SLA ! C:N Stem traits ! Sapwood density ! Trunk moisture ! Bark thickness Baraloto et al, in press, Ecol Lett

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SLIDE 10

Community phylogeny

611 species of French Guiana rain-forest trees using matK + rbcL

Gonzalez et al 2010 PLoS ONE, J. Ecology

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SLIDE 11

Modeling seedling survival

Stem traits Dissimilarity = Leaf traits Phylogeny

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Cox proportional hazard log(seedling size) ~ + log(seedling density) + log(adult basal area) * dissimilarity

( )

Seedling survival Also stratified by recruitment year

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SLIDE 12

Bigger seedlings survive better Seedling density has little effect

Seedling height (cm)

1 5 10 20 50 1 2 3 4

Ht: <0.0001

Seedling density (m-1)

15 30 50 80 1 2 3 4

N sdl: 0.0211

Median expected survival (years)

No important interactions between seedling height and dissimilarity

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SLIDE 13

1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 1 2 3 4

Leaf trait dissimilarity

BA: ns D: 0.0109 BAxD: 0.0063 Basal area 3.8 5.4 6.1 6.6 8.1

1.0 1.5 2.0 2.5 3.0 3.5 4.0 1 2 3 4

Stem trait dissimilarity

BA: 0.0023 D: ns BAxD: 0.0152

2 3 4 5 1 2 3 4

ALL trait dissimilarity

BA: ns D: ns BAxD: ns

180 220 260 300 1 2 3 4

Phylogenetic dissimilarity

BA: 0.0096 D: 0.0025 BAxD: 0.0023

Median expected survival (years)

Adult-seedling similarity affects survival

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SLIDE 14

How to explain?

Adults with... ! Dissimilar leaves ↑ survival ! Dissimilar stems ↓ survival ! (thus all traits together have no effect) ! Dissimilar evolutionary history ↑ survival

1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 1 2 3 4 Leaf trait dissimilarity BA: ns D: 0.0109 BAxD: 0.0063 Basal area 3.8 5.4 6.1 6.6 8.1 1.0 1.5 2.0 2.5 3.0 3.5 4.0 1 2 3 4 Stem trait dissimilarity BA: 0.0023 D: ns BAxD: 0.0152 2 3 4 5 1 2 3 4 ALL trait dissimilarity BA: ns D: ns BAxD: ns 180 220 260 300 1 2 3 4 Phylogenetic dissimilarity BA: 0.0096 D: 0.0025 BAxD: 0.0023 Median expected survival (years)
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SLIDE 15

How to explain?

JC predicts greater survival where adults very dissimilar from seedlings Support for JC for leaf traits and phylogeny NOT stem traits Environmental filtering on stem traits may counteract natural-enemy-mediated effects

1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 1 2 3 4 Leaf trait dissimilarity BA: ns D: 0.0109 BAxD: 0.0063 Basal area 3.8 5.4 6.1 6.6 8.1 1.0 1.5 2.0 2.5 3.0 3.5 4.0 1 2 3 4 Stem trait dissimilarity BA: 0.0023 D: ns BAxD: 0.0152 2 3 4 5 1 2 3 4 ALL trait dissimilarity BA: ns D: ns BAxD: ns 180 220 260 300 1 2 3 4 Phylogenetic dissimilarity BA: 0.0096 D: 0.0025 BAxD: 0.0023 Median expected survival (years)
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SLIDE 16

Conclusion

Strong support for Janzen-Connell hypothesis at the community level for leaf traits and phylogeny Confirms a long chain of literature Shows generality of JC hypothesis, even for rare spp Environmental filtering also limits seedling survival

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SLIDE 17

What’s next?

Trait/phylogenetic similarity of neighboring seedlings Repeat with minimum (rather than mean) dissimilarities or distance-weighting of adults Simultaneous evaluation of environmental filtering and JC?

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SLIDE 18