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Leaf variation in Ficus:
an evolutionary perspective

Kandlikar Gau-Lou
Nguyen Hai Huyen
Wu Wei-Huan
Yao Xin
Zeng Si-Jin
2...
Outline
•
•
•
•
•

Introduction to leaf diversity and Ficus
Our hypotheses
How we tested them
What we found
What it means
...
Leaf diversity
• Leaves are the
organs primarily
responsible for
photosynthesis
• Evolutionary
patterns of leaf
variation ...
Introduction to Ficus
• Ficus is a large pan-tropical genus with >800
species
• Lot of variation in life history patterns ...
Hemiepiphytic Ficus
• Many Ficus species have a “hemi-epiphytic” life
style
• Epiphytic stage
associated with
water stress...
Leaf variation in Ficus

6
Leaf size variation
• Large leaves require more “structure” to hold up
against gravity (Niklas 1999)
– Structure might com...
Hypotheses
• HI: Hemiepiphytic figs have evolved to deal
with water stressed environments
– Prediction: Hemiepiphytic figs...
Study site
• Xishuangbanna Tropical Botanical Garden
= sampled individuals

Ficus collection

9
Methods
• Sampling intensity:
– 2 trees/species; 3 leaves/tree

• Leaf traits measurements according to
published protocol...
F hispida
F fistulosa
F semicordata
●

F sur
F beipeiensis
F auriculata
F oligodon

Phylogenetic
Methods

• Used a publish...
Data Analysis
• Testing prediction 1
– T-Tests to compare leaf area and SLA of HE/Ter

• Testing prediction 2
– Test for r...
Results
F hispida
F fistulosa
F semicordata
F sur
F beipeiensis
F auriculata
F oligodon
F racemosa
F tikoua
F villosa
F sa...
Results
• Prediction 1 (Hemiepiphytes ~ smaller leaves with low SLA)
Log Area of HE and Terrestrial Ficus species

SLA of ...
Results, cont.
• Prediction 2 (Leaf area ~ Leaf toughness)
Log Area vs. Toughness

5
4
3

log Area (cm^2)

6

p=0.91

200
...
Results, cont.
• Prediction 3 (Leaf area ~ Leaf toughness across the
phylogeny)

16
Reconstruction of Toughness

Reconstruction of log Area

F hispida
F hispida
F hispida
F hispida
F fistulosa
F fistulosa
F...
Results, cont.
• Prediction 3
PICs of Log Area vs. PICs of Toughness

1
0
-1
-2

PICs of log area

2

p = 0.34
p=0.36

-50...
Discussion
• Prediction 1 (Hemiepiphytic figs ~ low leaf area and SLA)
– Not supported
– Environmental factors drive repor...
Discussion
• Prediction 2 (Leaf area ~ lamina toughness)
– Not supported
– Veins and petioles may be sufficient to support...
Discussion: Limitations
• Sampling intensity may not have accounted for
intraspecific variation in leaf traits
• Unbalance...
Future Directions
• The evolutionary history of leaves is not well
understood
• Ficus is an ideal group to study this dive...
Acknowledgements
• AFEC Instructors and Organizers
– Liu Jing-Xin, Drs. Richard Corlett, Alice Hughes, Kyle
Tomlinson, Uro...
References
•
•
•
•
•

•
•
•
•
•
•

Cruaud, A., et al. 2012. An extreme case of plant-insect codiversification: figs and fi...
3
1

2

4

5

1. South China Agricultural University
3,5. Xishuangbanna Tropical Botanical Garden
25
2. University of Scie...
Questions?

26
F. auriculata

F. ruficaulis

F. hirta

F. cyathistipula

F. deltoidea

F. ischnopoda
F. hispida

F. cyrtophylla

27
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Leaf traits of Ficus: an evolutionary prespective

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A presentation for the 2013 Applied Fieldcourse in Ecology and Evolution at the Xishuangbanna Tropical Botanical Garden.

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Leaf traits of Ficus: an evolutionary prespective

  1. 1. Leaf variation in Ficus: an evolutionary perspective Kandlikar Gau-Lou Nguyen Hai Huyen Wu Wei-Huan Yao Xin Zeng Si-Jin 28 November 2013
  2. 2. Outline • • • • • Introduction to leaf diversity and Ficus Our hypotheses How we tested them What we found What it means 2
  3. 3. Leaf diversity • Leaves are the organs primarily responsible for photosynthesis • Evolutionary patterns of leaf variation are not well understood (Jones et al. 2008) Leaf variation in Pelargonium. Image from Nicotra et al. 2011 3
  4. 4. Introduction to Ficus • Ficus is a large pan-tropical genus with >800 species • Lot of variation in life history patterns and ecology, with an equally complex evolutionary history (Harrison 2005) • Leaves within the genus exhibit tremendous diversity 4
  5. 5. Hemiepiphytic Ficus • Many Ficus species have a “hemi-epiphytic” life style • Epiphytic stage associated with water stress (Holbrook & Putz 1996, Hao et al. 2010) Epiphytic and Terrestrial F. concinna, from Hao G.Y., et al. 2010 5
  6. 6. Leaf variation in Ficus 6
  7. 7. Leaf size variation • Large leaves require more “structure” to hold up against gravity (Niklas 1999) – Structure might come in many ways (Westbrook et al., 2012) • For large leaves to evolve, ancestors needed to have leaves with greater structure 7
  8. 8. Hypotheses • HI: Hemiepiphytic figs have evolved to deal with water stressed environments – Prediction: Hemiepiphytic figs have smaller leaves with lower specific leaf area (SLA), even when grown in common gardens with terrestrial figs • HII: Large leaves are harder to hold up than small leaves; thus they need to be tougher – Prediction: Leaf size ~ Leaf lamina toughness – Prediction: Leaf size and leaf toughness show patterns of correlated evolution 8
  9. 9. Study site • Xishuangbanna Tropical Botanical Garden = sampled individuals Ficus collection 9
  10. 10. Methods • Sampling intensity: – 2 trees/species; 3 leaves/tree • Leaf traits measurements according to published protocols (Perez-Harguindeguy et al. 2013) – Leaf Area, Toughness, and SLA • Life history data (i.e. epiphytic/terrestrial) collected from the Flora of China (eFloras 2008) 10
  11. 11. F hispida F fistulosa F semicordata ● F sur F beipeiensis F auriculata F oligodon Phylogenetic Methods • Used a published Ficus phylogeny (Cruaud et al. ● F racemosa F tikoua F villosa F sarmentosa F deltoidea F stenophylla F ischnopoda F hirta F ruficaulis F tinctoria F subulata F cyrtophylla F henryi ● 2012) • Thirty species shared between XTBG Ficus collection and the phylogeny ● ● F concinna F cyathistipula ● ● F virens F religiosa ● ● ● ● F stricta ● F maclellandii F benjamina ● ● ● F altissima F annulata F glaberrima ● ● ● 11
  12. 12. Data Analysis • Testing prediction 1 – T-Tests to compare leaf area and SLA of HE/Ter • Testing prediction 2 – Test for relationship between Leaf Area and Toughness • Testing prediction 3 – Ancestral Trait Reconstruction and Phylogenetically Independent Contrasts (PICs) of Area and Toughness in R package phytools (Felsenstein 1985; Revell 2012) 12
  13. 13. Results F hispida F fistulosa F semicordata F sur F beipeiensis F auriculata F oligodon F racemosa F tikoua F villosa F sarmentosa F deltoidea F stenophylla F ischnopoda F hirta F ruficaulis F tinctoria F subulata F cyrtophylla F henryi F virens F religiosa F concinna F cyathistipula F stricta F maclellandii F benjamina F altissima F annulata F glaberrima SLA Area Toughness 13
  14. 14. Results • Prediction 1 (Hemiepiphytes ~ smaller leaves with low SLA) Log Area of HE and Terrestrial Ficus species SLA of HE and epiphytic and terrestrial Ficus specie SLA variation between Terrestrial Ficus species 250 p=0.40 200 150 SLA 5 100 4 3 Area log logArea 6 p=0.90 Epi Ter Epi Ter 14
  15. 15. Results, cont. • Prediction 2 (Leaf area ~ Leaf toughness) Log Area vs. Toughness 5 4 3 log Area (cm^2) 6 p=0.91 200 400 600 800 1000 1200 1400 Toughness (N/m^2) 15
  16. 16. Results, cont. • Prediction 3 (Leaf area ~ Leaf toughness across the phylogeny) 16
  17. 17. Reconstruction of Toughness Reconstruction of log Area F hispida F hispida F hispida F hispida F fistulosa F fistulosa F fistulosa F fistulosa F semicordata F semicordata F semicordata F semicordata F sur F sur F sur F sur F beipeiensis F beipeiensis F beipeiensis F beipeiensis F auriculata F auriculata F auriculata F auriculata F oligodon F oligodon F oligodon F oligodon F racemosa F racemosa F racemosa F racemosa F tikoua F tikoua F tikoua F tikoua F villosa F villosa F villosa F villosa F sarmentosa F sarmentosa F sarmentosa F sarmentosa F deltoidea F deltoidea F deltoidea F deltoidea F stenophylla F stenophylla F stenophylla F stenophylla F ischnopoda F ischnopoda F ischnopoda F ischnopoda F hirta F hirta F hirta F hirta F ruficaulis F ruficaulis F ruficaulis F ruficaulis F tinctoria F tinctoria F tinctoria F tinctoria F subulata F subulata F subulata F subulata F cyrtophylla F cyrtophylla F cyrtophylla F cyrtophylla F henryi F henryi F henryi F henryi F virens F virens F virens F virens F religiosa F religiosa F religiosa F religiosa F concinna F concinna F concinna F concinna F cyathistipula F cyathistipula F cyathistipula F cyathistipula F stricta F stricta F stricta F stricta F maclellandii F maclellandii F maclellandii F maclellandii F benjamina F benjamina F benjamina F benjamina F altissima F altissima F altissima F altissima F annulata F annulata F annulata F annulata F glaberrima F glaberrima F glaberrima F glaberrima 3.05 trait value 6.787 6.787 2 20 cm3.05trait value 800 cm2 length=0.023 length=0.023 239.572 1374.305 239.572 200 grams trait value 1500 trait value 1374.305 length=0.023 length=0.023 grams 17
  18. 18. Results, cont. • Prediction 3 PICs of Log Area vs. PICs of Toughness 1 0 -1 -2 PICs of log area 2 p = 0.34 p=0.36 -500 0 500 PICs of toughness 1000 18
  19. 19. Discussion • Prediction 1 (Hemiepiphytic figs ~ low leaf area and SLA) – Not supported – Environmental factors drive reported leaf variation (e.g. Holbrook & Putz 1996) – Hemiepiphytes may have evolved other mechanisms to deal with water stress 19
  20. 20. Discussion • Prediction 2 (Leaf area ~ lamina toughness) – Not supported – Veins and petioles may be sufficient to support large leaves. • Prediction 3 (Leaf area ~ lamina toughness across phylogeny) – Not supported – Evolution of leaf size and toughness not correlated across the phylogeny 20
  21. 21. Discussion: Limitations • Sampling intensity may not have accounted for intraspecific variation in leaf traits • Unbalanced data set to test Hypothesis I • Ficus phylogeny is still being developed (e.g. Yao et al. 2013), and our methods are sensitive to the phylogeny used 21
  22. 22. Future Directions • The evolutionary history of leaves is not well understood • Ficus is an ideal group to study this diversity – Morphological, distributional, and life history variation • Studies based on leaf morphology, structure, venation, and architecture (Nicotra et al., 2011) • Characterize leaf transcriptomes – Determine molecular basis for variation 22
  23. 23. Acknowledgements • AFEC Instructors and Organizers – Liu Jing-Xin, Drs. Richard Corlett, Alice Hughes, Kyle Tomlinson, Uromi Goodale, Eben Goodale, Ferry Slik – Other XTBG research groups, especially the Evolutionary Ecology, Plant Ecophysiology, and PlantAnimal Interaction groups, and Pan Bo for helping with Ficus identification • XTBG Ficus collection management staff • Numerous tutorials/guides to comparative phylogenetics in R 23
  24. 24. References • • • • • • • • • • • Cruaud, A., et al. 2012. An extreme case of plant-insect codiversification: figs and figpollinating wasps. Systematic Botany 61(6), 1029-1047 Felsenstein K. 1985. Phylogenetics and the comparative method. American Naturalist 125, 1-15.. Hao, G-Y., et al. 2010. Differentiation in leaf water flux and drought tolerance traits in hemiepiphytic and non-hemiepiphytic Ficus tree species. Functional Ecology 24(4), 731740 Harrison, R.D. 2005. Figs and the diversity of tropical rainforests. BioScience 55(12), 1053-1064. Holbrook, N.M., and Putz, F.E. 1996. From epiphyte to tree: differences in leaf structure and leaf water relations associated with the transition in growth form in eight species of hemiepiphytes. Plant, Cell and Environment 19, 631-642. Jones, C.S., et al. 2008. Leaf shape evolution in the South African genus Pelargonium L’He’R. (Geraniaceae). Evolution 63-2, 479-497 Niklas, K.J. A mechanical perspective on foliage leaf form and function. New Phytologist 143, 19-31. Nicotra, A.B., et al. The evolution and functional significance of leaf shape in the Angiosperms. Functional Plant Biology 38, 535-552 Revell, L.J. phytools: an R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution 3(2), 217-223. Westbrook, J.W. et al. 2011. What makes a leaf tough? Patterns of correlated evolution between leaf toughness traits and demographic rates among 197 shade tolerant woody species in a Neotropical forest. The American Naturalist 177(6), 800-811. Yao, X., et al. Exon-Primed Intron-Crossing (EPIC) Markers for Evolutionary Studies of Ficus and Other Taxa in the Fig Family (Moraceae). Appications in Plant Science 1(10). 24
  25. 25. 3 1 2 4 5 1. South China Agricultural University 3,5. Xishuangbanna Tropical Botanical Garden 25 2. University of Science, Vietnam National University 4. University of Minnesota
  26. 26. Questions? 26
  27. 27. F. auriculata F. ruficaulis F. hirta F. cyathistipula F. deltoidea F. ischnopoda F. hispida F. cyrtophylla 27

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