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Putting the algal tree of life to use
Evolutionary dynamics of ecological niches, physiology
and species’ diagnostic trait...
Algal Diversity
• architecture
• functional traits
• life history
• physiology
Algal Tree of Life
• systematics: species delimitation
and higher-level relationships
• phylogenies in evolutionary
enquir...
Talk contents
Systematics
Thermal Niche
Trace Elements
Thermal niche
van den Hoek. 1982. Biol. J. Linn. Soc. 18: 81-144 — Cambridge et al. 1984. Helgol. Meeresunt. 38: 349-363 —...
Thermal niche
Map: T. Schils
Species diversity map
Codium
How does the thermal niche evolve?
How fast?
Which direction?
Pulsed or gradual?
Effect on biodiversity?
Are there covaria...
Exploration
Which direction?
Codium Dictyotales
Tropics
Temperate
0.011
0.027
Tropics
Temperate
0.028
0.032
Are there covariates?
sheltered exposed
HalimedaMarcelino et al. 2014. submitted
Pulsed or gradual?
Dictyota
Thermal niche & diversification
DictyotaTyberghein et al. unpublished
Thermal niche evolution
• thermal niches evolve over geological timescales
• microhabitat preferences affect evolvability
...
Talk contents
Systematics
Thermal Niche
Trace Elements
Trace element utilization
Raven. 1999. Photosynthesis Research 60: 111-149
Thylakoid membrane with photosystems
Trace element utilization
Quigg et al. 2003. Nature 425: 291-294
Hypothesis
Exploration
Cya
Eug
Cha
Stra
Din
Din
Gla
Hap
Cry
Rho
Chl
time
(Ga) 00.511.5
Fe
utilization
low high
Hypothesis Model
Cya
Eug
Cha
Stra
Din
Gla
Hap
Cry
Rho
Chl
Cya
Eug
Cha
Stra
Din
Din
Gla
Hap
Cry
Rho
Chl
Host phylogeny Plas...
Results for Fe/P ratio
Pulsed Gradual
0 1
0.25
Nuclear Tree
0.40
Plastid Tree
Relative rates
σ 2ENDO = 73.5
σ 2OTHER = 447...
Simulation study
5.05!
7.55!
3.64!
7.94!
3.54!
4.23!
2.42!
5.92!
6.55!
9.78!
3.93!
4.95!
5.17!
5.14!
6.04!
5.10!
5.53!
4.2...
Power analysis
Trace element utilization
• results are inconclusive
• support for existing hypotheses is limited
• power to detect differ...
Talk contents
Systematics
Thermal Niche
Trace Elements
Role of ToL in Systematics
Simulation: extrapolate what we know about trait evolution
A
B
C
Problem: Cryptic Diversity
Zuccarello, G.C., West, J.A. 2003. J. Phycol. 39: 948-959. — Photo: J. West
How big is the prob...
Pseudochlorodesmis
Verbruggen, H. et al. 2009. J. Phycol. 45: 726-731 — Photo: H. Verbruggen
Is it more of a problem in si...
Complexity and diagnosability
Verbruggen, H. 2014. J. Phycol. 50: 26-31.
Simulation: extrapolate what we know about trait ...
Complexity and diagnosability
20 characters
theoretical maximum = 1,048,576
10 characters
theoretical maximum = 1,024
Verb...
Habitat selection
Verbruggen, H. 2014. J. Phycol. 50: 26-31.
A
B
C
D
E
➙ 1
➙ 3
➙ 5
➙ 7
➙ 9
Habitat selection
Verbruggen, H. 2014. J. Phycol. 50: 26-31.
Plasticity
Verbruggen, H. 2014. J. Phycol. 50: 26-31.
A
B
C
D
E
➙ 1
➙ 3
➙ 5
➙ 7
➙ 9
Plasticity
Verbruggen, H. 2014. J. Phycol. 50: 26-31.
Morphological evolution
• # unique morphologies << number of species
• null hypothesis: cryptic diversity abounds
• fundam...
Talk contents
Systematics
Thermal Niche
Trace Elements
Conclusions
• test evolutionary hypotheses
• insights from parameters
• measure uncertainty
• simulation to study behavior...
Acknowledgements
• Collectors
• Collaborators
• Previous workers
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Putting the algal tree of life to use

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The increase in resolution and taxon sampling of algal phylogenies resulting from the various algal tree of life projects and other initiatives worldwide opens tremendous opportunity to learn more about the evolution of all aspects of algal biology. Using evolutionary modeling techniques in a phylogenetic context, hypotheses about the evolution of particular traits and their interaction with speciation-extinction dynamics become testable. I will illustrate this with three case studies. First, I will investigate the evolution of the thermal niche of seaweeds, showing how it affects latitudinal diversity patterns. Second, I will test the hypothesis that the evolution of cellular trace element requirements (stoichiometry) is dominated by endosymbiosis events. Third, I will investigate the evolution of morphological traits typically used in species-level systematics, focusing on its implications for the prevalence of cryptic diversity. These case studies show the potential and limitations of the approach, and offer new insights in algal evolution from the very recent to the very ancient, and across the various subdisciplines of algal biology.

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Putting the algal tree of life to use

  1. 1. Putting the algal tree of life to use Evolutionary dynamics of ecological niches, physiology and species’ diagnostic traits Heroen Verbruggen School of Botany, University of Melbourne
  2. 2. Algal Diversity • architecture • functional traits • life history • physiology
  3. 3. Algal Tree of Life • systematics: species delimitation and higher-level relationships • phylogenies in evolutionary enquiry • endosymbiosis events: patchwork of genes → accumulation of genomic features • examples of phylogenetics applied in evolutionary questions
  4. 4. Talk contents Systematics Thermal Niche Trace Elements
  5. 5. Thermal niche van den Hoek. 1982. Biol. J. Linn. Soc. 18: 81-144 — Cambridge et al. 1984. Helgol. Meeresunt. 38: 349-363 — Eggert. 2012. In: Wiencke, C., Bischof, K. [Eds.] Seaweed Biology
  6. 6. Thermal niche Map: T. Schils Species diversity map Codium
  7. 7. How does the thermal niche evolve? How fast? Which direction? Pulsed or gradual? Effect on biodiversity? Are there covariates?
  8. 8. Exploration
  9. 9. Which direction? Codium Dictyotales Tropics Temperate 0.011 0.027 Tropics Temperate 0.028 0.032
  10. 10. Are there covariates? sheltered exposed HalimedaMarcelino et al. 2014. submitted
  11. 11. Pulsed or gradual? Dictyota
  12. 12. Thermal niche & diversification DictyotaTyberghein et al. unpublished
  13. 13. Thermal niche evolution • thermal niches evolve over geological timescales • microhabitat preferences affect evolvability • diversification relates to SST and its evolvability • results are taxon-specific • scale up to bigger datasets: more species • harder questions: adaptation, interactions, timescales
  14. 14. Talk contents Systematics Thermal Niche Trace Elements
  15. 15. Trace element utilization Raven. 1999. Photosynthesis Research 60: 111-149 Thylakoid membrane with photosystems
  16. 16. Trace element utilization Quigg et al. 2003. Nature 425: 291-294 Hypothesis
  17. 17. Exploration Cya Eug Cha Stra Din Din Gla Hap Cry Rho Chl time (Ga) 00.511.5 Fe utilization low high
  18. 18. Hypothesis Model Cya Eug Cha Stra Din Gla Hap Cry Rho Chl Cya Eug Cha Stra Din Din Gla Hap Cry Rho Chl Host phylogeny Plastid phylogeny Parfrey, L.W., Grant, J., et al. 2010. Syst. Biol. 59: 518-533 — Baurain, D., Brinkmann, H., et al. 2010. Mol. Biol. Evol. 27: 1698-1709.
  19. 19. Results for Fe/P ratio Pulsed Gradual 0 1 0.25 Nuclear Tree 0.40 Plastid Tree Relative rates σ 2ENDO = 73.5 σ 2OTHER = 447.3 Nuclear tree p = 0.09 Relative rates σ 2ENDO = 103.2 σ 2OTHER = 382.9 Plastid tree
  20. 20. Simulation study 5.05! 7.55! 3.64! 7.94! 3.54! 4.23! 2.42! 5.92! 6.55! 9.78! 3.93! 4.95! 5.17! 5.14! 6.04! 5.10! 5.53! 4.29! 3.76! 5.05! 2.98! 9.28! 3.49! 2.07! 5.67! 7.11! 6.07! 6.05! 7.95! 3.36! 3.95! 0.12! 3.80! 4.02! 5.63! 4.57! 5.22! 7.35! 5.55 non-ES rate ES rate 1 100 0.01 1 100 0.01
  21. 21. Power analysis
  22. 22. Trace element utilization • results are inconclusive • support for existing hypotheses is limited • power to detect differential evolution of continuous traits is low • limited effect of increase in taxon sampling • metalloproteomics approaches
  23. 23. Talk contents Systematics Thermal Niche Trace Elements
  24. 24. Role of ToL in Systematics Simulation: extrapolate what we know about trait evolution A B C
  25. 25. Problem: Cryptic Diversity Zuccarello, G.C., West, J.A. 2003. J. Phycol. 39: 948-959. — Photo: J. West How big is the problem?
  26. 26. Pseudochlorodesmis Verbruggen, H. et al. 2009. J. Phycol. 45: 726-731 — Photo: H. Verbruggen Is it more of a problem in simple organisms?
  27. 27. Complexity and diagnosability Verbruggen, H. 2014. J. Phycol. 50: 26-31. Simulation: extrapolate what we know about trait evolution 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 0 0 1 0 1 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 0 1 1 0 1 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 0 1 0 1 1 1 0 0 1 1 1 1 1 1 1
  28. 28. Complexity and diagnosability 20 characters theoretical maximum = 1,048,576 10 characters theoretical maximum = 1,024 Verbruggen, H. 2014. J. Phycol. 50: 26-31. • higher complexity more unique morphologies • # unique morphologies << # species in clade total # species #uniquemorphologies
  29. 29. Habitat selection Verbruggen, H. 2014. J. Phycol. 50: 26-31. A B C D E ➙ 1 ➙ 3 ➙ 5 ➙ 7 ➙ 9
  30. 30. Habitat selection Verbruggen, H. 2014. J. Phycol. 50: 26-31.
  31. 31. Plasticity Verbruggen, H. 2014. J. Phycol. 50: 26-31. A B C D E ➙ 1 ➙ 3 ➙ 5 ➙ 7 ➙ 9
  32. 32. Plasticity Verbruggen, H. 2014. J. Phycol. 50: 26-31.
  33. 33. Morphological evolution • # unique morphologies << number of species • null hypothesis: cryptic diversity abounds • fundamental inability to distinguish some species (possibly many) morphologically • problem more pronounced in simple organisms • plasticity blurs species boundaries
  34. 34. Talk contents Systematics Thermal Niche Trace Elements
  35. 35. Conclusions • test evolutionary hypotheses • insights from parameters • measure uncertainty • simulation to study behavior of methods • simulation to improve expectations • perspectives: genome dynamics, life histories, etc. Verbruggen, Marcelino, Costa (2014) Evolutionary dynamics of algal traits and diversity. Perspectives in Phycology 1: in press 
 [available from www.phycoweb.net]
  36. 36. Acknowledgements • Collectors • Collaborators • Previous workers

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