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On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird, Utah Monday, June 24 th , 2013
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On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

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Page 1: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

On patterns of coevolution

across cophylogenies

Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali

1

Evolution 2013 Snowbird, Utah Monday, June 24th, 2013

Page 2: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Species interactions

wormwould, (2010), Parasitic Wasp [ONLINE]. Available at: http://www.flickr.com/photos/onthespiral [Accessed 20 January 13]

Fir0002/Flagstaffotos, (2007), Leaf Hopper nymph of the Common Jassid (Eurymela fenestrata) [ONLINE]. Available at:

http://en.wikipedia.org/wiki/File:Common_jassid_nymphs_and_ants02.jpg [Accessed 20 January 13].

Dr Graham Beards, (2009), Electron micrograph of Bacteriophages [ONLINE]. Available at: http://commons.wikimedia.org/wiki/File:Phage.jpg [Accessed 20 January 13].

John Severns, (2006), Honey bee [ONLINE]. Available at: http://en.wikipedia.org/wiki/File:European_honey_bee_extracts_nectar.jpg [Accessed 20 January 13].

Richard Naylor, (2009), Parasitoid wasps [ONLINE]. Available at: http://news.bbc.co.uk/local/sheffield/hi/people_and_places/nature/newsid_8370000/8370816.stm [Accessed 20

January 13].

2

Page 3: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Symbiotic relationships

Persistent, recurring biological

interactions

Mutualism, commensalism, parasitism

Obligate and facultative relationships

Shared phylogenetic topology

Reciprocal speciation between the two

associated lineages; cospeciation

But what about when they don’t match?

Phylogenetic reconstruction

Superimpose phylogenies, explain

differences with events

Jan Derk, (2002), Common Clownfish [ONLINE]. Available at: http://en.wikipedia.org/wiki/File:Common_clownfish_curves_dnsmpl.jpg [Accessed 20 January 13].

Rafael Barbizan Sühs, (2006), Wasp [ONLINE]. Available at: http://en.wikipedia.org/wiki/File:Hymenoptera,_Vespidae,_Polistinae,_Mischocyttarus_rotundicollis_.jpg

[Accessed 20 January 13].

Brian Valentine, (2006), Small wasp parasitising aphids [ONLINE]. Available at: http://www.flickr.com/photos/lordv/2869916160/ [Accessed 20 January 13].

3

Page 4: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Coevolutionary Events

Event types

Cospeciation

Sorting

Duplication

Host switch

Cospeciation Sorting

Duplication Host switch

Host

Symbiont

4

Page 5: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Past research

Specific, small scale studies

Mostly single cophylogeny comparison

Tested individually for congruence

No direct comparisons between phylogenies

5

Page 6: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Questions

Is the symbiont phylogeny dependent on the

host phylogeny?

How can the coevolutionary history of

symbiotic species be quantified & compared

across groups?

How do profiles of coevolutionary events

differ across cophylogeny classifications?

Latitude, transmission (horizontal, vertical),

association (ecto, endo, intracellular), etc.

6

Page 7: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Overview of analysis

Co-phylogeny collection

Single, standardized Web of Science search

Both symbiont phylogenies, and associations

Transfer of cophylogenies to digital format

Co-phylogenetic history reconstruction

Core-PA (cost-based: fixed, parameter adaptive)

Develop permutation test

Use dirmult package in R

7

Page 8: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Symbiont categories

0

5

10

15

20

25

30

Josh Grosse, (2002), Roundworm [ONLINE]. Available at: http://en.wikipedia.org/wiki/File:Roundworm.jpg [Accessed 20 January 13].

CDC/Vector Ecology & Control Laboratory, (2007), Flea [ONLINE]. Available at: http://static.ddmcdn.com/gif/flea-1.jpg [Accessed 20 January 13].

Thomas Splettstoesser, (2012), Cowpea mosaic virus [ONLINE]. Available at: http://www.scistyle.com [Accessed 20 January 13].

Rafael Barbizan Sühs, (2006), Wasp [ONLINE]. Available at: http://en.wikipedia.org/wiki/File:Hymenoptera,_Vespidae,_Polistinae,_Mischocyttarus_rotundicollis_.jpg [Accessed 20 January

13].

8

Page 9: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Overview of analysis

Co-phylogeny collection

Single, standardized Web of Science search

Both symbiont phylogenies, and associations

Transfer of cophylogenies to digital format

Co-phylogenetic history reconstruction

Core-PA (cost-based: fixed, parameter adaptive)

Develop permutation test

Use dirmult package in R

9

Page 10: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Dirichlet distribution

How to compare cophylogeny event profiles?

Given total number of coevolutionary events,

event counts follow a multinomial distribution

Extension of binomial distribution for binary data

Assume event proportions among host-symbiont

pairs within a group (e.g. latitude zone) are similar

Expect variance within these groups

A Dirichlet distribution provides a model for how

these proportions vary

10

Page 11: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

2

8

4

1

Calculate LLRT

“Tropical” “Temperate”

2

1

2

1

Cospeciation Sorting Duplication Host-switch

Dirichlet model & permutation test

X 999 2

8

4

1

Cospeciation Sorting Duplication Host-switch

Calculate LLRT statistic

Tropical Temperate

2

1

2

1

11

2

8

4

1

Calculate LLRT

“Tropical” “Temperate”

2

1

2

1

Cospeciation Sorting Duplication Host-switch

2

8

4

1

Calculate LLRT statistic

“Tropical” “Temperate”

2

1

2

1

Cospeciation Sorting Duplication Host-switch

Fit Dirichlet distribution model to event counts

Log Likelihood Ratio Test (LLRT) statistic

Randomized permutations to build distribution

Page 12: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

12

LLRT statistic

Page 13: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Exploratory biological results (α = 0.2)

Of interest for further study

Ecto-endo symbionts (expected)

Innate-adaptive immune system (vert/invert)

Tropical vs. temperate

Not significant

Horizontal/vertical transmission (unexpected!)

Parasites/mutualist symbionts

Presence/absence of secondary hosts

Tropical-subtropical

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Page 14: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Parameter Adaptive results:

Ratios of Proportions Speciation Event

Comparison Cospeciation Sorting Duplication Host-switch p

Tropical vs

Temperate 0.96 1.08 0.90 0.54 0.10

Ecto- vs Endo-

symbiont 1.06 0.95 1.03 1.63 0.16

Adaptive vs

Innate *

1.31 0.90 0.98 4.53 0.03

14

* Vertebrate vs Invertebrate immune system

Less host-switching in the tropics

Page 15: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Parameter Adaptive results:

Ratios of Proportions Speciation Event

Comparison Cospeciation Sorting Duplication Host-switch p

Tropical vs

Temperate 0.96 1.08 0.90 0.54 0.10

Ecto- vs Endo-

symbiont 1.06 0.95 1.03 1.63 0.16

Adaptive vs

Innate *

1.31 0.90 0.98 4.53 0.03

15

* Vertebrate vs Invertebrate immune system

Less host-switching in the tropics

Page 16: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Parameter Adaptive results:

Ratios of Proportions Speciation Event

Comparison Cospeciation Sorting Duplication Host-switch p

Tropical vs

Temperate 0.96 1.08 0.90 0.54 0.10

Ecto- vs Endo-

symbiont 1.06 0.95 1.03 1.63 0.16

Adaptive vs

Innate *

1.31 0.90 0.98 4.53 0.03

16

* Vertebrate vs Invertebrate immune system

More host switching in ecto-symbionts

Page 17: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Parameter Adaptive results:

Ratios of Proportions Speciation Event

Comparison Cospeciation Sorting Duplication Host-switch p

Tropical vs

Temperate 0.96 1.08 0.90 0.54 0.10

Ecto- vs Endo-

symbiont 1.06 0.95 1.03 1.63 0.16

Adaptive vs

Innate *

1.31 0.90 0.98 4.53 0.03

17

* Vertebrate vs Invertebrate immune system

More cospeciation and host-switching in hosts with adaptive immune systems

Page 18: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Conclusions & future directions

Conducted meta-analysis of cophylogenies

Developed statistical test to compare event

profiles from several copylogenies

Identified knowledge gaps requiring study

Repeat comparisons within specific taxa

Perform Dirichlet regression test to

simultaneously include several factors into

one parsimonious model

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Page 19: On patterns of coevolution across cophylogenies 2013.pdf · On patterns of coevolution across cophylogenies Mark Merilo, Sarah J. Adamowicz, R. Ayesha Ali 1 Evolution 2013 Snowbird,

Acknowledgements

Jinzhong Fu, Alex Smith

Nicolas Wieseke

Adamowicz, Hebert labs

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