junglesm. a. charleston jungles a new solution to the host/parasite phylogeny reconciliation problem...
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Jungles M. A. Charleston
Jungles
A new solution to the host/parasite
phylogeny reconciliation problem
M. A. Charleston
University of Glasgow
Jungles M. A. Charleston
Problem Definition
Given: Host phylogeny H; Associate phylogeny P; Known associations between them .
What were the past associations of taxa?
Jungles M. A. Charleston
Host/Parasite Phylogeny Reconciliation
Associations
Host phylogeny Associate Phylogeny
Jungles M. A. Charleston
Multiple Solutions
Jungles M. A. Charleston
The Model
Parasites are independent new species after speciation.
There are four event types: cospeciation, duplication, lineage sorting, and host switching.
Jungles M. A. Charleston
The Four Event Types
time
Jungles M. A. Charleston
Cospeciation
Associate
Host
Lineage Sorting
Host switch
Duplication
Jungles M. A. Charleston
Event Costs
Event Cost
Cospeciation
Duplication
Lineage Sorting
Host Switch
c ≤ 0
d > 0
s > 0
w > 0
Jungles M. A. Charleston
Jungles
The Jungle is a digraph (directed graph). The vertices of the jungle are the extant &
hypothesized host/parasite associations. The arcs of the jungle correspond to arcs of
the associate tree. The inferred events are dependent only on
the arcs in the jungle.
Jungles M. A. Charleston
The Two J-Vertex Types
Type 1 vertex
h
p
Type 2 vertex
h
p
time
Jungles M. A. Charleston
Jungle Arc Types - 1
sorting events
cospeciation event (type 1 vertex)
(type 1 or 2 vertex)p
p'
time
Jungles M. A. Charleston
Jungle Arc Types - 2
sorting events
duplication event (type 2 vertex)
(type 1 or 2 vertex)p
p'
time
Jungles M. A. Charleston
Jungle Arc Types - 3sorting events
host switching event (type 2 vertex)
(type 1 or 2 vertex)p
p'
take-off
landing sitetermination
time
Jungles M. A. CharlestonJungles M. A. Charleston
Host Switching
Host switching induces a partial ordering on the host tree
time
landing site
take-off
termination
w
w'
x
x'
w’ precedes x
Jungles M. A. Charleston
Switch Incompatibility - 1
Weak Incompatibility:
Temporal incongruence can be fixed by moving landing sites back in time.
s1s2
u
u'
w
x
v
v'
x'
w'
s1 u’ precedes v
s2 w’ precedes x
Jungles M. A. Charleston
Switch Incompatibility - 2
s1 s2
induced sorting event
s1s2
induced sorting events
The order in which we consider host switches affects the number of induced sorting events.
Jungles M. A. Charleston
Switch Incompatibility - 3
Strong Incompatibility:
Moving the landing sites back cannot solve the temporal incongruity.
s1 u’ precedes w
s2 v’ precedes x
s1s2
xx'
w
w'
v
v'
u
u'
- Impossible!
Jungles M. A. Charleston
From Tanglegram...
1
2
3
4
5
6
7
a
b
c
d
e
f
g
Jungles M. A. Charleston
... to Jungle
Lineage sorting
Duplication
Cospeciation
Host switch✝
(f:7)*
(g:6)
(g:5)
(e:7)*
(a:1)
(c:2)
(b:3)
(d:4)
(f:4)
(e:1)
(g:7)*
(e:5)
(g:7)
(f:6)
(f:2)
(f:5)
(e:3)
(e:6)
✝
✝✝
✝✝
✝ ✝
✝
✝
✝
✝✝
✝✝
✝
✝
✝
✝
✝
✝
✝✝✝
✝
✝✝✝
✝
s12
s 1
s 3 s 4
s 5s 6
s 7
s 8
s 9s 10
s 11
s 13 s 14
2s
1
2
3
4
5
6
7
a
b
c
d
e
f
g
Jungles M. A. Charleston
(f:7)*
(g:6)
(g:5)
(e:7)*
(a:1)
(c:2)
(b:3)
(d:4)
(f:4)
(e:1)
(g:7)*
(e:5)
(g:7)
(f:6)
(f:2)
(f:5)
(e:3)
(e:6)
1 1
221
3
2 21
3
4 44
4
4
444
6
6 5
7
88
38
8
8
83
88
6
65
7
1
2 21
331
2 2
1
10
10
3
0 0
00
2
33
5 5
8 6
5 5
3
2
JungleTraversal
1
2
3
4
5
6
7
a
b
c
d
e
f
g
Jungles M. A. Charleston
Optimal Solutions Embedded
1
2
3
4
5
6
7
a
c
b
d
e
f
g
Jungles M. A. Charleston
1
2
3
4
5
6
7
a
c
b
d
e
f
g
All Solutions Embedded
Jungles M. A. Charleston
Gophers and Lice: Tanglegram
a:T.wardi
b:T.minor
c:G.thomomyus
d:G.actuosi
e:G.ewingi
f:G.chapini
g:G.panamensis
h:G.setzeri
i:G.cherriei
j:G.costaricensis
k
l
m
n
o
p
q
r
s
louse tree1:T.talpoides
2:T.bottae
3:G.busarius
4:O.hispidus
5:O.cavator
6:O.underwoodi
7:O.cherriei
8:O.heterodus
9
1011
12
13
14
15
gopher tree
Jungles M. A. Charleston
Gophers and Lice: Jungle
(m:12)*(p:11)*
(p:12)
(p:12)*
(m:6)
(g:5)
(h:6)
(m:5)
(i:7)(j:8)
(n:10)*
(p:5)
(f:4)
(q:11)
(q:5)
(q:13)*
(q:4)
(l:14)
(o:3)
(o:14)
(o:15)*
(o:15)
(o:9)
(o:9)*
(r:15)*
(r:3)
(r:14)
(r:14)*
(r:4)
(r:11)
(r:12)
(r:5)
(r:13)
(s:12)
(s:5)
(s:13)
(s:4)
(s:3)
(s:15)*
(s:15)
(k:9)*
(l:15)*
(d:2)
(e:3)
(l:2)
(l:9)
(l:3)
(c:1)
(o:1)(r:1)
(r:9)
(r:15)
(a:1)
(b:2)
(s:9)
(s:11)
Page's solutionRonquist's solutionBoth solutionsRequired
Jungles M. A. Charleston
1:T.talpoides
2:T.bottae
3:G.busarius
4:O.hispidus
5:O.cavator
6:O.underwoodi
7:O.cherriei
8:O.heterodus
a:T.wardi
b:T.minor
c:G.thomomyus
d:G.actuosi
e:G.ewingi
f:G.chapini
g:G.panamensis
h:G.setzeri
i:G.cherriei
j:G.costaricensis
Page’s Solution
Cospeciation
Duplication
Lineage Sorting
Host Switch
6
1
2
2
Jungles M. A. Charleston
Ronquist’s Solution
1:T.talpoides
2:T.bottae
3:G.busarius
4:O.hispidus
5:O.cavator
6:O.underwoodi
7:O.cherriei
8:O.heterodus
a:T.wardi
b:T.minor
c:G.thomomyus
d:G.actuosi
e:G.ewingi
f:G.chapini
g:G.panamensis
h:G.setzeri
i:G.cherriei
j:G.costaricensis
Cospeciation
Duplication
Lineage Sorting
Host Switch
5
2
2
2
Jungles M. A. Charleston
Conclusions
Complete representation of solution space in one figure Fast solution through dynamic programming Guarantees global optimality Correct accounting for host switches General model, extendable to maximum likelihood