MULTIDISCIPLINRY NATURE OF THE ENVIRONMENTAL STUDIES.pptx
Detecting evolutionary regime shifts with comparative phylogenetics
1. Introduction Parrotfish: a story A new method Challenges Opportunities
Detecting evolutionary
regime shifts with
comparative
phylogenetics
@cboettig
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 1/61
2. Introduction Parrotfish: a story A new method Challenges Opportunities
Once upon a time. . .
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 2/61
3. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogeny was a nuisance.
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 3/61
4. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogeny was a nuisance.
something we “corrected for”
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 3/61
5. Introduction Parrotfish: a story A new method Challenges Opportunities
Then we turned our attention to the parameters
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 4/61
6. Introduction Parrotfish: a story A new method Challenges Opportunities
Then we turned our attention to the parameters
2 Diversification
σ
α Stabilizing selection
θ Adaptive Optimum
λ phylogenetic signal
κ speciational change
δ accelerating / decelerating rates
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 4/61
7. Introduction Parrotfish: a story A new method Challenges Opportunities
The parameters were allowed to change,
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 5/61
8. Introduction Parrotfish: a story A new method Challenges Opportunities
The parameters were allowed to change, we became interested
in where the changes occur.
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 5/61
9. Introduction Parrotfish: a story A new method Challenges Opportunities
Regime
Shifts
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 6/61
10. Introduction Parrotfish: a story A new method Challenges Opportunities
The (Updated) Phylogeny of Phylogenetic Models
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 7/61
11. Introduction Parrotfish: a story A new method Challenges Opportunities
Outline
1 A Method:
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 8/61
12. Introduction Parrotfish: a story A new method Challenges Opportunities
Outline
1 A Method: Detecting a release of constraint
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 8/61
13. Introduction Parrotfish: a story A new method Challenges Opportunities
Outline
1 A Method: Detecting a release of constraint
2 Challenges:
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 8/61
14. Introduction Parrotfish: a story A new method Challenges Opportunities
Outline
1 A Method: Detecting a release of constraint
2 Challenges: The edge of identifiability
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 8/61
15. Introduction Parrotfish: a story A new method Challenges Opportunities
Outline
1 A Method: Detecting a release of constraint
2 Challenges: The edge of identifiability
3 Opportunities:
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 8/61
16. Introduction Parrotfish: a story A new method Challenges Opportunities
Outline
1 A Method: Detecting a release of constraint
2 Challenges: The edge of identifiability
3 Opportunities: Large, synthetic data sets
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 8/61
17. Introduction Parrotfish: a story A new method Challenges Opportunities
A key innovation?
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 9/61
18. Introduction Parrotfish: a story A new method Challenges Opportunities
A key innovation?
This guy
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 9/61
19. Introduction Parrotfish: a story A new method Challenges Opportunities
A key innovation?
This guy These fish
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 9/61
20. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 10/61
21. Introduction Parrotfish: a story A new method Challenges Opportunities
Fish Suck
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 11/61
22. Introduction Parrotfish: a story A new method Challenges Opportunities
Fish Suck
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 12/61
23. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 13/61
24. Introduction Parrotfish: a story A new method Challenges Opportunities
These fish don’t suck
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 14/61
25. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 15/61
26. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 16/61
27. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 17/61
28. Introduction Parrotfish: a story A new method Challenges Opportunities
Did this innovation
release the constraints
of suction feeding?
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 18/61
29. Introduction Parrotfish: a story A new method Challenges Opportunities
. . . and release a burst of jaw evolution?
Parrotfish
Intramandibular Pharyngeal
Joint Jaw
Wrasses
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 19/61
30. Introduction Parrotfish: a story A new method Challenges Opportunities
An evolutionary regime shift?
a large change in the model parameters
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 20/61
31. Introduction Parrotfish: a story A new method Challenges Opportunities
An evolutionary regime shift?
a large change in the model parameters
In which parameters?
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 20/61
32. Introduction Parrotfish: a story A new method Challenges Opportunities
Candidate models
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 21/61
33. Introduction Parrotfish: a story A new method Challenges Opportunities
Unconstrained Evolution
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 22/61
34. Introduction Parrotfish: a story A new method Challenges Opportunities
Constrained Evolution
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 23/61
35. Introduction Parrotfish: a story A new method Challenges Opportunities
Release of constraint
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 24/61
36. Introduction Parrotfish: a story A new method Challenges Opportunities
Accelerating rates
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 25/61
37. Introduction Parrotfish: a story A new method Challenges Opportunities
2
Brownian Motion 2
Ornstein-Uhlenbeck
1 1
trait value
trait value
0 0
-1 -1
-2 -2
0 20 40 60 80 100 0 20 40 60 80 100
time time
2
Accelerated Evolution 2
Release of Constraint
1 1
trait value
trait value
0 0
-1 -1
-2 -2
0 20 40 60 80 100 0 20 40 60 80 100
time time
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 26/61
38. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 27/61
39. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 28/61
40. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 29/61
41. Introduction Parrotfish: a story A new method Challenges Opportunities
XT1 = X0 e−α1 T1 +
where you start
θ1 1 − e−α1 T1 +
Where you are going
T1
σ1 eα1 (s−T1 ) dWs
0
Random shocks along the way
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 30/61
42. Introduction Parrotfish: a story A new method Challenges Opportunities
XT1 = X0 e−α1 T1 +
where you start
θ1 1 − e−α1 T1 +
Where you are going
T1
σ1 eα1 (s−T1 ) dWs
0
Random shocks along the way
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 30/61
43. Introduction Parrotfish: a story A new method Challenges Opportunities
XT2 = X1 e−α2 T2 +
where you were
θ2 1 − e−α2 T2 +
possibly new optimum
T2
σ2 eα2 (s−T2 ) dWs
T1
possibly new shocks
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 31/61
44. Introduction Parrotfish: a story A new method Challenges Opportunities
XT2 = X1 e−α2 T2 +
where you were
θ2 1 − e−α2 T2 +
possibly new optimum
T2
σ2 eα2 (s−T2 ) dWs
T1
possibly new shocks
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 31/61
45. Introduction Parrotfish: a story A new method Challenges Opportunities
XT2 = X1 e−α2 T2 +
where you were
θ2 1 − e−α2 T2 +
possibly new optimum
T2
σ2 eα2 (s−T2 ) dWs
T1
possibly new shocks
n
γi = − αj t j
j=i+1
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 31/61
46. Introduction Parrotfish: a story A new method Challenges Opportunities
Generalized OU model
n
XN = eγi θi 1 − eαi ti +
i=0
ti
σi eαi (s−ti ) dWs
i
0
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 32/61
47. Introduction Parrotfish: a story A new method Challenges Opportunities
Some Mathematical Conveniences
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 33/61
48. Introduction Parrotfish: a story A new method Challenges Opportunities
Some Mathematical Conveniences
Gaussian process!
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 33/61
49. Introduction Parrotfish: a story A new method Challenges Opportunities
Some Mathematical Conveniences
Gaussian process! Need only
E(Xi ) and Cov(Xi , Xj )
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 33/61
50. Introduction Parrotfish: a story A new method Challenges Opportunities
Some Mathematical Conveniences
Gaussian process! Need only
E(Xi ) and Cov(Xi , Xj )
Evolution on different branches is
independent
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 33/61
51. Introduction Parrotfish: a story A new method Challenges Opportunities
Averaging removes the random shocks
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 34/61
52. Introduction Parrotfish: a story A new method Challenges Opportunities
Averaging removes the random shocks
n
E(Xi ) = eγi θi 1 − eαi ti
i=0
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 34/61
53. Introduction Parrotfish: a story A new method Challenges Opportunities
Averaging removes the random shocks
n
E(Xi ) = eγi θi 1 − eαi ti
i=0
Itô Isometry makes stochastic integrals into
regular integrals!
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 34/61
54. Introduction Parrotfish: a story A new method Challenges Opportunities
Averaging removes the random shocks
n
E(Xi ) = eγi θi 1 − eαi ti
i=0
Itô Isometry makes stochastic integrals into
regular integrals!
E( f (Xi )dWi × f (Xj )dWj )
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 34/61
55. Introduction Parrotfish: a story A new method Challenges Opportunities
Averaging removes the random shocks
n
E(Xi ) = eγi θi 1 − eαi ti
i=0
Itô Isometry makes stochastic integrals into
regular integrals!
E( f (Xi )dWi × f (Xj )dWj )
Cov(Xi , Xj ) =
n ti
eγi σi eαi (u−ti ) dWu ×
i
i=1 0
m sj
eγj σj eαj (v−sj ) dWv
j
j=1 0
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 34/61
56. Introduction Parrotfish: a story A new method Challenges Opportunities
Covariance matrix determines the likelihood
Cov(Xi , Xj ) =
k 2
σs
eγs +γs 1 − e−2αs ts
s=0
2αs
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 35/61
57. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 36/61
58. Introduction Parrotfish: a story A new method Challenges Opportunities
OUwie
Beaulieu et al. 2012, Evolution
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 36/61
59. Introduction Parrotfish: a story A new method Challenges Opportunities
Challenges
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 37/61
60. Introduction Parrotfish: a story A new method Challenges Opportunities
Where the sidewalk ends
Generalizing models any further,
we will not have analytic likelihoods
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 38/61
61. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 39/61
62. Introduction Parrotfish: a story A new method Challenges Opportunities
Limits of Identifiability?
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 40/61
63. Introduction Parrotfish: a story A new method Challenges Opportunities
Can our data support so many parameters?
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 41/61
64. Introduction Parrotfish: a story A new method Challenges Opportunities
How do we choose between multiple complex models?
Before we can ask which is the right model,
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 42/61
65. Introduction Parrotfish: a story A new method Challenges Opportunities
How do we choose between multiple complex models?
Before we can ask which is the right model,
we must ask if we have enough data to tell.
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 42/61
66. Introduction Parrotfish: a story A new method Challenges Opportunities
How do we choose between multiple complex models?
Before we can ask which is the right model,
we must ask if we have enough data to tell.
Information citeria don’t address this.
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 42/61
67. Introduction Parrotfish: a story A new method Challenges Opportunities
How do we choose between multiple complex models?
Before we can ask which is the right model,
we must ask if we have enough data to tell.
Information citeria don’t address this.
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 42/61
68. Introduction Parrotfish: a story A new method Challenges Opportunities
AIC scores for four model comparisons
(a) BM vs. OU.3 (b) OU.3 vs. OU.4
0.15
0.08
0.10
Density
Density
0.04
0.05
0.00
0.00
0 10 20 30 40 50 -10 0 10 20 30
δ δ
(c) OU.3 vs. OU.15 (d) BM vs. OU.1
0.04
0.20
0.03
0.15
Density
Density
0.02
0.10
0.01
0.05
0.00
0.00
0 50 100 150 -5 0 5 10 15
δ δ
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 43/61
69. Introduction Parrotfish: a story A new method Challenges Opportunities
AIC scores for four model comparisons
(a) BM vs. OU.3 (b) OU.3 vs. OU.4
False Pos (37%) False Pos (43.75%)
0.15
False Neg (0%) False Neg (8.25%)
0.08
0.10
Density
Density
0.04
0.05
0.00
0.00
0 10 20 30 40 50 -10 0 10 20 30
δ δ
(c) OU.3 vs. OU.15 (d) BM vs. OU.1
0.04
False Pos (47.75%) False Pos (19.95%)
0.20
False Neg (0%) False Neg (76.65%)
0.03
0.15
Density
Density
0.02
0.10
0.01
0.05
0.00
0.00
0 50 100 150 -5 0 5 10 15
δ δ
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 44/61
70. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
m
fro →
ate l
mul mode
Si ple
sim
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 45/61
71. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
m
fro →
ate l
mul mode
Si ple
sim del s
h mo
ts bot a
← Fi is dat
to th
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 46/61
72. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
m
fro →
ate l
mul mode
Si ple
sim del s
h mo
ts bot a
← Fi is dat
to th
LogLik(complex) LogLik(simple)
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 47/61
73. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
m
fro →
ate l
mul mode
Si ple
sim del s
h mo
ts bot a
← Fi is dat
to th
LogLik(complex) LogLik(simple)
frequency
Likelihood ratio
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 48/61
74. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
m
fro →
ate l
mul mode
Si ple
sim del s
h mo
ts bot a
← Fi is dat
to th
LogLik(complex) LogLik(simple)
frequency
Likelihood ratio
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 49/61
75. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
m
fro →
ate l
mul mode
Si ple
sim del s
h mo
ts bot a
← Fi is dat
to th
LogLik(complex) LogLik(simple)
frequency
Likelihood ratio
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 50/61
76. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
x
p le
m comdel →
fro → mo
ate l
mul mode
Si ple
sim del s
h mo
ts bot a
← Fi is dat
to th
LogLik(complex) LogLik(simple)
frequency
Likelihood ratio
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 51/61
77. Introduction Parrotfish: a story A new method Challenges Opportunities
Phylogenetic Monte Carlo
x
p le
om comdel →
e fr → mo
ulat odel
Sim ple m
sim l s
mode
ts both
← Fi is data
to th
LogLik(complex) LogLik(simple)
frequency
Likelihood ratio
Boettiger et al. 2012, Evolution, pmc R package
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 52/61
78. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 53/61
79. Introduction Parrotfish: a story A new method Challenges Opportunities
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 54/61
80. Introduction Parrotfish: a story A new method Challenges Opportunities
Software has made
comparative methods
ubiquitous. . .
So how about data?
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 55/61
81. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic Access to data
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 56/61
82. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic Access to data
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 56/61
83. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic Access to data
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 56/61
84. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic Access to data
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 56/61
85. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic access for everyone
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 57/61
86. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic access for everyone
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 57/61
87. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic access for everyone
Building tools, building community
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 57/61
88. Introduction Parrotfish: a story A new method Challenges Opportunities
Programmatic access for everyone
Building tools, building community
Follow @ropensci for more
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 57/61
89. Introduction Parrotfish: a story A new method Challenges Opportunities
A wealth of repositories
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 58/61
90. Introduction Parrotfish: a story A new method Challenges Opportunities
and what about the Parrotfish?
Was it a release of constraint after all?
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 59/61
91. Introduction Parrotfish: a story A new method Challenges Opportunities
and what about the Parrotfish?
Was it a release of constraint after all?
Of course!
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 59/61
92. Introduction Parrotfish: a story A new method Challenges Opportunities
Conclusions
1 Can distinguish mechanisms such as release of
constraint.
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 60/61
93. Introduction Parrotfish: a story A new method Challenges Opportunities
Conclusions
1 Can distinguish mechanisms such as release of
constraint.
2 Pushing the limits of analytic solutions,
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 60/61
94. Introduction Parrotfish: a story A new method Challenges Opportunities
Conclusions
1 Can distinguish mechanisms such as release of
constraint.
2 Pushing the limits of analytic solutions,
3 and more worryingly, the limits of our data
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 60/61
95. Introduction Parrotfish: a story A new method Challenges Opportunities
Conclusions
1 Can distinguish mechanisms such as release of
constraint.
2 Pushing the limits of analytic solutions,
3 and more worryingly, the limits of our data
4 New and diverse data sources would alleviate, but not
solve these issues
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 60/61
96. Introduction Parrotfish: a story A new method Challenges Opportunities
Conclusions
1 Can distinguish mechanisms such as release of
constraint.
2 Pushing the limits of analytic solutions,
3 and more worryingly, the limits of our data
4 New and diverse data sources would alleviate, but not
solve these issues
5 Continue to foster a community of data sharing and code
sharing
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 60/61
97. Introduction Parrotfish: a story A new method Challenges Opportunities
Acknowledgments
Jeremy Beaulieu
Peter Wainwright
Graham Coop
Peter Ralph
Brian O’Meara
Duncan Temple
Lang
Slides, source code, and more
and many more
http://carlboettiger.info
Carl Boettiger, UC Davis Detecting Evolutionary Regime Shifts 61/61