2017 Simons
from nanostructure to macroevolution
Rafael Maia Columbia University
Charley Harper
diversity across the tree of life
Taxa 1 5 10 50 100 500 1000 5000 10000 20000 500
400
300
200 MYA
100
0
why are some groups
more diverse than others?
Joanna Barnum
adaptive radiations
adaptive radiations
adaptive radiations: ecological opportunity
adaptive radiations: key innovations
Pharyngeal jaw
not a product of
stronger selection but of
greater opportunity
Evolution:
Sex & Death
sexual selection & diversification
Can we think about
ornamental traits and diversification in the same way?
COLORS
COLORS are not created equal
pigment-based colors
McGraw et al 2006, illustration: Quinn Burrell
pigment-based colors
pigment-based colors
pheomelanin
pigment-based colors
pheomelanin
eumelanin
reconstructing dino color
Anchiornis huxleyi
reconstructing dino color
Anchiornis huxleyi
pigment-based colors
pigment-based colors
structural colors
structural colors
structural colors
iridescence in feathers
iridescence in feathers
iridescence in feathers melanin-containing organelle (melanosome)
form & function: structure determines color
form & function: structure determines color
form & function: structure determines color
form & function: structure determines color
form & function: structure determines color
Reflectance (%)
form & function: structure determines color
300
400
500
600
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
form & function: structure determines color
300
400
500
600
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
form & function: structure determines color
300
400
500
600
n1
d1
n2
d2
n1
d1
n2
d2
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
form & function: structure determines color
300
400
500
600
n1
d1
n2
d2
n1
d1
n2
d2
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
structural colors
variation through shared mechanisms
How do color-producing mechanisms influence color diversification?
an integrative approach to the study of avian coloration
an integrative approach to the study of avian coloration
Development
an integrative approach to the study of avian coloration
Function
Reflectance (%)
Development
300
400
500
600
Wavelength (nm)
700
an integrative approach to the study of avian coloration
Reflectance (%)
Development
Function
300
400
500
600
700
s13
s12
Wavelength (nm)
s11
s14
s10
s15
s9
s16
s8
s17
s7
s18
s19
s6
s20
s5
s21
s3
s23
s2
s24
s1
s25
s50
s49
s26
s48
s27
s47
s28
s46
s29
s45
s30
s44
s31
s32
s43
s33
s42
s41
s34
s35
s40
s39
s38
s37
s36
Evolution
s4
s22
?
How do color-producing structures become organized?
feather development Earliest Barb Ridges
keratinocyte (barbule)
Later
Rachis
Follicle Collar
melanocyte
♂
♀
Blue-black grassquit (Volatinia jacarina)
1
melanin deposition and organization are decoupled
2
organization likely not cell-mediated
3
likely related to the amount of melanin and speed of keratinization
organization through self-assembly
University of Pennsylvania, Department of Physics and Astronomy
organization through self-assembly
University of Pennsylvania, Department of Physics and Astronomy
?
How does the structural template influence color evolution?
color is a multidimensional trait
A
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Lamprotornis fischeri Saroglossa spiloptera Lamprotornis bicolor Grafisia torquata Lamprotornis regius Speculipastor bicolor Lamprotornis chalcurus Poeoptera sharpii Lamprotornis nitens Poeoptera femoralis Lamprotornis purpureus Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis iris Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis elisabeth Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis shelleyi
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Notopholia corruscus Lamprotornis fischeri Saroglossa spiloptera Hylopsar cupreocauda Lamprotornis bicolor Grafisia torquata Hylopsar purpureiceps Lamprotornis regius Speculipastor bicolor Lamprotornis australis Lamprotornis chalcurus Poeoptera sharpii Lamprotornis caudatus Lamprotornis nitens Poeoptera femoralis Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis unicolor Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis mevesii Lamprotornis iris Lamprotornis splendidus Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis ornatus Lamprotornis elisabeth Lamprotornis pulcher Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis superbus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis albicapillus Lamprotornis shelleyi
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Notopholia corruscus Lamprotornis fischeri Saroglossa spiloptera Hylopsar cupreocauda Lamprotornis bicolor Grafisia torquata Hylopsar purpureiceps Lamprotornis regius Speculipastor bicolor Lamprotornis australis Lamprotornis chalcurus Poeoptera sharpii Lamprotornis caudatus Lamprotornis nitens Poeoptera femoralis Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis unicolor Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis mevesii Lamprotornis iris Lamprotornis splendidus Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis ornatus Lamprotornis elisabeth Lamprotornis pulcher Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis superbus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis albicapillus Lamprotornis shelleyi
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Notopholia corruscus Lamprotornis fischeri Saroglossa spiloptera Hylopsar cupreocauda Lamprotornis bicolor Grafisia torquata Hylopsar purpureiceps Lamprotornis regius Speculipastor bicolor Lamprotornis australis Lamprotornis chalcurus Poeoptera sharpii Lamprotornis caudatus Lamprotornis nitens Poeoptera femoralis Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis unicolor Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis mevesii Lamprotornis iris Lamprotornis splendidus Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis ornatus Lamprotornis elisabeth Lamprotornis pulcher Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis superbus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis albicapillus Lamprotornis shelleyi
starlings (Sturnidae)
reconstructing evolution
reconstructing evolution
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
Evolutionary stability of optical complexity
Reflectance (%) 300
400
500
600
Wavelength (nm)
700
520 540
560
500
CIE Y
580
` 490
600 620 700
480 470 460
380
CIE X
Absorbance
how birds see the world
uv
300
s
400
m
500
l
600
Wavelength (nm)
700
how birds see the world
how birds see the world
747 individuals from 47 species 10-15 body patches
Reflectance (%)
creating an avian color space
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
creating an avian color space
300
400
500
600
700
Wavelength (nm)
Absorbance
+
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
creating an avian color space
300
400
500
600
700
Wavelength (nm)
=
Absorbance
+
300
400
500
600
Wavelength (nm)
700
PC2
PC1
non-iridescent ● ●
● ●● ● ●
● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ●● ●
● ●
●
● ●
● ● ● ● ●
● ● ●
● ● ● ● ● ● ● ● ● ● ● ●● ●●● ●● ●● ●● ● ● ●● ● ●● ● ● ● ● ● ● ● ● ● ●●● ● ● ●●● ● ● ● ● ●●● ● ● ●● ●●● ●●●● ● ● ● ● ●● ● ●●●● ● ●● ● ● ● ●● ● ● ● ● ●●● ●● ● ● ●● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ● ●●●●● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ●●● ● ●● ●● ● ●● ●
●
●
PC2
●
●
PC1
●
●
non-iridescent + ● ●
● ●● ● ●
● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ●● ●
● ●
● ●
PC2
● ●
●
● ● ● ●● ● ●● ● ● ● ● ● ●● ● ● ● ●● ● ●● ● ●●● ● ● ● ● ● ● ●● ●●●● ●● ● ● ● ● ● ● ● ● ● ●● ● ● ●● ● ● ●● ● ● ● ● ● ● ● ●● ● ●●● ● ● ● ● ● ●● ● ●● ● ● ●● ● ● ●● ●●●●● ● ●● ●● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ●●●● ●● ●● ●● ● ●● ●●● ● ● ●● ● ● ● ● ● ●●● ● ● ● ● ●● ● ● ● ●● ●●● ●●●● ● ●● ● ●● ● ●● ● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ●● ● ● ●● ●● ● ● ●●●●● ● ● ● ● ● ●● ● ● ●● ● ● ● ●● ● ● ● ●●●● ● ● ●● ● ● ● ●● ●● ● ● ● ●●● ● ● ●● ● ● ●● ● ● ●●●● ●●● ●●● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ●●●●● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●●●●● ●●● ● ●● ● ● ● ● ● ● ●● ● ●●●●● ● ● ● ● ● ● ● ●
● ●
PC1
●
non-iridescent +
+ ● ●
●
● ●
● ●● ● ●
● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ●● ● ●
● ●
● ●
PC2
● ●
●
●
● ●
● ●
●
●
●
●
●
●
●●
●
●
● ●● ●
● ● ● ● ● ●
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● ●
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PC1
●●
●
How fast is evolution?
modeling evolution: Brownian Motion σ2 = 1.0
Phenotype
σ2 = 0.1
0.0
rate parameter
2 σ
Time
1.0
0.0
Time
1.0
modeling evolution: Brownian Motion
σ21
Phenotype
σ2 = 0.1 σ2 = 1.0
σ
2 2
0.0
Time
1.0
modeling evolution: Ornstein-Uhlenbeck σ2 = 1.0 α = 10
Phenotype
σ2 = 1.0 α = 0.0
0.0
Time
1.0
0.0
Time
rate parameter σ2, attraction parameter α, optimal trait value θ
1.0
modeling evolution: Ornstein-Uhlenbeck
θ1
Phenotype
θ=1 θ=3
θ2 0.0
Time
1.0
non-iridescent +
+ ● ●
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● ●
● ●
PC2
● ●
innovations result in
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●
●
●●
●
●
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● ●
40x faster evolution of color
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●
●
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PC1
●●
●
non-iridescent +
+ ● ●
●
● ●
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● ●
● ●
PC2
● ●
innovations result in
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●
●●
●
●
● ●● ●
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● ●
●●
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and faster
40x faster evolution of color
●
● ●
●
●
●
●
speciation
●
PC1
Melanosome morphology underlies
diversification
Proximate and developmental factors are critical to color evolution
Proximate and developmental factors are critical to color evolution
optical innovations: key component of ornamental diversification
current & future directions
The genomic and molecular basis of phenotypic novelty
A
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
current & future directions
The genomic and molecular basis of phenotypic novelty
A
B
Linking microevolution and macroevolution of coloration
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
current & future directions
The genomic and molecular basis of phenotypic novelty
A
B
Linking microevolution and macroevolution of coloration
The role of color innovations in avian diversification
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
Tiffany Bozic, “Oil Slick"
Dustin Rubenstein Luke Harmon Matthew Shawkey Chad Eliason Regina Macedo
Rafael Maia Columbia University
Charley Harper
diversity across the tree of life
Taxa 1 5 10 50 100 500 1000 5000 10000 20000 500
400
300
200 MYA
100
0
why are some groups
more diverse than others?
Joanna Barnum
adaptive radiations
adaptive radiations
adaptive radiations: ecological opportunity
adaptive radiations: key innovations
Pharyngeal jaw
not a product of
stronger selection but of
greater opportunity
Evolution:
Sex & Death
sexual selection & diversification
Can we think about
ornamental traits and diversification in the same way?
COLORS
COLORS are not created equal
pigment-based colors
McGraw et al 2006, illustration: Quinn Burrell
pigment-based colors
pigment-based colors
pheomelanin
pigment-based colors
pheomelanin
eumelanin
reconstructing dino color
Anchiornis huxleyi
reconstructing dino color
Anchiornis huxleyi
pigment-based colors
pigment-based colors
structural colors
structural colors
structural colors
iridescence in feathers
iridescence in feathers
iridescence in feathers melanin-containing organelle (melanosome)
form & function: structure determines color
form & function: structure determines color
form & function: structure determines color
form & function: structure determines color
form & function: structure determines color
Reflectance (%)
form & function: structure determines color
300
400
500
600
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
form & function: structure determines color
300
400
500
600
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
form & function: structure determines color
300
400
500
600
n1
d1
n2
d2
n1
d1
n2
d2
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
form & function: structure determines color
300
400
500
600
n1
d1
n2
d2
n1
d1
n2
d2
700
Reflectance (%)
Wavelength (nm)
300
400
500
600
Wavelength (nm)
700
structural colors
variation through shared mechanisms
How do color-producing mechanisms influence color diversification?
an integrative approach to the study of avian coloration
an integrative approach to the study of avian coloration
Development
an integrative approach to the study of avian coloration
Function
Reflectance (%)
Development
300
400
500
600
Wavelength (nm)
700
an integrative approach to the study of avian coloration
Reflectance (%)
Development
Function
300
400
500
600
700
s13
s12
Wavelength (nm)
s11
s14
s10
s15
s9
s16
s8
s17
s7
s18
s19
s6
s20
s5
s21
s3
s23
s2
s24
s1
s25
s50
s49
s26
s48
s27
s47
s28
s46
s29
s45
s30
s44
s31
s32
s43
s33
s42
s41
s34
s35
s40
s39
s38
s37
s36
Evolution
s4
s22
?
How do color-producing structures become organized?
feather development Earliest Barb Ridges
keratinocyte (barbule)
Later
Rachis
Follicle Collar
melanocyte
♂
♀
Blue-black grassquit (Volatinia jacarina)
1
melanin deposition and organization are decoupled
2
organization likely not cell-mediated
3
likely related to the amount of melanin and speed of keratinization
organization through self-assembly
University of Pennsylvania, Department of Physics and Astronomy
organization through self-assembly
University of Pennsylvania, Department of Physics and Astronomy
?
How does the structural template influence color evolution?
color is a multidimensional trait
A
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Lamprotornis fischeri Saroglossa spiloptera Lamprotornis bicolor Grafisia torquata Lamprotornis regius Speculipastor bicolor Lamprotornis chalcurus Poeoptera sharpii Lamprotornis nitens Poeoptera femoralis Lamprotornis purpureus Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis iris Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis elisabeth Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis shelleyi
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Notopholia corruscus Lamprotornis fischeri Saroglossa spiloptera Hylopsar cupreocauda Lamprotornis bicolor Grafisia torquata Hylopsar purpureiceps Lamprotornis regius Speculipastor bicolor Lamprotornis australis Lamprotornis chalcurus Poeoptera sharpii Lamprotornis caudatus Lamprotornis nitens Poeoptera femoralis Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis unicolor Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis mevesii Lamprotornis iris Lamprotornis splendidus Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis ornatus Lamprotornis elisabeth Lamprotornis pulcher Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis superbus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis albicapillus Lamprotornis shelleyi
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Notopholia corruscus Lamprotornis fischeri Saroglossa spiloptera Hylopsar cupreocauda Lamprotornis bicolor Grafisia torquata Hylopsar purpureiceps Lamprotornis regius Speculipastor bicolor Lamprotornis australis Lamprotornis chalcurus Poeoptera sharpii Lamprotornis caudatus Lamprotornis nitens Poeoptera femoralis Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis unicolor Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis mevesii Lamprotornis iris Lamprotornis splendidus Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis ornatus Lamprotornis elisabeth Lamprotornis pulcher Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis superbus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis albicapillus Lamprotornis shelleyi
A
A
B
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Notopholia corruscus Lamprotornis bicolor Hylopsar cupreocauda Lamprotornis regius Hylopsar purpureiceps Lamprotornis chalcurus Lamprotornis australis Lamprotornis nitens Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis unicolor Lamprotornis iris Lamprotornis mevesii Lamprotornis acuticaudus Lamprotornis splendidus Lamprotornis elisabeth Lamprotornis ornatus Lamprotornis chloropterus Lamprotornis pulcher Lamprotornis hidebrandti Lamprotornis superbus Lamprotornis shelleyi Lamprotornis albicapillus Neocichla gutturalis Notopholia corruscus Lamprotornis fischeri Saroglossa spiloptera Hylopsar cupreocauda Lamprotornis bicolor Grafisia torquata Hylopsar purpureiceps Lamprotornis regius Speculipastor bicolor Lamprotornis australis Lamprotornis chalcurus Poeoptera sharpii Lamprotornis caudatus Lamprotornis nitens Poeoptera femoralis Lamprotornis purpuroptera Lamprotornis purpureus Lamprotornis unicolor Lamprotornis chalybaeus Poeoptera stuhlmanni Poeoptera kenricki Lamprotornis mevesii Lamprotornis iris Lamprotornis splendidus Lamprotornis acuticaudus Poeoptera lugubris Onychognathus nabouroup Lamprotornis ornatus Lamprotornis elisabeth Lamprotornis pulcher Lamprotornis chloropterus Onychognathus fulgidus Lamprotornis superbus Lamprotornis hidebrandti Onychognathus walleri Onychognathus salvadorii Lamprotornis albicapillus Lamprotornis shelleyi
starlings (Sturnidae)
reconstructing evolution
reconstructing evolution
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
Evolutionary stability of optical complexity
Reflectance (%) 300
400
500
600
Wavelength (nm)
700
520 540
560
500
CIE Y
580
` 490
600 620 700
480 470 460
380
CIE X
Absorbance
how birds see the world
uv
300
s
400
m
500
l
600
Wavelength (nm)
700
how birds see the world
how birds see the world
747 individuals from 47 species 10-15 body patches
Reflectance (%)
creating an avian color space
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
creating an avian color space
300
400
500
600
700
Wavelength (nm)
Absorbance
+
300
400
500
600
Wavelength (nm)
700
Reflectance (%)
creating an avian color space
300
400
500
600
700
Wavelength (nm)
=
Absorbance
+
300
400
500
600
Wavelength (nm)
700
PC2
PC1
non-iridescent ● ●
● ●● ● ●
● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ●● ●
● ●
●
● ●
● ● ● ● ●
● ● ●
● ● ● ● ● ● ● ● ● ● ● ●● ●●● ●● ●● ●● ● ● ●● ● ●● ● ● ● ● ● ● ● ● ● ●●● ● ● ●●● ● ● ● ● ●●● ● ● ●● ●●● ●●●● ● ● ● ● ●● ● ●●●● ● ●● ● ● ● ●● ● ● ● ● ●●● ●● ● ● ●● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ● ●●●●● ● ● ● ● ● ● ● ●● ● ● ● ● ● ● ●●● ● ●● ●● ● ●● ●
●
●
PC2
●
●
PC1
●
●
non-iridescent + ● ●
● ●● ● ●
● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ●● ●
● ●
● ●
PC2
● ●
●
● ● ● ●● ● ●● ● ● ● ● ● ●● ● ● ● ●● ● ●● ● ●●● ● ● ● ● ● ● ●● ●●●● ●● ● ● ● ● ● ● ● ● ● ●● ● ● ●● ● ● ●● ● ● ● ● ● ● ● ●● ● ●●● ● ● ● ● ● ●● ● ●● ● ● ●● ● ● ●● ●●●●● ● ●● ●● ● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ●●●● ●● ●● ●● ● ●● ●●● ● ● ●● ● ● ● ● ● ●●● ● ● ● ● ●● ● ● ● ●● ●●● ●●●● ● ●● ● ●● ● ●● ● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ●● ● ● ●● ●● ● ● ●●●●● ● ● ● ● ● ●● ● ● ●● ● ● ● ●● ● ● ● ●●●● ● ● ●● ● ● ● ●● ●● ● ● ● ●●● ● ● ●● ● ● ●● ● ● ●●●● ●●● ●●● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ● ● ● ● ● ● ● ●● ● ● ●●●●● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●●●●● ●●● ● ●● ● ● ● ● ● ● ●● ● ●●●●● ● ● ● ● ● ● ● ●
● ●
PC1
●
non-iridescent +
+ ● ●
●
● ●
● ●● ● ●
● ● ● ● ● ● ● ● ● ● ●● ●● ● ● ● ● ●● ● ●
● ●
● ●
PC2
● ●
●
●
● ●
● ●
●
●
●
●
●
●
●●
●
●
● ●● ●
● ● ● ● ● ●
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● ●
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PC1
●●
●
How fast is evolution?
modeling evolution: Brownian Motion σ2 = 1.0
Phenotype
σ2 = 0.1
0.0
rate parameter
2 σ
Time
1.0
0.0
Time
1.0
modeling evolution: Brownian Motion
σ21
Phenotype
σ2 = 0.1 σ2 = 1.0
σ
2 2
0.0
Time
1.0
modeling evolution: Ornstein-Uhlenbeck σ2 = 1.0 α = 10
Phenotype
σ2 = 1.0 α = 0.0
0.0
Time
1.0
0.0
Time
rate parameter σ2, attraction parameter α, optimal trait value θ
1.0
modeling evolution: Ornstein-Uhlenbeck
θ1
Phenotype
θ=1 θ=3
θ2 0.0
Time
1.0
non-iridescent +
+ ● ●
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● ●
● ●
PC2
● ●
innovations result in
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●
●
●●
●
●
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● ●
40x faster evolution of color
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●
●
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PC1
●●
●
non-iridescent +
+ ● ●
●
● ●
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● ●
● ●
PC2
● ●
innovations result in
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●
●●
●
●
● ●● ●
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● ●
●●
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and faster
40x faster evolution of color
●
● ●
●
●
●
●
speciation
●
PC1
Melanosome morphology underlies
diversification
Proximate and developmental factors are critical to color evolution
Proximate and developmental factors are critical to color evolution
optical innovations: key component of ornamental diversification
current & future directions
The genomic and molecular basis of phenotypic novelty
A
B
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
current & future directions
The genomic and molecular basis of phenotypic novelty
A
B
Linking microevolution and macroevolution of coloration
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
current & future directions
The genomic and molecular basis of phenotypic novelty
A
B
Linking microevolution and macroevolution of coloration
The role of color innovations in avian diversification
Notopholia corruscus Hylopsar cupreocauda Hylopsar purpureiceps Lamprotornis australis Lamprotornis caudatus Lamprotornis purpuroptera Lamprotornis unicolor Lamprotornis mevesii Lamprotornis splendidus Lamprotornis ornatus Lamprotornis pulcher Lamprotornis superbus Lamprotornis albicapillus Lamprotornis fischeri Lamprotornis bicolor Lamprotornis regius Lamprotornis chalcurus Lamprotornis nitens Lamprotornis purpureus Lamprotornis chalybaeus Lamprotornis iris Lamprotornis acuticaudus Lamprotornis elisabeth Lamprotornis chloropterus Lamprotornis hidebrandti Lamprotornis shelleyi Neocichla gutturalis Saroglossa spiloptera Grafisia torquata Speculipastor bicolor Poeoptera sharpii Poeoptera femoralis Poeoptera stuhlmanni Poeoptera kenricki Poeoptera lugubris Onychognathus nabouroup Onychognathus fulgidus Onychognathus walleri Onychognathus salvadorii Onychognathus blythii Onychognathus frater Onychognathus tristramii Onychognathus albirostris Onychognathus morio Onychognathus neumanni Onychognathus tenuirostris Cinnyrincinclus leucogaster Hartlaubius aurata
Tiffany Bozic, “Oil Slick"
Dustin Rubenstein Luke Harmon Matthew Shawkey Chad Eliason Regina Macedo
