First published online January 31, 2006
Journal of Experimental Biology 209, 748-765 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02051
Anatomically diverse butterfly scales all produce structural colours by coherent scattering
Richard O. Prum1,*,
Tim Quinn2 and
Rodolfo H. Torres3
1 Department of Ecology and Evolutionary Biology, and Peabody Museum of
Natural History, Yale University, PO Box 208105, New Haven, Connecticut 06250,
USA
2 Department of Ecology and Evolutionary Biology, University of Kansas,
Lawrence, KS 66045, USA
3 Department of Mathematics, University of Kansas, Lawrence, KS 66045,
USA

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Fig. 1. Photographs of the butterfly species examined. Uraniidae: (A) Urania
fulgens; Papilionidae: (B) Papilio ulysses, (C) Papilio
zalmoxis, (D) Parides sesostris, (E) Troides brookiana,
(F) Troides urvillianus, (G) Troides priamus priamus
(phenotypically similar to the green T. p. hecuba examined);
Lycaenidae: (H) Celastrina ladon, (I) Callophrys dumetorum,
(J) Mitoura gryneus and (K) Parrhasius m-album (very similar
and closely related to the P. moctezuma examined); Nymphalidae: (L)
Morpho aega. Photos are of upper wing surfaces, except for
Callophrys dumetorum (I) and Mitoura gryneus (J), which are
underwing surfaces. Not to scale. Specimens courtesy of the Yale Peabody
Museum of Natural History Department of Entomology.
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Fig. 2. Light microscope photographs of the structurally coloured scales of a
sample of the Lepidoptera examined. (A) Urania fulgens blue, (B)
Urania fulgens green, (C) Urania fulgens showing the
transition between blue and green patches, (D,E) Papilio ulysses, (F)
Papilio zalmoxis, (G) Parides sesostris, (H) Troides
urvillianus, (I) Troides priamus. All images at 63x,
except C (20x) and D (40x). Classification of species examined is
given in Table 1.
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Fig. 3. Light microscope photographs of the structurally coloured scales of a
sample of the Lepidoptera examined. (A) Celastrina ladon, (B)
Callophrys dumetorum, (C) Mitoura gryneus, (D)
Parrhasius moctezuma and (E,F) Morpho aega. All images at
63x, except D (70x) and E (7.5x). Classification of species
examined is given in Table
1.
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Fig. 4 Transmission electron micrographs (TEMs) of sections of the structurally
coloured scales of a sample of the lepidopteran species examined. (A,B)
Urania fulgens blue, (C) Urania fulgens green, (D,E)
Papilio ulysses, (F,G) Papilio zalmoxis, (H-J) Parides
sesostris, (K,L) Troides brookiana. Scale bars, 500 nm (A,D,
F,G), 200 nm (B,C,E,J,L), 2 µm (H), and 1 µm (K).
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Fig. 5. Transmission electron micrographs (TEMs) of the structurally coloured
scales of a sample of the Lepidoptera examined. (A-B) Troides
urvillianus, (C) Troides priamus, (D) Celastrina ladon,
(E,F) Callophrys dumetorum, (G) Mitoura gryneus, (H)
Parrhasius moctezuma, (I) Morpho aega. Scale bars, 500 nm
(A,D,E,H), 200 nm (B,C,F,G), and 2 µm (L).
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Fig. 6. Two-dimensional Fourier power spectra of transmission electron micrographs
of structural colour producing butterfly scale nanostructures (Figs
4,
5). (A) Urania fulgens
blue (Fig. 4B), (B) Papilio
ulysses (Fig. 4E), (C)
Papilio zalmoxis (Fig.
4G), (D) Parides sesostris
(Fig. 4J), (E) Troides
brookiana (Fig. 4L), (F)
Troides urvillianus, (G) Callophrys dumetorum
(Fig. 5F), (H) Parrhasius
moctezuma (Fig. 5H) and
(I) Morpho aega (not illustrated). Colour scale (from blue to red)
indicates the relative magnitude of the squared Fourier components, which are
dimensionless quantities. Direction from the origin indicates the direction of
the 2D component waves in the image, and the distance from the origin
indicates the spatial frequency (cycles/nm) of each Fourier component. The
Fourier power peaks (red pixels) demonstrate predominant periodicities at
intermediate spatial frequencies. The distance from the origin is inversely
proportional to the wavelength of the coherently scattered colour.
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Fig. 7. Radial averages of two-dimensional Fourier power spectra from TEM
micrographs of structural colour producing butterfly scale nanostructures
(Fig. 6). (A) Urania
fulgens blue, (B) Parides sesostris, (C) Troides
brookiana, (D) Troides urvillianus, (E) Mitoura
gryneus. The shaded zone shows the range of spatial frequencies that are
likely to produce coherent scattering of visible light wavelengths.
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Fig. 8. Measured reflectance spectra (blue) and Fourier predicted reflectance
spectra (orange) for the structurally coloured butterfly nanostructures
illustrated in Fig. 4. (A)
Urania fulgens blue, (B) Urania fulgens green, (C)
Papilio ulysses, (D) Papilio zalmoxis, (E) Parides
sesostris, (F) Troides brookiana.
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Fig. 9. Measured reflectance spectra (blue) and Fourier predicted reflectance
spectra (orange) for the structurally coloured butterfly nanostructures
illustrated in Fig. 5. (A)
Troides urvillianus, (B) Troides priamus, (C) Celastrina
ladon, (D) Callophrys dumetorum, (E) Mitoura gryneus,
(F) Parrhasius moctezuma and (G) Morpho aega.
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Fig. 10. Fourier analysis of a concavity from a structurally coloured scale of
Papilio ulysses. (A) Transmission electron micrograph of a single
scale surface concavity from Papilio ulysses. (B) 2D Fourier power
spectrum of A showing an arc-shaped distribution of Fourier power peaks (red
pixels) above and below the origin that is created by the concave distortion
of the fundamentally laminar array of air bubbles in the scale. The result is
a broadening of the range angles over which the light back-scattered to the
observer will maintain the same peak hue.
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© The Company of Biologists Ltd 2006