Materials Science and Engineering C 27 (2007) 941 – 946

www.elsevier.com/locate/msec

Living photonic crystals: Butterfly scales —

Nanostructure and optical properties
L.P. Biró a,⁎, K. Kertész a , Z. Vértesy a , G.I. Márk a , Zs. Bálint b , V. Lousse c , J.-P. Vigneron d
a
Research Institute for Technical Physics and Materials Science, H-1525, Budapest, POB 49, Hungary
b
Hungarian Natural History Museum, H-1088, Budapest, Baross utca 13, Hungary
c
Department of Electrical Engineering, Stanford University, Stanford, California 94305, California, USA and Facultes Universitaires Notre Dame de la Paix,
B-5000, Namur, Rue de Bruxelles 61, Belgium
d
Facultes Universitaires Notre Dame de la Paix, B-5000, Namur, Rue de Bruxelles 61, Belgium
Received 3 May 2006; received in revised form 22 September 2006; accepted 25 September 2006
Available online 15 November 2006

Abstract

The photonic crystal type nanostructures in the scales of male individuals of two butterfly species: Cyanophrys remus and Albulina metallica
were investigated by electron microscopy and reflectance measurements. While the colors of C. remus arise from structures with rigorous long
range (dorsal) or short range (ventral) three dimensional (3D) order, the colors of A. metallica are produced by quasi-ordered, layered structures.
Surprisingly, the most efficient photonic band gap reflector is the quasi-ordered structure giving the shiny, yellowish green color of the ventral
hind wings of A. metallica. All four investigated structures are based on a moderate refractive index contrast between chitin (n = 1.58) and air, the
various structures achieve a wide range of biological functions.
© 2006 Elsevier B.V. All rights reserved.

Keywords: Nanostructures; Butterfly wing scales; Photonic crystals; Electron microscopy; Reflectance

1. Introduction
Photonic crystals [1, 2]] may be regarded as a special case of
composite, built from two materials with refractive indices n1 and
n2, characterized by a refractive index invariant under the spatial
translations of a crystalline lattice, Fig. 1. The periodicity of the
structure has to be comparable with the wavelength range in
which it will exhibit the photonic crystal type behavior. In well-
defined frequency ranges, called “stop bands”, these materials,
also called photonic band gap materials (PBG), can prohibit
electromagnetic wave propagation along specific directions, even
with the very moderate refractive index contrasts found with
biological materials [3]. It should be emphasized, that for a
complete photonic gap, usually more complex structures are
needed than the structure shown in Fig. 1, which is only illus-
trating the principle of a three dimensional PBG.
PBG materials may have numerous applications ranging
from optical computing [4] through tunable photonic circuits
⁎ Corresponding author. Tel.: +36 1 3922681; fax: +36 1 3922226.
E-mail address: biro@mfa.kfki.hu (L.P. Biró).
0928-4931/$ - see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.msec.2006.09.043
[5], more efficient lasers [6] and fibers for optical communica-
tions [7], less harmful pigments [8] to photonic paper [9] to
mention only a few.
While the manufacturing of large area photonic crystals
operating in the visible spectrum is still a somewhat challenging
and expensive task for laboratory techniques, as the result of
extremely refined biological evolutionary paths, periodic di-
electric structures which operate in this wavelength range, have
evolved in living beings like certain butterflies, for example the
exclusively Neotropical butterfly genus Morpho [10–12], birds
[13], beetles [14], marine organisms [15] and even in plants
[16]. A more comprehensive overview was published recently
by Welch [17], the palaeontological aspects were reviewed
recently by Parker [18].
In the present paper the characteristic nanostructures in the
wing scales, generating the blue and green colors of the lycaenid
butterflies: Cyanophrys remus (Hewitson, 1868) and Albulina
metallica (Felder and Felder, 1865) (family Lycaenidae), will be
examined by transmission electron microscopy (TEM) and
scanning electron microscopy (SEM), the reflectance of the
investigated structures and their biological role will be
942 L.P. Biró et al. / Materials Science and Engineering C 27 (2007) 941–946

ing especially in the early morning hours, before the butterfly

Fig. 1. Schematic model of a 3D photonic crystal. As a rule of thumb the lattice
parameter a has to be of the same order of magnitude as the wavelength for which the
PBG will form. Note that for a full PBG in practical cases, usually more complicated
structures are needed than the simple cubic case used here for illustration.
discussed. Both butterfly species are colored by PBG
nanostructures on both the dorsal and ventral sides and the
nanostructures in the scales exhibit different degrees of order.
starts flying. It is interesting to examine more closely in which
way evolution, using the same two materials: air and chitin
(n = 1.58) produced several different structures, which all exhibit
PBG behavior and have very different reflectance and different
roles in the survival of the individual and of the species.
The nano- and micro-structure of the butterfly scales are
revealed by SEM and TEM as shown in Figs. 3 and 4. From the
examination of the electron micrographs one may conclude that
the four structures represent four different cases, in the sequence
of the decreasing order in the structure: i) the dorsal wing
surface scales of C. remus are characterized by well developed
ridges, which are filled by the same nanostructure as the body of
the scale, as shown by the TEM image, the structure has long
range order in the full three dimensions (3D), this observation is
quantitatively supported by detailed Fourier analysis of the
images, published elsewhere [20]; ii) the ventral wing surface
scales of the C. remus exhibit locally in the grains a highly
ordered structure, a so called face centered cubic (fcc) inverse
opal, but there is no long range order as the orientation of the
grains is random; iii) the dorsal wing surface scales of A.

2. Experimental results and discussion

The butterfly samples used in the present work were obtained

from the collections of the Hungarian Natural History Museum,
Budapest, Hungary. All the specimens investigated were males.
The species A. metallica belongs to the subfamily Polyomma-
tinae and inhabits high altitude open habitats in the Himalayas.
The species C. remus belongs to the subfamily Theclinae and
occurs in the tropical and subtropical forest zone of south-
east Brazil, where it lives in forest edges at low or moderate
elevations.
For SEM examination pieces of wings were attached to the
sample holders and covered by 20 nm of sputtered gold to avoid
charging. A LEO 1540XB FESEM/FIB was used for SEM
investigations. The TEM samples were prepared by incorpora-
tion of wing pieces in plastic blocks, followed by ultramicro-
tome sectioning which resulted in 70 nm thick slices. The
samples were examined in TECNAI 10 transmission electron
microscope. Reflectivity measurements were carried out with an
Avaspec 2048/2 fiber-optic spectrometer, both in specular ar-
rangement under normal incidence and using a 3 cm integrating
sphere in order to collect all the light reflected under any angle
of emergence. An Avaspec diffuse, white standard was used as
comparison sample for reflectance measurements.
As seen in Fig. 2a and b, both butterfly species have blue
dorsal color (with role in sexual signaling) [19], while their
ventral coloration, used as cryptic color, to help avoiding the
attacks of predators, is greenish. The ventral wing surfaces of
the C. remus have a matt pea green coloration, while only the
ventral hind wing surface of A. metallica is shiny greenish-
yellow. It is worth pointing out that C. remus lives in forests and
the matt pea green color is helpful in hiding the butterfly against
the diffuse green background of the vegetation. On the other Fig. 2. Composed photographs of male butterfly individuals representing C. remus
hand A. metallica lives in a dew-rich, grassy environment at (a) and Albulina metallica (both Lepidoptera: Lycaenidae) (b). Left side shows the
high altitude, the shiny greenish color is useful for camouflag- dorsal surface of the wings, while the right side the ventral side, respectively.
 L.P. Biró et al. / Materials Science and Engineering C 27 (2007) 941–946 943

Fig. 3. Electron micrographs of the scales on the wings of the C. remus: (a) dorsal scale SEM micrograph, note the well developed ridges; (b) dorsal scale TEM
micrograph, note the filled ridges and long range order in the structure; (c) ventral scale SEM micrographs, note the granular filling below the plane of the thin ridges;
(d) ventral scale TEM micrographs, note the local order within each grain and the random orientation of neighboring grains. All scale bars correspond to 5 μm.

metallica show a characteristic, quasi-ordered structure, which
is often called by the entomologists “pepper pot” structure [21];
iv) the ventral wing surface scales of A. metallica also have a
quasi-ordered pepper pot type structure, but characterized by
different dimensions as compared with the dorsal side. As seen
from the TEM images, in fact both the dorsal and ventral scales
of A. metallica exhibit a layered structure in which the thickness
of the chitin filled layers is different: 80 nm for the dorsal scales
and 60 nm for the ventral scales. Additionally, within the air/
chitin filled layers the effective refractive index depends on the
average size of the holes and the walls separating them, which is
again different for the dorsal and ventral scales. This quantity,
which we call filling factor, i.e., the ratio of hole diameter to the
width of the chitinous walls separating the holes is: 1.8 for the
dorsal scales and 2.2 for the ventral scales.
The normal incidence specular reflectance measurements
(Fig. 5) revealed that for each of the above structures a char-
acteristic reflectance peak can be found, Table 1. To allow the
comparison of the spectra measured, each spectrum was nor-
malized to its maximum in the range of 250–850 nm.
In fact, the spectral distribution of intensities in the light
reflected from the scales of butterflies built of chitin – a trans-
parent bio-polymer – is determined by the content of pigment: in
our case, brown melanin [3] (also a bio-polymer and a very
efficient absorber in the UV and blue range of the visible [22,
23]). A second factor is the structure itself when the scale
contains PBG material. Melanin alone would yield a continu-
ously increasing reflectance from 500 nm – the value of ab-
sorption edge determined from Fig. 1 of Ref. [22] – towards
800 nm, this would result in a brownish color. The PBG material
producing the structural, or “physical color” [24] will cause that
in a certain wavelength region, the incoming light cannot
penetrate into the scale volume. The light will be reflected by the
photonic crystal type structure (PBG), therefore it cannot be
absorbed by the melanin distributed in the volume of the scale. In
a scale containing only melanin or other pigments and no PBG
structure, the light may penetrate in the volume of the scale, due
to the absorption its intensity will be reduced according to the
absorption coefficient of melanin (other pigment). Therefore,
when illuminated with white light, it will appear brown in
reflected light. In the region of the PBG a reflectance peak will be
superimposed on the reflectance curve characteristic for the
melanin, i.e., when a PBG is present, in the wavelength region of
the photonic band gap, the reflectance will increase (from the
usually low value, characteristic for melanin) as the light is
reflected before being absorbed by the melanin. The combined
reflectance due to the melanin and the PBG will produce together
the blue or green coloration seen in Fig. 2 and the spectra in Fig.
5. As a consequence, the color will depend on the melanin
content, the efficiency of the PBG material in the scales – the
amplitude of the reflectance maximum – and on the bandwidth of
the photonic band gap – the full width at half maximum (FWHM)
of the peak – these factors together will decide the intensity and
the hue of the observed color. To facilitate the comparisons, all
spectra in Fig. 5 were normalized to the maximum measured in
the range of 250–850 nm, thus relative reflectances are given in
the figure. It is worth to point out that three of the four spectra,
although not all belong to the same butterfly species, show
relative reflectance of the order of 100% at 850 nm, while for the
ventral scales of A. metallica the relative reflectance is only 40%
944 L.P. Biró et al. / Materials Science and Engineering C 27 (2007) 941–946

Fig. 4. Electron micrographs of the scales on the wings of Albulina metallica: (a) dorsal scale SEM micrograph; (b) dorsal scale TEM micrograph; (c) ventral scale
SEM micrograph, note the similar but not identical structure with the SEM micrograph in (a); (d) ventral scale TEM micrograph, note similar but not identical structure
with the TEM micrograph in (b). The scale bars in (a) and (c) correspond to 2 μm; the scale bars in (b) and (d) correspond to 1 μm.

at this wavelength. This deviation from the general behavior is
associated with a strong and broad maximum at 545 nm with the
relative amplitude of 100%. This indicates that the ventral scales
of A. metallica contain an extremely efficient reflector, which is
in agreement with the observation by the naked eye, or the
photographic image in Fig. 2b.
Fig. 5. Relative reflectance data under normal incidence specular measurement
for the four butterfly wings. To facilitate comparison, each curve was normalized
to its maximum in the presentation window.
The two ordered structures, corresponding to the ventral (blue,
Fig. 3a and b) scales and to the dorsal (green, Fig. 3c and d) scales
of C. remus were modeled structurally and on the basis of the
models numerical computations successfully reproduced the
experimentally observed reflectance peaks. The model calcula-
tion yielded: a strong blue peak at 450 nm for the dorsal scales,
while the green peak is composed of weaker blue (469 nm), strong
green (498 nm) and strong yellow (550 nm) reflections occurring
on the (110), (100) and (111) faces of the fcc inverted opal [20].
The modeling of quasi-regular structures is a more difficult task,
the same is valid for the numerical computation of the reflectance.
On the other hand the experimental reflectance curves shown in
Fig. 5, indicate that the quasi-regular, layered structures – which
may be less difficult to manufacture than the structures based on
three dimensional long, or short range order – are more efficient
when normal incidence specular reflectance is measured. A
similar behavior is found when the integrating sphere is used, the
PBG material from the ventral scales of A. metallica is the most
efficient one, but the differences are reduced as compared with the
specular case Table 1. Another interesting feature is that in these
layered structures the refractive index contrast that could be
achieved by sharply separated chitin/air layers is reduced by the
“mixed layers” of air and chitin, still the reflectance achieved in
the case of ventral scales of A. metallica is rather strong and has a
large bandwidth. To check for possible directional effects reflec-
tance measurements were carried out using an integrating sphere,
the peak positions and FWHM values are given in Table 1. The
comparison on specular and integrated values in Table 1 shows,
 L.P. Biró et al. / Materials Science and Engineering C 27 (2007) 941–946 945

Table 1
Reflectance peak positions and intensities as measured in normal incidence specular geometry and with the integrating sphere for the two butterflies under
investigation, λnormal is the peak position for normal incidence, λintegrated is the peak position for integrated measurements, NRnormal and NRintegrated are the normalized
reflectance intensities (see text for details) for specular and integrated measurements respectively, FWHMn and FWHMi are the full width at half maximum values for
the peaks for normal incidence and integrated measurements respectively
Butterfly Side λnormal (nm) λintegrated (nm) NRnormal (%) NRintegrated (%) FWHMn (nm) FWHMi (nm)
C. remus Dorsal 480 460 55 93 110 106
Ventral 545 541 64 58 87 91
A. metallica Dorsal 450 – 81 – 152 a
415 – 77 – 152 a
416 – 92 133
Ventral 560 545 100 100 185 227
a
Due to the close overlap of the two peaks the FWHM value is given for the two peaks together.

that although the relative amplitude of the blue maximum cor-
responding for the dorsal scales of C. remus increased signifi-
cantly when measured with the integrating sphere – indicating
strong non-specular reflectance –, still its amplitude did not
increase over the amplitude of the green maximum of the A.
metallica. The blue maximum of C. remus has a narrow band-
width, while the green maximum of the A. metallica, has a very
large bandwidth which makes these structures suitable for dif-
ferent applications.
As regards the biologic function of these colors, the blue
color of the dorsal scales has a role in sexual signaling [19], both
in attracting the females and repelling other males. It is perhaps
more interesting here to concentrate on the very different,
cryptic green of the ventral scales. Two very different tasks were
achieved by appropriately modified PBG nanostructures built
from the very same materials chitin and air. The species C.
remus, living in forests or at forest edges evolved a matt green,
diffuse reflector built of randomly oriented photonic crystal
grains with fcc inverse opal structure, thanks to which it may
blend very efficiently in the diffuse green background of the
forest. The A. metallica, living in a high altitude, grassy habitat
with rich dew in the morning hours, developed a shiny cryptic
color, which may help it to blend efficiently in the glittering dew
covered grass. The production of these two greens with very
different aspect (matt versus shiny) clearly shows the great
flexibility of the natural “designs”, which can be used as tem-
plates or blueprints for artificial structures [14].
3. Conclusions
The photonic crystal type nanostructures built of chitin, oc-
curring in the butterfly wing scales of the male individuals
representing the species C. remus and A. metallica were inves-
tigated by electron microscopy and reflectance spectroscopy.
While C. remus has a dorsal metallic blue color arising from a
fully ordered 3D structure and a ventral mat green color pro-
duced by randomly oriented fcc inverse opal photonic crystal
grains, with perfect short range order, but lacking long range
order, the A. metallica has metallic blue color on its dorsal wings
and a very shiny goldish green color on the ventral side. Sur-
prisingly, both colors of A. metallica are achieved with similar
quasi-ordered layered structures with slightly different structural
parameters. The examination of four different PBG materials
built with the same two components: air and chitin, allowed the
comparison of the efficiency of different structures. The PGB
material found in the ventral scales of A. metallica is the most
efficient reflector from the four investigated ones. It is worth
pointing out that it is not a rigorously ordered structure and it
is based on a moderate refractive index contrast. Such quasi-
ordered nanostructures, which may be more easily manufactured
than the perfectly ordered 3D nanostructures could prove useful
in various practical applications. The investigation of natural
designs may yield very useful templates for artificial structures.
Acknowledgements
The work was supported by EU6 NEST/PATHFINDER/
BioPhot-01915. The work in Hungary was partly supported by
OTKA No. 042972. The work in Belgium was partly carried out
with support from EU5 Centre of Excellence ICAI-CT-2000–
70029 and from the Inter-University Attraction Pole (IUAP P5/1)
on “Quantum-size effects in nanostructured materials” of the
Belgian Office for Scientific, Technical, and Cultural Affairs.
Correspondence and requests for materials should be addressed to
László P. Biró, biro@mfa.kfki.hu.
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