J. R. Soc.

Interface (2009) 6, S165–S184

doi:10.1098/rsif.2008.0354.focus
Published online 28 October 2008

REVIEW

Gold bugs and beyond: a review of

iridescence and structural colour
mechanisms in beetles (Coleoptera)
Ainsley E. Seago1,*, Parrish Brady2, Jean-Pol Vigneron3
and Tom D. Schultz4
1
Department of Environmental Science, Policy, and Management, University of California,
Berkeley, CA 94720-3112, USA
2
Department of Physics, University of Texas at Austin, 1 University Station C1600, Austin,
TX 78712-0264, USA
3
Department of Physics, University of Namur (Notre-Dame de la Paix Namur),
Rue de Bruxelles 61, 5000 Namur, Belgium
4
Department of Biology, Denison University, Granville, OH 43023, USA

Members of the order Coleoptera are sometimes referred to as ‘living jewels’, in allusion to the
strikingly diverse array of iridescence mechanisms and optical effects that have arisen in
beetles. A number of novel and sophisticated reflectance mechanisms have been discovered in
recent years, including three-dimensional photonic crystals and quasi-ordered coherent
scattering arrays. However, the literature on beetle structural coloration is often redundant
and lacks synthesis, with little interchange between the entomological and optical research
communities. Here, an overview is provided for all iridescence mechanisms observed in
Coleoptera. Types of iridescence are illustrated and classified into three mechanistic groups:
multilayer reflectors, three-dimensional photonic crystals and diffraction gratings. Taxonomic
and phylogenetic distributions are provided, along with discussion of the putative functions
and evolutionary pathways by which iridescence has repeatedly arisen in beetles.

Keywords: iridescence; structural colors; Coleoptera; multilayer reflectors;

diffraction gratings; photonics

1. INTRODUCTION have been formally described (Erwin 1982; Lawrence &

Britton 1994). In addition to the striking iridescence,
Optical mechanisms in the natural world have fascin-
beetles have also evolved facultative luminescence,
ated researchers since the science of optics began;
Newton, Young, Michelson and Rayleigh all used their ultraviolet signals, polarized reflectance and complex
increased understanding of light rays to explain photonic crystals analogous to the fibre-optic tech-
iridescence in the feathers of birds and the scales of nology used to deliver high-speed Internet connections
butterflies and moths (Rayleigh 1930; Greenewalt et al. (Parker 1998; Galusha et al. 2008). Although iridescent
1960; Kinoshita et al. 2008). Butterflies have long been beetles have long been collected and used in human
the focus of iridescence research and industrial biomi- ornamentation, we still do not know what purpose
micry, probably due to their conspicuous structural iridescence may serve to the beetles themselves. This
colours, large body size and diurnal activity. However, question cannot be approached without a coherent
an equally important array of optical mechanisms grasp of both iridescence (as an optical phenomenon)
exists in another group of insects, the enormously and beetles (as organisms within an evolutionary context).
diverse order Coleoptera (figure 1). Beetles comprise an In recent years, researchers in both the physics and
estimated 5–30 million species, over 350 000 of which biology fields have discovered and characterized wholly
novel iridescence mechanisms in insects (e.g. Parker
et al. 2003; Prum et al. 2006). Unfortunately, these
*Author and address for correspondence: CSIRO Entomology, GPO
Box 1700, Canberra, Australian Capital Territory 2601, Australia.
disciplines have a bimodal distribution of interests and
([email protected]). rarely intersect: publications on beetle iridescence
One contribution of 13 to a Theme Supplement ‘Iridescence: more have appeared either in entomological literature
than meets the eye’. without optical context (e.g. Pope & Hinton 1977;

Received 19 August 2008

Accepted 6 October 2008 S165 This journal is q 2008 The Royal Society
S166 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

overview of beetle iridescence mechanisms, (ii) to

(a) (b) (c)
provide taxonomic, phylogenetic and functional context
to these optical phenomena, and (iii) to create a
robust, accurate and accessible vocabulary with
which to diagnose and discuss iridescence in beetles
and other insects.

1.1. Conventions
We adopt a definition of iridescence based on that of
Mason (1927): ‘iridescence has for its main characteristic
(d ) (e) (f) a change in the hue of the object exhibiting it as the angle
of vision is varied’. Although the ‘entomological’
definition of iridescence implies strictly spectral
(rainbow-like) reflectance ( Torre-Bueno 1989), the
term has been so widely and variously applied that a
broad interpretation is preferable in order to avoid
confusion. We have designated three main classes of
iridescence mechanism observable in beetles (multilayer
reflectors, three-dimensional photonic crystals and
diffraction gratings), and propose that these terms be
considered in future descriptive and experimental work.
Figure 1. Examples of beetle iridescence. (a) Loxandrus rectus Taxonomic groups are discussed in the narrowest
(Carabidae: Harpalinae), (b) Phalacridae gen. sp., (c) Cicindela sense applicable: we attempt to list the known
scutellaris scutellaris (Carabidae: Cicindelinae), (d ) Amaryg-
distribution of structural colour mechanisms as
minae gen. sp. (Tenebrionidae), (e) Phanaeus vindex
(Scarabaeidae: Phanainae), (f ) Eupholus sp. (Curculionidae: thoroughly as possible, but (space being limited) do
Entiminae). not list every species known or observed to possess a
particular mechanism. Names and classification used for
weevils (Curculionidae) follow Oberprieler et al. (2007);
Robertson et al. 2004; Doberski & Walmesley 2007) or
names for ruteline scarabs follow Hawks (2001).
in the physics literature without a strong organismal
We recognize that some of the anatomical terms
context (e.g. Parker 2000, 2002; Goldstein 2005;
used here may not be familiar to readers without an
Parker & Martini 2006; Jewell et al. 2007b; Vukusic
entomological background; however, we believe that
et al. 2007; Kinoshita et al. 2008; Liu et al. 2008).
incorporating the correct vocabulary is necessary to
The optical literature on beetle iridescence displays a lack
improve communication throughout a disparate litera-
of synthesis, with repeated ‘discovery’ or description of
ture. Thus, we present a brief review of relevant
virtually identical structures, misidentification of
structures and terminology in appendix B.
study organisms, outdated nomenclature and occasional
misunderstandings of insect anatomy (e.g. Hariyama
et al. 2002; Parker et al. 2003; De Silva et al. 2005; Hegedüs 2. MULTILAYER REFLECTORS
et al. 2006b; Jewell et al. 2007a,b; Liu et al. 2008).
The entomological literature on iridescence suffers 2.1. Mechanism
from both misunderstandings of optical mechanisms Multilayer reflectors are without question the most
and a severe terminological confusion. Although common and the best understood iridescence
methods and terms have been proposed for accurate mechanism in beetles (Parker et al. 1998; Noyes et al.
diagnosis of pigment colours (Paclt 1983; Aguiar 2005), 2007; Kinoshita et al. 2008). During the formation of
there is no consistent vocabulary for structural colours; insect integument, thin parallel layers of chitin (some-
‘iridescence’ is used to describe a number of dissimilar times interspersed with other materials) that differ in
phenomena, including metallic colours, spectral irides- refractive index are secreted by the epidermis and later
cence and opal-like effects. (As a point of reference, harden during sclerotization. If the spacing of these
the standard entomological vocabulary contains layers approaches one-quarter the wavelength of visible
approx. 30 terms for various shades of brown, but no light (approx. 380–750 nm), one or more colours will be
way to distinguish more than one type of iridescence produced by constructive interference (Land 1972;
(Smith 1909; Torre-Bueno 1989)). This long-standing figures 1c–e, 2–6).
confusion can be extremely detrimental when the word Contrary to the reports in some textbooks and
iridescence is used to connote a diagnostic character in articles (e.g. Daly et al. 1998), multilayer reflectors of
taxonomic works or treated as a single homologous beetles can be located at different layers within the
character state in cladistic analyses (e.g. Robertson integument (figure 2d– f ). Mason (1927) noted that
et al. 2004). beetle iridescence arose from different depths within the
We provide a much-needed comprehensive review of cuticle and could be removed from meloid, carabid and
known iridescence mechanisms in Coleoptera, synthe- buprestid beetles by scraping the (epicuticular) surface;
sizing the entomological and optical perspectives. Our transmission electron microscopy (TEM) later revealed
aims are threefold: (i) to provide a comprehensive multilayer reflectors in the outer 1–2 mm of the

J. R. Soc. Interface (2009)

 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al. S167

incident (white) light

(a) reflected (coloured light) (b) (c)

high index

low index

(d ) (e) (f)

Figure 2. Multilayer reflectors. (a) A schematic of simple cuticular multilayer reflector, (b) Cicindela scutellaris, TEM cross
section of cuticular reflector, (c) simple multilayer colour in a buprestid; schematic of (d ) epicuticular reflector, (e) exocuticular
reflector and ( f ) endocuticular reflector.

(a) (b)

(c) 2 µm (d) 1.0

0.9
0.8
relative reflectance

0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
400 450 500 550 600 650 700
wavelength (nm)

Figure 3. Additive coloration systems in Cicindelinae. (a) Cicindela repanda, habitus view, (b) Cicindela repanda elytral
surface with punctae, and (c) close-up view, SEM of elytron showing epicuticular reflector (arrow) and surface microsculpture,
(d ) reflectance spectra of punctae and surrounding areas (5 m2) of the elytron, the entire elytron and sand substrate
(T. D. Schultz 1994, unpublished data). Green triangles, puncta; yellow triangles, perimeter; red triangles, field; open circles, wet
sand; filled circles, elytron.

J. R. Soc. Interface (2009)

S168 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

(a) (b) (c)

Figure 4. Circularly polarized multilayer reflectors. (a) Schematic of helical multilayer reflector, (b) Chrysina boucardi viewed
through quarter wave plate rotated 08, and (c) C. boucardi viewed through quarter wave plate rotated 908 clockwise.

exoskeleton of buprestids and carabids (Durrer & (a) (b)

Villiger 1972; Mossakowski 1979, 1982). Using electron
microscopy and chemical analysis, Schultz & Rankin
(1985a) identified the reflectors of tiger beetles
(Cicindelinae) as non-chitinous epicuticle laminated
with ultra-thin layers of melanin. A similar epicuti-
cular reflector occurs in the iridescent chrysomelid
Plateumaris sericea (Kurachi et al. 2002). Layers of
chitin fibrils and protein in the exocuticle form the
interference reflectors of many scarabs ( Neville &
Caveney 1969), while the facultative, ‘switchable’
iridescence of some tortoise beetles (Chrysomelidae:
Cassidinae; see discussion in ‘Other mechanisms’
section) appears to arise from thin layers in the
endocuticle adjacent to the epidermis (Hinton 1973a). Figure 5. Broadband multilayer reflectors. (a) Schematic of
The colour reflected by a multilayer structure chirped reflector and (b) broadband reflectance in Chrysina
depends on the refractive index of the component chrysagyrea (Scarabaeidae: Rutelinae).
layers and their periodicity. Layers with a greater
optical thickness reflect longer wavelengths than
thinner layers and the peak wavelengthmax is equal to the pronotum and elytra will be blue shifted with
2 (nadaCnbdb), where n is the refractive index; d is the respect to the top of the segment (figure 2c). Deparis
actual layer thickness; and a and b are the alternating et al. (2008) have proposed a novel measure by which to
layers in the reflector (Land 1972). When reflector directly quantify iridescence in structurally coloured
thickness varies between body regions, multiple colours objects; this metric, termed ‘spectral richness’,
of different hue may be reflected (e.g. figure 1c,d ). describes the change in apparent wavelength as the
Polymorphism and geographical variations in metallic viewing angle departs from normal.
colour within species have been shown to result from
differences in the periodicity of reflecting layers
( Knisley & Schultz 1997; Kurachi et al. 2002). 2.2. Modifications
Estimates for the average refractive index of individual 2.2.1. Additive or ‘pointillistic’ colour mixing. An
reflecting layers in beetles range between 1.4 and 1.73 interesting modification of multilayer reflectors appears
(Caveney 1971; Parker et al. 1998; Kurachi et al. 2002; in tiger beetles (Carabidae: Cicindelinae). These pre-
Noyes et al. 2007). datory beetles are often found running on exposed soil
The apparent colour of a simple multilayer reflector or sand, where bright metallic colours would render
also varies with the angle of observation; given a them conspicuous to any natural enemy reliant on
constant angle of illumination, as the angle of obser- visual cues. In an apparent adaptation for camouflage,
vation increases (i.e. deviates from normal), the the elytra of many cicindeline species are covered with
reflected colour will undergo a ‘blue shift’ to a shorter an array of closely packed, minute dimples or punctae
wavelength, i.e. towards the blue end of the spectrum (figure 3a–c). Each dimple constructively reflects a
(see Vigneron et al. (2006) and Kinoshita et al. (2008) narrow band of wavelengths that differ from the
for a photographic illustration). The colour shift occurs surrounding field owing to differences in periodicity
because reflected light rays are travelling a shorter within the epicuticular reflector (Schultz & Rankin
distance through each layer, which means that con- 1985a). Generally, the reflecting layers within the
structive interference occurs for shorter wavelengths. punctae are thinner and reflect shorter wavelengths
This relationship between viewing angle and apparent than the layers around the perimeter. When viewed at a
hue results in a multicoloured appearance in convex distance, these ‘pixels’ of bright iridescent colour blend
beetles: when viewed from above, the lateral regions of to create a diffuse, matte brown, green or similarly

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 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al. S169

Circularly polarized light is also rare in nature;

among insects, it has been documented only among
scarabs and in the photic organs of certain lampyrid
larvae (Hegedüs et al. 2006b). In adult beetles, a
circularly polarizing reflector is formed when birefrin-
gent chitin layers are deposited helically (i.e. with
successive rotation), with a periodicity (one rotation of
the helix) equal to a wavelength of visible light
(figure 4a). According to Neville & Caveney (1969)
this ‘helical stack’ of chitin microfibrils is optically
analogous to a cholesteric liquid crystal. Under incident
unpolarized white light (i.e. sunlight), polarized
coloured light is reflected; the majority of green ruteline
scarabs reflect polarized light with a left-handed
(anticlockwise) rotation ( Neville & Caveney 1969;
Caveney 1971; Kattawar 1994).
Circular polarization appears in combination with
additive colour mixing in certain scarabs (Rutelinae:
Chrysina boucardi, Melolonthinae: Pyronota festiva;
De Silva et al. 2005; Jewell et al. 2007a,b). In these
beetles, polarizing multilayer reflectors take the shape
of closely packed, bowl-shaped ‘micromirrors’. Under
normal microscopy, these reflectors resemble the
Figure 6. Phanolinus sp. (Staphylinidae: Staphylininae) tiger beetle structures described above; however,
specimen exposed to sunlight (including UV ) on left side for the colours they reflect are circularly polarized, and
20C years. the surface of the dimples is sometimes covered by
a smooth, transparent wax layer (Chrysina, Pyronota).
In wax-covered reflectors, the overall chromatic effect is
unsaturated hue, often matching the colour of natural deeper in colour (e.g. C. boucardi, figure 4b), evocative
backgrounds (figure 3d; Schultz 1986; Schultz & of the ‘metal flake’ paint used on automobiles.
Bernard 1990; Acorn 1992). In some tiger beetles, a The human eye cannot readily discern circular
‘structural black’ is produced by the additive mixing polarization in ordinary light, but polarized colours in
of magenta and green iridescence. In addition to the beetles can be easily identified by viewing specimens
punctae, the surface microsculpture of the elytron is through a ‘circular analyser’, essentially a linear
composed of smaller hexagonal pits or alveoli, 10 mm in polarizer placed over a quarter wave plate (Kattawar
diameter (figure 3c) that enhance the matte-like 1994). This tool can be used to block out the polarized
reflectance. Owing to the effect of angle on interference green reflectance, while allowing the unpolarized
colour (see above), the walls of the alveoli reflect specular and pigment reflectance to pass through,
shorter wavelengths than the floor (Berthier 2007) which causes the normally virescent Chrysina to appear
contributing to the additive mixture of colours. Similar an ordinary ‘beetle brown’ (figure 4c).
colour mixing systems have recently been documented
in lagriine tenebrionids (Chlorophila; Liu et al. 2008)
and melolonthine and ruteline scarabs (see following 2.2.3. Broadband reflectors. A broadband reflector is
discussion of circular polarization). Unlike tiger beetles, any multilayer structure that reflects most or all
these mechanisms use identically bicoloured dimples wavelengths of light simultaneously, hence a ‘broad-
(e.g. Jewell et al. 2007b), instead of the ‘pointillism’ of band’ of the visible spectrum. A multilayer reflector
differently hued punctae. with many layers of different thicknesses (regardless of
the order in which they are arranged) will produce
broadband reflectance; however, only chirped broad-
band reflectors (those in which cuticle layers gradually
2.2.2. Circularly polarizing reflectors. Reflectance
increase or decrease in thickness; figure 5a) have been
of circularly polarized light by beetle cuticle was
reported in beetles (Parker 1998). The colour of broad-
first reported in beetles by Michelson (1911) in
band reflectors is also less directionally dependent,
Chrysina resplendens (see Goldstein (2005) for an
because the full range of wavelengths is reflected
in-depth analysis of reflectance and polarization prop-
(as opposed to the ‘narrow-band’ reflectance of chromatic
erties); this phenomenon appears to be restricted to
multilayers, wherein the apparent hue changes dramatic-
Scarabaeidae, occurring predominantly in Rutelinae,
ally with a change in viewing angle). The broader
Scarabaeinae and Cetoniinae.
the range of bandwidths, the closer to pure gold or
Whereas linearly polarized light is light in which the
silver (mirror-like) the cuticle appears (figure 5b).
electric vectors (e-vectors) are all propagating in the
same plane, the e-vectors in circularly polarized light all
propagate with the same clockwise or anticlockwise 2.2.4. Colour change in multilayers. The colours of
rotation. Unlike linearly polarized light, the detection beetles produced by multilayer reflectors may change
of circular polarization is not angle dependent. during development or as the result of environmental

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S170 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

conditions, which should be considered when examining stack, quarter wave stack, interference reflector and
and comparing the colours of specimens. After eclosion, dielectric mirror.
some beetles will exhibit a gradual increase in reflected We propose that the term multilayer reflector be
wavelengths as layers attain their maximal optical applied to such structures in Coleoptera; this describes
thickness; dehydration of the cuticle during this period the multilayered nature of cuticular chitin lamellae
may limit or reduce the wavelengths (Schultz & Rankin (which are not true films) and the reflective mechanism
1985b). In some beetle multilayer reflectors with porous by which colour is produced.
regions, air or fluid can be introduced in order to change The terms ‘metallic colours’ or ‘metallic iridescence’
the colour reflected (see §4, ‘Other structural colours’ can be used to distinguish multilayer effects from those
for an in-depth discussion of this phenomenon, which is produced by other optical structures. Multilayer
unique to polyphagan beetles). Permanent colour reflectance can typically be diagnosed as such by its
change in multilayers can be induced only under limited palette (usually one or two apparent hues per
extreme stress of sufficient force to alter the thickness reflector), blue shift with decreased observation angle
or refractive index of the cuticular chitin layers. Adachi and fixed position on the cuticle surface.
(2007) found that the metallic colour of buprestid
(Chrysochroa) elytra could be changed by heating the
specimen to 2008C or by soaking it in bromoform for 3. THREE-DIMENSIONAL PHOTONIC CRYSTALS
one month. A staphylinid specimen partially exposed (INCLUDING OPAL ANALOGUES)
to sunlight for 20C years lost the multilayer colours
on one side of its body, presumably due to long-term 3.1. Mechanism
exposure to UV (figure 6; S. Chatzimanolis 2008, Three-dimensional crystalline structures producing
personal communication). Under normal museum con- scintillating, gem-like reflectance were described by
ditions, most beetle multilayer reflectors retain their Parker et al. (2003) in the entimine weevil Metapocyr-
colours in perpetuity; the oldest known beetle reflector tus sp. (initially misidentified as Pachyrrhynchus argus);
is that described by Parker & McKenzie (2003), still by Welch et al. (2007) in Pachyrrhynchus congestus, and
reflecting metallic blue after an estimated 49 recently in another entimine weevil, Lamprocyphus
million years. augustus, by Galusha et al. (2008). The photonic
crystals found in the scales of pachyrrhynchine weevils
(Pachyrrhynchus and Metapocyrtus) have a close-packed
2.3. Taxonomic distribution
hexagonal arrangement analogous to (mineral) opal,
Multilayer colours occur in many families of adephagan while the photonic crystal of Lamprocyphus has a
and polyphagan beetles, and in too many species to count. diamond-based lattice (i.e. a face-centred cubic system
In the majority of beetle families where they occur, rather than a hexagonal one).
multilayer reflectors are restricted to a few lineages (e.g. Similar to the broadband multilayer reflectors, the
within Trogossitidae, Tenebrionidae, Silphidae, Meloi- three-dimensional photonic crystals found in beetles
dae, Staphylinidae); they are most commonly encoun- are an ‘iridescence-reducing’ colour mechanism: that is,
tered in the metallic wood-boring beetles (Buprestidae, they reflect vivid, saturated interference colours but
figure 2c), scarabs (Scarabaeidae, particularly Phanaei- reduce the angle dependency of the chromatic effect.
nae, figure 1e; and Rutelinae, figures 4b and 5b), ground Colour is produced by a highly ordered lattice of
beetles (Carabidae, figure 1c), leaf beetles (Chrysome- nanoscale spheres in the interior of flattened scales
lidae), some longhorn beetles (Cerambycidae) and the (figures 1f and 7a–d). The inverse of this structure,
weevil tribe Baridinae. In many families, there is a spherical lacunae in a chitin matrix, produces the same
distinct phylogenetic pattern to the distribution of optical effect; Galusha et al. (2008) have interpreted the
multilayer reflectors: some clades are almost entirely latter as ‘hexagonally ordered air cylinders’. Parker &
metallic, e.g. the scarab tribe Rutelini and the carabid Martini (2006) described the scales in Metapocyrtus as
subfamily Cicindelinae. filled with ‘transparent spheres, each 250 nm in
Additive colour mixing is widespread in the ground diameter . arranged in flat layers, [with] a precise,
beetle subfamilies Cicindelinae and Elaphrinae; colour hexagonal-close-packing order’. On a higher level of
mixing with polarized reflectors has been documented only organization, each scale contains a dense array of
in scarabs and tenebrionids, but may be more widespread. differently oriented micrometre-scale single-crystalline
‘domains’ (Galusha et al. 2008).
Although they are frequently compared with highly
2.4. Terminology and diagnosis
regular mineral structures, photonic crystals found in
Multilayer reflectors in beetles have also been described living organisms are not ‘perfect’ from a mathematical
as ‘thin-layer stacks’, ‘one-dimensional photonic crys- or engineering perspective. However, if we assume that
tals’ and ‘thin-film reflectors’ (e.g. Parker 1998, 2002; these optical mechanisms have been selected for, their
Vigneron et al. 2006). The vocabulary used to describe inherent imperfections may be considered to be part of
these structures is somewhat dispersive, as the var- an optimized diffuse reflection device. In weevils, the
iously intersecting disciplines of entomology, physics three-dimensional photonic structures are usually
and applied optics (e.g. laser technology, fibre-optic divided into an irregular assemblage of regular domains
data transmission, telescopes and microscopy) have all (e.g. figure 7a). Thus, the interior of the scale is better
developed slightly different suites of terminology. Other described as a photonic polycrystal than as a mono-
synonyms for ‘multilayer reflector’ include multilayer crystal: each grain in the polycrystal is cut from a

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 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al. S171

(a) (b)

(c) (d)

Figure 7. Three-dimensional photonic crystals in weevils and longhorn beetles. (a) Pachyrrhynchus congestus pavonius
(Curculionidae: Entiminae), SEM of crystal structure from scale interior (from Welch & Vigneron 2007), (b) SEM of the interior
structure of Prosopocerus lactator (Cerambycidae: Lamiinae), (c) Pachyrrhynchus gemmatus, light photograph of opalescent
scale patch, (d ) P. gemmatus, habitus view.

highly ordered photonic crystal (short-range order), 12 000 species (Anderson & Lanteri 2000). Entimine
but the orientation of the different grains varies across adults (many of which are flightless) feed on flowers or
the scale (long-range disorder). The size of the domains green foliage, which suggests that their coloration may
ranges from the full size of the scale (monocrystal) to be important in mimicry or crypsis.
the size of the photonic crystal period in the grain In cerambycids, the African lamiine species
(amorphous structure). These various levels of disorder Prosopocera lactator has slender, pearlescent scales
can be quantified by the grain size, i.e. the coherence with an internal ‘ball-and-stick’ crystal structure
length of the structure. The visual effects produced (figure 7b). Another African lamiine, Sternotomis
by structures with different coherence lengths include virescens (as well as some other Sternotomis), has
iridescence (e.g. figures 1f and 7c,d ), dull metallic elytral scale patterns with a greenish, opalescent
colours and whites. appearance, strongly reminiscent of the reflectance in
entimine weevils; the structure of the scale interior has
yet to be investigated for this genus.
3.2. Taxonomic distribution
Most members of the ‘higher weevils’ (Curculionidae
3.3. Terminology and diagnosis
sensu stricto, Brentidae and Attelabidae) bear pigmen-
ted or structurally coloured scales; species with Although the term ‘photonic crystal’ applies to any
opalescent green scales appear in most of the tribes of ordered subwavelength structure that affects the
Curculionidae, albeit most commonly in the subfamilies propagation of specific wavelengths of light (Parker &
Curculioninae and Entiminae. The most striking Townley 2007), it is the three-dimensionally ordered
iridescence occurs in various members of Entiminae, structures to which the term is most commonly applied.
the tribes Leptopiini and Pachyrrhynchini in particular. We recommend use of the term ‘three-dimensional
Entiminae is a large, tremendously diverse and taxono- photonic crystal’, which distinguishes these structures
mically problematic weevil clade, comprising over from the one-dimensional periodicity of multilayer

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S172 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

(a)
(b) (c)

0
1
2
3

Magn Det WD Exp 20 µm

normal 1500× GSE 11.3 0 0.3 Torr

Figure 8. Diffraction gratings. (a) Schematic of cuticular grating, (b) SEM of diffraction grating, Sphaeridiinae gen. sp.
(Hydrophilidae), and (c) Sphaeridiinae gen. sp., habitus view with zero, first, second and third spectral orders labelled.

(a) (b)

Magn Det WD Exp 20 µm

1500× GSE 11.9 0 0.3 Torr

(c) (d )

Figure 9. Modified diffraction gratings. (a) Pallodes sp. (Nitidulidae), SEM of bidirectional grating, (b) Pallodes sp. (Nitidulidae),
‘double’ spectral reflectance, (c) Aglyptinus tumerus (Leiodidae: Leiodinae), SEM of ‘interrupted’ grating, and (d ) A. tumerus,
habitus view.

reflectors or Bragg gratings. The terms ‘opal’ and Gibbs (1969a,b, 1971; see Hinton (1973a) for a complete
‘diamond based’ have been used to describe iridescence list of beetle taxa with cuticular diffraction gratings).
in weevil scales, but refer to phenomena that are They have recently been discovered in five additional
relatively similar from an organismal perspective; it is families, all within the suborder Polyphaga (A. E. Seago
important to note that these terms refer to crystalline 2008, unpublished data; e.g. figures 1a,b, 8a–c and 9a–d ).
lattice morphology and not the appearance of the scales A diffraction grating is any nanoscale array of
themselves. Maldovan & Thomas (2004) provided an parallel ridges or slits that disperses white light into
excellent overview of diamond-based lattice morphology its constituent wavelengths (figure 8a shows a grating
(as observed in Lamprocyphus) in photonic crystals; in cross section). Because white light consists of many
Yablonovitch (1993) provided a thorough introduction different wavelengths, it diffracts into full spectra,
to the photonic band-gap mechanism by which colours creating the rainbow-like reflectance shown in figures
are produced in three-dimensional photonic crystals. 1a,b, 8c and 9b,d. While man-made diffraction gratings
can disperse light via reflection or transmission, all
beetle gratings are strictly reflection mechanisms.
4. DIFFRACTION GRATINGS Diffraction gratings in beetles can be derived from
strigulose microsculpture, microtrichiae or modified
4.1. Mechanism setae (see Harris (1979) for useful basic classification of
Diffraction gratings were first described in a scarab surface microsculpture in insects). Iridescence arising
beetle by Anderson & Richards (1942) and in six other from diffraction gratings always takes the form of one or
beetle families by Hinton (1969, 1973b) and by Hinton & more ordered spectra—that is, colours are ordered in

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 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al. S173

the same sequence as the colours in the spectrum of independent origins of diffraction gratings, although
visible light, red–orange–yellow–green–blue–violet. explicit phylogenetic and evolutionary hypotheses are
This distinguishes diffraction colours from the unor- available only for Carabidae (e.g. Lindroth 1974; Allen &
dered ‘faux spectra’ produced in some beetles by Ball 1980).
variable thickness multilayer reflectors (figure 1d ).
The spectra produced by diffraction gratings are also
4.4. Terminology and diagnosis
‘ordered’ in a different sense: when compared with a
standard birefringence chart used in mineral analysis, Diffraction gratings give rise to iridescence in its
they follow the same sequence of spectral orders strictest sense, literally rainbow like (from the Greek
(labelled in figure 8), from zero-order (pure specular goddess Iris, associated with the rainbow). We use the
reflectance), first-order (saturated red/yellow/blue) to narrower term ‘spectral iridescence’ (in reference to
high-order spectra of secondary and tertiary colours. ordered spectra) to distinguish diffraction colours
Diffraction grating iridescence can easily be charac- from the metallic or jewel-like colours produced by
terized in terms of spectral order: for example, other structures. Diffraction gratings can be recognized
loxandrine carabids (figure 1a) display predominantly by the reflectance of one or more ordered spectra,
first-order spectra, while most phalacrid gratings running parallel or perpendicular to the longitudinal
(figure 1b) display long series of spectra, usually up to body axis and appearing to change position with the
the third order or higher. This difference is less likely to angle of observation.
be due to taxonomic differences and more likely to be
due to differences in grating morphology and curvature
of the elytron itself; basic literature on the effects of 5. OTHER STRUCTURAL COLOURS IN
convexity on diffraction gratings is sparse. COLEOPTERA
Although iridescence is the most familiar mode of
structural coloration in beetles, several other notable
4.2. Modifications
structural colour mechanisms exist in this group. The
Modification refers here to deviations from a ‘standard’ optical mechanisms discussed below produce colours
diffraction grating, not to evolutionary modification that do not vary significantly with viewing angle or give
from a shared ancestral morphology. Diffraction rise to spectral reflectance.
gratings in beetles occur in a wide variety of forms,
from weakly organized arrays of parallel ridges (e.g.
Aglyptinus, figure 9c,d ) to dense strigulae (figure 8b); 5.1. Whites and UV reflectance
see Hinton (1976a,b) and Ball & Shpeley (1983) for All whites in beetles are structural, arising from non-
additional examples of grating morphology, including ordered, broadband or Mie scattering of incident light
the ‘fringed’ gratings of sericine scarabs and the by nanoscale particles. White light may be scattered by
stretched sculpticells found in carabids, respectively. unpigmented cuticle, setae, scales and surface waxes.
Neotropical members of Nitidulidae (e.g. some The white maculations of tiger beetles arise from
Pallodes) have evolved a particularly curious grating broadband scattering by areas of the cuticle that lack
microsculpture not hitherto observed in nature: in these melanin (Schultz & Rankin 1985a). Vukusic et al.
species, iridescence on the pronotum arises from two (2007) described particularly bright white reflectance
intersecting, perpendicular diffraction gratings; this in the melolonthine scarab Cyphochilus; this whiteness
results in the reflectance of ordered spectra along both arises from fine (5 nm diameter) unordered cuticular
longitudinal and lateral axes, forming a spectral ‘halo’ filaments on the interior of scales, which in cross
around the point of specular reflectance (figure 9a,b). section form a slender but extremely effective array
of scattering structures. Several desert tenebrionid
species are white or pale blue due to scattering
4.3. Taxonomic distribution
from wax filaments that prevent water loss, reduce
Diffraction gratings are remarkably widespread through- radiation absorption and provide crypsis (Hadley 1979;
out Coleoptera; families in which gratings have been McClain et al. 1985).
observed include Torridincolidae (Ytu, Reichardtia, Iapir UV reflectance in beetles is a special case of ‘insect
and Hintonia), Carabidae (Loxandrus, Seleophrus, many white’ that also results from broadband scattering
Loxandrini and Oodini), Noteridae, Gyrinidae (Gyretes), (contra Pope & Hinton (1977), who wrongly attributed
Dytiscidae, Hydrophilidae (Sphaeridiinae), Scarabaeidae UV reflectance to pigments). Both white and UV
(Serica), Staphylinidae (Staphylininae and Scaphidii- wavelengths may be reflected by microtrichiae (Hinton
nae), Silphidae, Leiodidae (Aglyptinus and Agathidium), 1973b); however, species that appear similar in white
Mordellidae (Boatia), Phalacridae (many genera), Niti- light may produce dramatically different reflectance
dulidae (Pallodes), Bothrideridae (Ogmoderes, Prolyctus patterns in the ultraviolet. Pope & Hinton (1977)
and others) and Erotylidae (Mycotretus) (Hinton 1969; documented several cases of species that are super-
Hinton & Gibbs 1969a,b; N. Lord 2008, personal ficially similar in this way in Carabidae, Scarabaeidae,
communication; A. E. Seago 2008, unpublished data). Tenebrionidae, Cerambycidae, Curculionidae and
In several of these families diffraction gratings have a Chrysomelidae; such ‘UV-cryptic’ signals are relatively
polyphyletic distribution, i.e. are present in two or more widespread in insects (Silberglied 1979). Ultraviolet
non-sister lineages. The families Carabidae, Phalacridae vision (or at least sensitivity) is well documented in
and Staphylinidae appear to contain the most beetles; it is therefore plausible that UV reflectance

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S174 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

(a) (c)

Det WD 10 µm
GSE 9.7 0.3 Torr

(b)

L ×6.0k 10 µm

Figure 10. Quasi-coherent scattering. (a) Heikertingerella sp. (Chrysomelidae: Alticinae), SEM of scattering structure,
(b) Mychocerus sp. (Cerylonidae), SEM of scattering structure, and (c) Mychocerus sp. (Cerylonidae), habitus view showing
diffuse reflectance.

functions in visual recognition of conspecifics or mate Rassart et al. 2008). Hygrochromic colours were first
location ( Frantsevich et al. 1977; Dacke et al. 2002). noted in Dynastes hercules by Beebe (1947), and later
described in detail by Hinton & Jarman (1972) in the
same species. In D. hercules, the ‘resting state’ elytral
5.2. Quasi-ordered scattering coloration of greenish-grey changes to black when
Quasi-ordered two- or three-dimensional micro- hydrated; this change is purported to occur only in
structures can give rise to vivid, non-iridescent colours, males (Rassart et al. 2008) and can also be induced by
most frequently blues and greens. Colours are produced strong compression or mechanical stress to the elytra.
when identically sized nanoscale light-scattering The hygrochromic structure itself is a light-scattering,
objects (particles or lacunae) are evenly spaced in a weakly ordered three-dimensional lattice with air-
transparent or translucent matrix; the wavelength that filled porous regions. As moisture infiltrates the spaces,
will be constructively reflected is determined by the size the difference in refractive index between chitin
and spacing of the particles (Prum & Torres 2003). The and lacunae—and thus the amount of scattering/reflec-
semi-ordered nature of this phenomenon distinguishes tion—is decreased dramatically, more light is absorbed,
it from Tyndall or Rayleigh scattering, which arises and the elytron appears black in colour (Hinton &
from a completely unordered distribution of particles Jarman 1972; Hinton 1973a; Rassart et al. 2008).
(Prum & Torres 2003). This effect has been most Dramatic reversible colour change has also been
extensively documented in birds, Lepidoptera (Prum documented in the cassidine chrysomelids Aspidomorpha
et al. 2006, see also Prum & Torres 2003), Odonota tecta (Hinton (1976a), Charidotella egregia (Vigneron
(Prum et al. 2004) and Hemiptera (Miyamoto & et al. 2007), Deloyala and Metriona (ZCharidotella;
Kosaku 2002), but has not hitherto been described in Barrows 1979), all of which change from ‘resting’ gold
Coleoptera. Among beetles, quasi-ordered arrays of colour to red or reddish when disturbed. Vigneron et al.
surface tubercles (figure 10a,b) produce a blue or purple (2007) performed an extremely thorough study of the
sheen in some chrysomelids, scaphidiine staphylinids colour change mechanism in C. egregia, concluding that
and cerylonids (e.g. Mychocerus, figure 10c). The the ‘default’ gold colour of this species is produced in the
function of this mechanism in beetles is unknown; the exocuticle by a broadband reflector comprising high-index
faint, diffuse reflectance is unlikely to act as a strong chitin layers interspersed with irregular, porous patches.
visual signal. Unlike the colour-change mechanism in D. hercules
(in which reflectance is disrupted by the addition of
moisture to the exocuticle) Vigneron et al. (2007) found
5.3. Reversible colour change that red colour is exposed via dehydration of the porous
In a few species of beetles, reversible colour change is layers, thereby collapsing the broadband cuticular reflec-
effected by partially hydrating or dehydrating multi- tor into a ‘translucent stack’. In this configuration, the
layer structures, thus changing the refractive index of change in refractive index between adjacent layers is low
porous layers; this mechanism is also known as enough to permit most incident light to reach the pigment
‘hygrochromic colour’ (Mason 1929; Hinton 1973a,b; layer, where all but red wavelengths are absorbed.

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 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al. S175

Crowson (1981) noted that this phenomenon is anti-predator defence is provided by the observation that
widespread among cassidine chrysomelids. The func- cicindeline species often exhibit geographical variation in
tion of such a colour change (particularly in leaf- structural coloration that is correlated with the colour of
perching insects) seems likely to be aposematism or soils on which they occur (Schultz 1986; Hadley et al. 1988;
warning; however, as the transition takes place rather Pearson & Vogler 2001).
slowly, over at least 2 min (Hinton 1976a), it may be a
generalized response to disturbance rather than a
warning signal directed at a particular predator. 6.1.3. Aposematic (warning) colours. Although tiger
Non-reversible structural colour change co-occurs beetles are, as all carabids, chemically defended to some
with ageing in some donaciine chrysomelids, in which extent, they have also evolved a remarkable diversity of
adults undergo a permanent shift from metallic blue defensive coloration mechanisms from the same pre-
to metallic green over the course of several months sumably ancestral interference colours. Multilayer
(M. Barclay 2008, personal communication). The reflectors are used in aposematic colour signalling in
mechanism by which this occurs has not been investi- some cicindelines (Shelly & Pearson 1978; Pearson
gated, but is probably related to hydration or dehy- 1985). Acorn (1988) reported that multilayer colours in
dration of a cuticular multilayer reflector. some tiger beetles are used in the mimicry of sympatric,
chemically defended blister beetles (Meloidae) and
velvet wasps (Mutillidae). Vogler & Kelley (1998)
6. DISCUSSION analysed cicindeline colour pattern evolution in a
phylogenetic context, concluding that bright colours
Iridescence in Coleoptera clearly arises from a wide
in Cicindela appear to have two functions: either as an
variety of mechanisms. Although types of beetle
aposematic cue in strongly chemically defended beetles
iridescence can be classified into three major groups of
(subgenus Cicindelidia) or as a cryptics or disorienta-
analogous structures (multilayer reflectors, three-
tion colour in brightly iridescent but weakly defended
dimensional photonic crystals and diffraction gratings),
species (e.g. C. sexguttata). Iridescence may serve to
iridescence across taxa is neither identical in function
deceive visual predators when tiger beetles that appear
nor evolutionarily homologous. We now discuss the
bright in the Sun fly into shade and the colours are
demonstrated and theoretical functions of iridescence
extinguished (Knisley & Schultz 1997; Schultz 2001).
in Coleoptera, and discuss the hypothesized evolutionary
patterns and pathways by which it has most likely arisen.
6.1.4. Sexual signals. Although behavioural and ecologi-
cal observations are relatively scarce for Coleoptera,
6.1. Visual functions
several non-crypsis functions of multilayer colours have
6.1.1. Crypsis in foliage. The most common multilayer been posited. The following studies provide evidence for
colour in beetles is green, which is widely speculated to metallic colours as a visual signalling component in male–
function in crypsis (Crowson 1981; Parker 1998); this is male competition and mate location.
also the most common hue in coloured weevil scales, The pronotal shield of phanaeine scarabs (figure 1e)
which may derive their colours either from multilayer produces both bright red and bright ultraviolet reflec-
structures or crystalline arrays. Parker (1998) suggested tance when viewed head-on. Vulinec (1997) argued that
that multilayer reflectors coupled with a diffusing (light- this surface acts as a ‘backdrop’ emphasizing the size
scattering) surface texture could be a particularly of the horn in males, allowing competitors to assess
effective means of substrate matching in beetles, each other visually; it may also allow females to estimate
particularly those on substrates for which no insect the parasite load of males. Thery et al. (2008) have
pigment exists (e.g. green leaves; some flower surfaces). documented a similar system in Coprophanaeus lancifer,
In ruteline scarabs (see circular polarization section which has similar male horn morphology but subtler
below), vivid, saturated green hues are produced by pronotal coloration. Gwynne & Rentz (1983) documen-
helically arranged multilayer reflectors. Although these ted strongly vision-based mate location behaviour in
colours are highly conspicuous in a museum context, metallic gold buprestids; however, colour cues appear to
they may well be cryptic in a rainforest or cloud forest be less important than chemical cues in other buprestid
environment (Crowson 1981; Thomas et al. 2007). taxa for which mating habits are known (Alcock 1976).
Thomas et al. (2007) noted that Chrysina gloriosa, Kurachi et al. (2002) also noted that metallic colours
while vividly coloured in isolation, is one of a suite of are sexually dimorphic in some leaf beetles (Chrysome-
green-and-white-striped insects that are relatively lidae), and suggested that coloration plays a role in
cryptic in juniper foliage, their frequent natural perch. sexual signalling.

6.1.2. Substrate matching through additive colour. Most 6.1.5. Polarized signalling. Polarized reflectance may
tiger beetle (Cicindelinae) species show an inconspic- act as a receiver-dependent signal system, detectable
uous dorsal coloration due to the additive mixing of by polarization-sensitive conspecifics but invisible to
interference colours as described earlier (figure 3a,d). vertebrate predators. Experimental evidence indicates
These colours provide crypsis either by matching the that scarabs, as well as desert tenebrionids and many
general colour of the substrate or by mimicking the colour non-beetle insects, can detect polarized light in various
of small stones (Schultz 1986; Schultz & Bernard 1990; wavelength ranges (Dacke et al. 2002; Barta &
Knisley & Schultz 1997). Indirect evidence of this Horvath 2004). Hegedüs et al. (2006a) discovered that

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S176 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

melolonthine scarabs (Melolontha melolontha) are sensi- able to forage longer without overheating than the
tive to green polarized light, and Frantsevich et al. (1977) resident black morphs, which escaped the heat by
reported UV-polarized vision in the scarab genus Lethrus burrowing in the sand (Hadley et al. 1992).
(Geotrupidae). Dacke et al. (2002) found that the flightless
desert scarab Pachysoma striatum has particularly UV
and green polarization-sensitive ommatidia in the dorsal 6.2.2. Anti-adhesive/friction reduction. Hinton (1969,
region of the eye, and argue that a UV-polarized skylight is 1970, 1973a,b, 1976a,b) suggested that the spectral
used as an ‘optical compass’. Under laboratory conditions, iridescence arising from diffraction gratings serves
C. gloriosa with a choice of two polarized light sources a visual function, either as a warning colour or as
(circular and linear) preferentially moved towards the a facultatively conspicuous signal that helps to confuse
former (P. Brady & M. Cummings 2008, unpublished the depth-perception ability of predators. He also
data). Thus, while various scarabs do appear to use suggested that the usefulness of spectral iridescence as
polarization cues to navigate, the possible mate- a visual signal is contingent on availability of bright
recognition function of green polarized reflectance has direct illumination, and noted that ‘the highest
yet to be tested. percentage of beetles with diffraction gratings is found
in the sunniest parts of the world’ (Hinton 1973b);
however, beetle diversity (and insect diversity, in
6.1.6. Function of three-dimensional photonic crystals. general) is markedly higher near the equator, so the
Parker et al. (2003), Welch & Vigneron (2007) and apparently skewed distribution of diffraction gratings
Galusha et al. (2008) all emphasize the angle- may be an artefact of overall diversity.
independent nature of the iridescent reflectance in Some spectral iridescence is clearly a secondary result
weevils (a similar effect is produced in the hopliine of other functions. The abdominal plastron of many
scarab Hoplia, members of which have bright green torridincollids is brightly iridescent when illuminated;
and blue structural colours arising from multilayer similarly, stridulatory files (documented by Hinton
reflectors in thousands of minute, body-coating scales; (1969) in the hymenopteran family Mutillidae) are
Vigneron et al. 2005). In the case of weevils, while the often spectrally iridescent when exposed. The majority
‘multifaceted’ arrangement of crystal domains is of external diffraction gratings in Coleoptera occur on
important, it should be recognized that the distri- exposed regions of the dorsum and, more rarely, the
bution of a reflectance mechanism across thousands of entire body (Hinton 1973a); their function is not known.
scales also contributes to direction independence and However, it is notable that the beetle groups in which
could thus function effectively in crypsis among foliage diffraction gratings are most common share similar
or in producing a dispersive visual signal. It is also of microhabitats and thus similar mechanical challenges.
note that the multilayer reflectors that produce bright Nearly all adephagan and polyphagan beetle species
greens in other beetles (e.g. ruteline scarabs, many with diffraction gratings are associated with a moist,
buprestids) are almost entirely absent from weevils, sticky or colloidal substrate, e.g. mud-burrowing
occurring only in a few baridines and rarely in carabids (Oodinae (Spence 1982), swamp- or wet-
pachyrrhynchines. Weevils may therefore have forest-dwelling carabids (Loxandrinae (Ball & Shpeley
replaced elytral multilayer reflectors with scale- 1983; Ball 1985)), aquatic beetles (Dytiscidae, Gyrini-
based reflectors; the diversity of colours produced by dae and Noteridae) and small slime-mould and rotten-
the latter is certainly comparable to that of the former, fungus-feeding beetles, often from wet tropics or
and many scale patterns of ‘opal’ weevils seem closer cloud forests (Mordellidae, Nitidulidae; Phalacridae,
to warning coloration than to crypsis (e.g. Pachyr- Leiodidae, Staphylinidae: Scaphidiinae (Steiner 1984;
rhynchus gemmatus, figure 7d ). Franciscolo 1985; Leschen 1999; Seago & Wheeler
2004))). Interestingly, diffraction gratings are relatively
common in adephagan water beetles, but less so in
6.2. Non-visual functions
polyphagan water beetles: in Hydrophilidae, for
6.2.1. Thermoregulation. Any form of dorsal coloration example, diffraction gratings have only been observed
has consequences for thermoregulation in diurnal in the largely terrestrial and semiaquatic subfamily
beetles; these effects have been most thoroughly Sphaeridiinae. Members of this subfamily are found
investigated in tiger beetles. In a study of the in dung, rich humusy soil and moist decaying leaves
polymorphic Cicindela horni, Schultz & Hadley (Van Tassel 2001).
(1987) demonstrated that metallic green morphs Conspicuous iridescence arising from diffraction
attained the same body temperatures as black, pig- gratings is also common among burrowing snakes and
mented morphs under controlled conditions. Expanded lizards (Monroe & Monroe 1968; Arnold 2002). Gans &
white maculations significantly lowered the body Baic (1977) asserted that diffraction gratings in burrow-
temperatures of Cicindela formosa when compared ing snakes function in shedding mud or damp soil, but did
with morphs with reduced maculations. In populations not test this across many species nor in any experimental
of Neocicindela perhispida, the elytra of beetles that fashion. In carabid beetles, Erwin (1979) and Ball (1985)
occur on white beaches are entirely white while those on noted that diffraction gratings are prevalent among
black beaches have greatly reduced maculations and swamp-, marsh- and mud-inhabiting species, and rare in
appear black through the additive mixing of species from dry environments. The strong correlation
interference colours ( Hadley et al. 1988). When between presence of diffraction gratings and habitat type
transplanted to black beaches, the white morphs were (moist, concealed, largely nocturnal and crepuscular)

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 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al. S177

suggests that the primary function of this iridescence 6.3.3. Evolution of three-dimensional photonic crystals.
mechanism is of a friction-reducing or water-repelling Scales are a common feature in higher weevils, and
nature, and may aid locomotion in many species. often form conspicuous colour patterns that are likely
A second possible function of ‘blazed’ gratings (those to function as crypsis or visual signals. Although scales
in which the grating ridges are ratchet-like in cross seem to be a prerequisite for the evolution of photonic
section) is to aid forward movement in compressed crystals, lattices such as those discussed here have not
microhabitats. Carabids are known to engage in been observed in other scale-bearing beetle groups
‘wedge-pushing’ behaviour with enough force to move outside the clade Phytophaga (cf. blue and white scales
through the fibres of wood; Schmalfuss (1978) docu- in the scarabs Hoplia and Cyphochilus). This suggests
mented similar ‘terraces’ (essentially large-scale blazed that there may be some self-assembly mechanism
gratings) in burrowing crustaceans, and suggested that acting in weevils and cerambycid that is not present
these structures function in aiding movement through a in other beetles. Self-assembling opal-like crystal
dense substrate. Crowson (1981) noted that blazed lattices occur elsewhere in nature and have been
gratings are relatively common in beetles that move manufactured from a variety of compounds (Parker
through tight under-bark, rotting wood or packed soil et al. 2003; Parker & Martini 2006; Pursiainen et al.
conditions, and uncommon in species found in loosely 2007); those in weevils most likely arise from molecular
packed bark or leaf litter. self-assembly during scale formation.
The crystal lattices described by Galusha et al. (2008)
have laminar domains in certain directions and pre-
6.3. Evolution of structural colours dominantly reflect green light; most non-green structu-
6.3.1. Phylogenetic distribution of iridescence in rally coloured weevil scales are white, forming an
Coleoptera. A conservative estimate of phylogenetic important component of crypsis and shape disruption
distribution of the colour mechanisms described here is and creating false texture used to mimic rough bark, bird
given in figure 11. Again, we do not aim to list every droppings, etc. White reflectance in insects can only be
species or genus known to possess a particular colour produced by the scattering of light rays by an unordered
mechanism, but wish to indicate the breadth and particulate array (see ‘Other mechanisms’ section).
frequency with which different types of iridescence Thus, a plausible evolutionary pathway for three-
appear throughout the order Coleoptera. It is clear that dimensional photonic crystals could proceed along lines
iridescence mechanisms have independently evolved of increasing organization, from unordered particles
many times, with so great a frequency as to suggest (white reflectance) to two-dimensional ordered laminar
that they are favoured by selection in many circum- structures (green reflectance) to three-dimensional
stances. A number of hypotheses have been advanced ordered crystal lattices (opal-like reflectance). In this
as to how specific mechanisms have evolved. scenario, existing structural colours used in crypsis are
exapted to form the conspicuous reflectors of Pachyr-
rhynchus and other entimines; this hypothesis can be
6.3.2. Evolution of multilayer reflectors. Even by the tested using electron microscopy of scale interiors and a
most conservative estimate, there are many repeated tribe- or genus-level phylogeny for higher weevils.
evolutionary origins of multilayer reflectors in beetles.
The ‘armoured’ body plan of beetles is probably a key
innovation that subsequently allowed the group to 6.3.4. Evolution of diffraction gratings. Diffraction
radiate into the enormous diversity we see today. This gratings have evolved many times within Coleoptera
feature requires a many-layered cuticle thicker than that (‘a hundred times or more’ by Hinton’s (1973b)
of most other insects. Thus, it is possible that beetles estimate), and from many different morphological pre-
are preadapted for multilayer optics. As noted by Parker cursors. Diffracting structures have arisen from plastron
(1998), selection can act on colour by changing relative microtrichiae (Torridincolidae), stretched sculpticells
thicknesses of chitin or melanin layers in the cuticle, (Carabidae, Staphylinidae), what may be modified
rather than changing materials (i.e. pigments or tagma) setae or vestigial sculpticells (Leiodidae: Scotocryptini;
outright. The amount of selective pressure needed to Scarabaeidae: Sericini), and unknown microsculptural
drive evolutionary colour change may, therefore, be elements (Nitidulidae). The selective pressures that
lower for multilayer structures. Once a laminar structure favour the evolution of diffraction gratings are unlikely
exists in the cuticle, modification to a broadband to be related to visual function. Although ‘macrograt-
reflector is relatively straightforward, requiring enough ings’ such as strigae and dense recumbent pubescence
compression or attenuation during development to effect may provide a mild crypsis by reducing specular reflect-
an increase or decrease in the thickness of chitin layers. ance, gratings that are fine enough to produce iridescence
Similarly, circularly polarizing reflectance requires a have little effect on specular reflectance (figure 1a,b).
fairly simple modification of the deposition of cuticle, Interestingly, many groups that have diffraction
one which already exists in beetle larvae: Crowson gratings do not have multilayer metallic colours.
(1981, citing Neville 1970) reported that chitin layers in Rather, in these groups pigment colours predominate,
the cuticle of larvae are normally deposited in a chiefly the brown or black of melanin. Beetle lineages
helicoidal fashion, while the cuticle layers in adult with diffraction gratings have markedly similar eco-
beetles are not. Retention of larval cuticle-formation logical proclivities, with the majority of spectrally
patterns could be the mechanism by which helicoidal iridescent taxa living and feeding in concealed, moist,
multilayer reflectors in adult beetles have evolved. epigean habitats. Although their iridescent reflectance

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S178 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

Elateroidea

Elateridae in part

Dascilloidea Tenebrionoidea in part

Byrrhidae

Dryopoidea

Buprestidae Tenebrioninae
Meloidae
Bostrichoidea Tenebrionoidea in part
Lymexyloidea
Mordellidae
Dermestoidea
Coccinellidae
Staphylinidae in part
Cerylonid series
Bothrideridae
Scaphidiinae

Hydrophilidae Melyridae
Cleroidea
Lucanidae + Scarabaeidae in part
Trogossitidae
Dvnastinae + Rutelinae
Melolonthinae Erotylidae in part
Cetoniinae
Scarabaeoidea in part Erotylidae in part
Leiodidae
Nitidulidae
Staphylinoidea in part Cucujiforms including Phalacridae

Histeroidea
Chrysomelidae
Scirtoidea
Chrysomeloidae in part
Polyphaga
Dytiscidae Cerambycidae incl. Lamiinae

Noteridae Curculioninae
Gyrinidae
Adephaga Cicindelinae
Harpalinae
Carabinae Curculionoidea
Myxophaga + Carabidae in part
Archostemata Myxophage in part
Archostemata, Myxophaga in part
Sialidae
Figure 11. Approximate phylogenetic distributions of iridescence mechanisms in Coleoptera. Triangles represent diffraction
gratings; squares represent multilayer reflectors; circles represent three-dimensional photonic crystals. Tree simplified from that
presented by Hunt et al. (2007).

may not serve a visual function as that of leaf feeders, function for iridescence. Similar iridescence and colour
this optical mechanism is no less widespread and has by mechanisms are likely to serve different functions in
any estimate evolved many times within Polyphaga different taxa, not only crypsis but also sexual signal-
and Adephaga alike. ling, aposematism, mimicry, locomotory enhancement
and thermoregulation; some may simply be an artefact
of other structural constraints, such as the iridescence
7. CONCLUSIONS AND RECOMMENDATIONS
of the torridincollid plastron or the thin-film colour of
FOR FUTURE STUDIES
an exposed flight wing.
There is no single evolutionary pathway by which Multilayer reflectors are the most widespread
structural colour evolves in beetles, and no single iridescence mechanism, and have evolved many times

J. R. Soc. Interface (2009)

 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al. S179

Table 1. Relationships between structural colours, optical mechanisms and visible reflectance in Coleoptera.

structural colour mechanism wavelengths reflected

simple metallic hues multilayer reflector discrete band of visible spectrum

silver or gold colour variable thickness broadband multilayer reflector all visible
circularly polarized colour helically arranged multilayer reflector yellow–greena
spectral iridescence diffraction grating all visible, as ordered spectra
opal or diamond effects three-dimensional photonic crystal all visible
UV and white reflectance Tyndall scattering all visible, plus ultraviolet
most non-metallic blues quasi-ordered array typically blues and purplesa
a
Only these colours have been observed in nature; any colour is possible.

from similar precursors: stacked chitin layers (optically Because colour perception is subjective, varying
active or not) are present in all beetles, more so than between organisms, and even between human observers,
other, less well-armoured insects. In this sense, the and since we currently lack a fixed vocabulary for
order Coleoptera can be considered preadapted for this structural colours, it is particularly important to
particular form of iridescence. characterize chromatic features in the most objective
It is clear that multilayer coloration is asymmetri- fashion possible. The quickest and most objective
cally distributed across the beetle phylogeny: it occurs method is to measure reflectance peaks with a spectro-
throughout the order but is most common in the large radiometer (e.g. Endler 1990); this method is effective
clades Phytophaga, Scarabaeidae and Buprestidae. In with colours produced by multilayer reflectors and
adults of all three groups, surface feeding and daytime three-dimensional photonic crystals. Measured reflec-
activity on plant surfaces is common (Farrell 1998; tance spectra can then be used for precise character-
Bellamy & Nelson 2002). Although there are some ization of multilayer colour effects with or without
dazzling exceptions, the most common colour in these polarization (e.g. De Silva et al. 2005; Deparis et al.
groups is green, with apparent disruptive coloration, 2008). A less expensive alternative is that proposed by
aposematism and mimicry less widespread. Aguiar (2005), who proposed sampling a colour from
It is conceivable that, as various lineages of within a digital photograph of the specimen in question;
polyphagan beetles evolved to take full advantage of graphics programs such as Adobe Photoshop and
land plants (and in the process departing from their CorelDraw or Corel PhotoPaint automatically display
likely ancestral habits of concealed saprophagy ( Farrell the RGB coordinates of the sampled colour. This
1998; Marvaldi et al. 2002)) they were also subject to approach is more affordable than the former, but has
increased predation pressure from other invertebrates two notable limitations: unlike spectroradiometry, it
and vertebrates. If crypsis is the first (and arguably cannot detect UV and IR (nor quantify the amount of
metabolically cheapest) line of defence against preda- reflectance of a colour); it will also be affected by the
tion, and green pigments are rare or unavailable in light environment used to take the photograph.
animals, it can then be argued that multilayer reflectors When evaluating the proposed function of a color-
and photonic crystals are inextricably linked to major ation mechanism, it is also important to take into
evolutionary shifts towards plant feeding in polyphagan account not only the visual system of study organisms
beetles. As reflectors that produce green coloration in and their potential predators but also the organism’s
any exposed condition (i.e. when sunlight is present), natural context, including optical properties of the
structural colour mechanisms grant their bearers an substrate and the ambient light conditions (Endler
efficient and lasting camouflage among the green foliage 1990, 1993). This approach has been applied broadly to
of plants. the study colorations in vertebrates and butterflies, but
rarely with beetles. Schultz (2001) quantified the
conspicuousness of iridescent anti-predator colorations
of two tropical tiger beetles against natural back-
7.1. Recommendations for future research
grounds and under forest light. Using photoreceptor
When describing the coloration of an organism (in sensitivities obtained from the related species Onitis
ecological, behavioural and taxonomic works alike), we alexis, Thery et al. (2008) have determined that the
encourage the biologist to determine the underlying iridescent colour of the crepuscular scarab C. lancifer
colour mechanism(s); this is particularly important in would be most effective as an intraspecific signal at
iridescent taxa, where similar chromatic effects can be dusk when the beetles were most active. The visual
caused by very different and non-homologous system of O. alexis is dichromatic with receptor peaks
structures. In this review, we have aimed to provide in the UV and green (Warrant & McIntyre 1990); the
accurate descriptions, recognitory diagnoses and ancestral condition for beetles is trichromatic vision
known distribution of the major mechanisms of with peaks in UV, blue and green (Briscoe & Chittka
iridescence and structural colours in Coleoptera. We 2001, based on measurements for lampyrid beetles and
also provide a glossary of optical terms of potential use owlflies). However, more research is needed on the
to entomologists (appendix A) and a proposed termi- spectral sensitivities of beetles for which iridescent
nology of iridescence mechanisms (table 1). colours may serve as signals.

J. R. Soc. Interface (2009)

S180 Review. Iridescence mechanisms in Coleoptera A. E. Seago et al.

There is a long entomological tradition of consider- — Dielectric constant: essentially, the square of the
ing beetles and other insects within the context of the refractive index; also an electrical property of any
museum and laboratory, prizing large and seemingly insulating material.
conspicuous specimens for their jewel-like reflectance or — Diffraction grating: a series of parallel nanoscale ridges
unusual morphology. A similar habit persists in the that disperses light into ordered spectra.
fields of optics and biomechanics; individual structures — Iridescence: colour that changes in hue and intensity
are treated in isolation and evaluated on the basis of with change in viewing angle.
their material properties or as potential source material — Liquid crystal: substances with a phase of matter in
for the expanding technological field of biomimicry. between liquid and crystal; different LC phases are
Although a mechanistic, experimental approach is distinguished by their optical properties.
critical to elucidate how insect reflectors function, it is — Metallic iridescence or metallic colours: structural
important to connect these structures to their role in colours arising from a cuticular multilayer reflector.
the organism, which in turn necessitates a phylogenetic — Modes: wavelengths of EM that are allowed to
and ecological context. propagate through a photonic crystal.
The order Coleoptera is by any standard a prodi- — Nanoscale: at the same scale as wavelengths of visible
gious showcase for the extraordinary creativity and light, usually measured in nanometres.
flexibility of the evolutionary process. Many optical — Photonic crystal: any dielectric material with
mechanisms in beetles remain to be fully explored; with nanoscale refractive index periodicity affecting the
more diligent integration of the experimental and propagation of light waves; can be one-, two- or three-
evolutionary perspectives in research, we can achieve dimensional. In entomological literature, most com-
a fuller understanding of insect chroma. monly used to describe three-dimensionally ordered
lattice structures.
We thank the ASU Frontiers in Life Sciences programme and
— Refractive index: measure of a material’s ability to
the organizers of the conference ‘Iridescence: More than Meets
the Eye’, as well as A. Slipinksi (CSIRO Entomology, slow down (bend) a beam of transmitted light.
Canberra) and M. Brandley (University of California, — Spectral iridescence: reflectance of one or more
Berkeley) for their comments on an early draft of ordered spectra, the apparent position of which
the manuscript. changes with angle of observation; caused by diffrac-
tion grating.

APPENDIX A. GENERAL DEFINITIONS OF

OPTICAL TERMS USED IN THIS REVIEW
AND/OR COMMONLY ENCOUNTERED IN APPENDIX B. ANATOMICAL TERMS
IRIDESCENCE LITERATURE IMPORTANT IN LOCATING AND DESCRIBING
BEETLE IRIDESCENCE MECHANISMS
— Anisotropic: having direction-dependent optical
properties, i.e. transmitted light travels differently The most important distinguishing character of beetles
depending on its angle of incidence. is the presence of elytra, the hardened, armour-like
— Band gaps: wavelengths (band or range of wavelengths) forewings that protect the insect’s body. Most of the
that are not allowed to propagate through a medium. structural studies discussed here have focused on these
— Birefringence: ‘double refraction’; splitting of a single appendages, which are easily removed from dried
light ray into two rays when travelling through specimens for examination. The cross-sectional dia-
anisotropic medium (e.g. crystal or dielectric material). grams provided in figure 2d–f illustrate a section of the
— Bragg grating: can be a surface structure or a integument, the material of which the exoskeleton is
transmission structure. Diffracts light but with selec- composed. The integument comprises a soft, inner
tive reflectance; differs from transmission or reflection epidermis (nucleate cells shown at the bottom of
(diffraction) grating in that it does not disperse figure 2d– f ) and a hard, outer cuticle, which is typically
an ordered spectrum; unlike multilayer reflectors, divided into three layers, the inner endocuticle, the
Bragg gratings can propagate wavelengths for exocuticle and the waxy outer epicuticle. Beetle
long distances. cuticle derives its strength and flexibility from cross-
— Bragg wavelength: wavelength of light that is reflected linked fibres of the polysaccharide molecule chitin,
by a multilayer structure. which are deposited in a layered matrix exterior to
— Broadband reflectance: bright silver or gold appear- the epidermis.
ance caused by chirped or chaotic multilayer reflector
(sensu Parker 1998).
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