[Palaeontology, Vol. 56, Part 3, 2013, pp.
557–575]
THE TAPHONOMY OF COLOUR IN FOSSIL INSECTS
AND FEATHERS
by MARIA E. MCNAMARA 1 , 2 , 3
1
Department of Geology & Geophysics, Yale University, New Haven, CT 06520, USA
2
UCD School of Geological Sciences, University College Dublin, Belfield, Dublin 4, Ireland
3
Current address: School of Earth Sciences, University of Bristol, Bristol, BS8 1RJ, UK; e-mail: maria.mcnamara@bristol.ac.uk
Typescript received 19 October 2012; accepted in revised form 5 March 2013
Abstract: Colouration is an important multifunctional
attribute of modern animals, but its evolutionary history is
poorly resolved, in part because of our limited ability to rec-
ognize and interpret fossil evidence of colour. Recent studies
on structural and pigmentary colours in fossil insects and
feathers have illuminated important aspects of the anatomy,
taphonomy, evolution and function of colour in these fossils.
An understanding of the taphonomic factors that control the
preservation of colour is key to assessing the fidelity with
which original colours are preserved and can constrain inter-
pretations of the visual appearance of fossil insects and
theropods. Various analytical approaches can identify ana-
tomical and chemical evidence of colour in fossils; experi-
mental taphonomic studies inform on how colour alters
C O L O U R is a phylogenetically important and multifunc-
tional attribute of extant animals (Hill and McGraw
2006), with important roles in inter- and intraspecific
signalling (e.g. sexual and social display (Parker 2000;
Bokony et al. 2003; Seago et al. 2009), camouflage (Riley
1997), UV protection (Butler et al. 2005), thermoregula-
tion (Riley 1997), sequestration of toxic metal ions
(McGraw 2003) and as a sink for free radicals (C
esarini
and INSERM 1996)). Visual evidence of colour associ-
ated with fossils can be conspicuous, for example metal-
lic colours in fossil insects (Fig. 1; Lutz 1990; Parker
and McKenzie 2003; Tanaka et al. 2010; Wedmann et al.
2010) and colour patterning in fossil molluscs (Hagdorn
and Sandy 1998), insects (Fig. 2; Wang et al. 2010) and
some feathers (Fig. 3; Vinther et al. 2008; Li et al. 2010;
Wogelius et al. 2011), and can yield taxonomic and
palaeoecological information (Blodgett et al. 1983; Turek
2009). Despite their palaeobiological potential, such ‘col-
oured’ fossils have, until recently, received little atten-
tion. The original colouration of most fossil organisms
thus remains speculative (Labandeira 2005; Li et al.
2010; Carney et al. 2012). A profusion of recent studies
on the colour of ancient insects and feathers, however,
has resulted in major advances in our understanding of
© The Palaeontological Association
during diagenesis. Preservation of colour is controlled by a
suite of factors, the most important of which relate to the
diagenetic history of the host sediment, that is, maximum
burial temperatures and fluid flow, and subsurface weather-
ing. Future studies focussing on key morphological and
chemical aspects of colour preservation relating to cuticular
pigments in insects and keratinous structures and nonmela-
nin pigments in feathers, for example, will resolve outstand-
ing questions regarding the taphonomy of colour and will
enhance our ability to infer original colouration and its
functions in fossil insects and theropods.
Key words: structural colour, multilayer reflectors, mela-
nin, pigments, taphonomy, fossil preservation.
fossil colouration and its functional evolution. Key
discoveries include fossilized melanin-bearing organelles
(melanosomes) in feathers (Fig. 4; Vinther et al. 2008)
and structural colours in fossil insects from many locali-
ties (Figs 1, 5; McNamara et al. 2012b). Recent studies
have also contributed novel methods for analysing fossil
colour (Figs 6, 7; Li et al. 2010; Wogelius et al. 2011;
McNamara et al. 2011, 2012a, b), provided insights into
the taphonomy of colour (McNamara et al. 2011, 2012a,
b, 2013a, b), constrained interpretations of enigmatic
anatomical features (Zhang et al. 2010) and allowed
inferences on behaviour and evolution (Clarke et al.
2010; Li et al. 2010, 2012; Zhang et al. 2010; McNamara
et al. 2011). Insects and feathers each have an important
fossil record: insects have been important members of
continental ecosystems since at least the Early Devonian
(Grimaldi and Engel 2005); feathers, a key derived avian
character (Bergmann et al. 2010), were diverse by the
late Cretaceous and may extend to the base of the
archosaur tree (Norell 2011). Evidence of original colour
in fossil examples of these taxa thus has the potential to
inform on the evolution of colour and its functions in
important fossil groups, and the role of colour, in par-
ticular visual signalling, in ancient ecosystems. Even
doi: 10.1111/pala.12044 557
558 PALAEONTOLOGY, VOLUME 56

A B C

D E F

FIG. 1. Metallic colours in fossil insects. A, leaf beetle (Coleoptera: Chrysomelidae) from the middle Eocene of Eckfeld, Germany.
NHMM PE1997 003a. B, weevil (Coleoptera: Curculionidae) from the middle Eocene of Messel, Germany. SMF MeI13011. C, jewel
beetle (Coleoptera: Buprestidae) from Messel showing patterning on thorax and elytra. SMF MeI14586. D, partial specimen of a jewel
beetle from the late Oligocene of Enspel, Germany. GDKE 1996 PE 1592. E, moth from Messel with, inset, reconstruction of original
wing colours (modified from McNamara et al. 2011). SMF MeI12269. F, detail of coprolite from Messel showing well-defined moth
scales. SMF MeI11808. Scale bars represent: A, 2 mm; B–E, 5 mm; F, 5 mm.

where evidence of colour is preserved in fossils, however, Interpretations of the colours and colour patterns of fos-
it may be the result of diagenetic alteration (Klug et al. sil organisms thus require an understanding of the
2009; Turek 2009; McNamara et al. 2011, 2012a, b). processes leading to their preservation. The purpose of

A B
FIG. 2. Monotonal colour pattern-
ing in fossil insects. A, leaf beetle
from the late Eocene of Florissant,
USA, showing symmetrical spotted
patterning on elytra. UCM 51311a.
B, unidentified bug (Hemiptera)
from the late Miocene of Oeningen,
Switzerland with, inset, detail of
area indicated showing submillime-
tre-scale spotted pattern. MCZ
14431. C, leaf beetle from the late
Eocene of Florissant, Colorado,
USA, showing banding on elytra
and abdomen. UCM 51324. D,
hindwing from the Neuropteran
C D
Limnogramma mongolicum (Kalli-
grammatidae) from the Jurassic of
Daohuguo, China, showing promi-
nent eyespot. NIGPAS NND11021.
Scale bars represent: A, C, 2 mm; B,
5 mm; D, 10 mm.
 MCNAMARA: FOSSIL COLOUR 559

A B C D

FIG. 3. Fossil feathers showing monotonal colour banding. A, feather from the Oliogocene Creede Formation, USA, showing proxi-
mal to distal gradation in tone. UCMP 169013. B, feather from Archaeopteryx lithographica (from Carney et al. 2012) showing proxi-
mal to distal gradation in tone. MfN MB.Av.100. C, feather from the Miocene Alvord Creek Formation, Oregon, USA, showing distal
to proximal gradation in tone. UCMP 391006. D, left forelimb of the troodontid Anchiornis huxleyi from the late Jurassic Taojishan
Formation, China, showing variation in tone across the wing. BMNHC PH828 (from Li et al. 2010). Scale bars represent: A, 2 mm; B,
5 mm; C, D, 10 mm.

this paper is to review recent developments in the study
of colour in fossil insects and feathers and summarize
how they have transformed our understanding of how
colour-generating mechanisms can be recognized in the
fossil record, how colours are modified by diagenetic
processes and how ‘coloured’ fossils can shed light on
the ecology and evolution of ancient insects and
theropods.
Light visible to humans is represented by a narrow
region of the electromagnetic spectrum characterized by
wavelengths between c. 380 and 750 nm (Farrant 1997).
Certain animals, including many insects and birds, are
sensitive to colours in the ultraviolet (10–400 nm)
regions of the spectrum (Shi and Yokoyama 2003); rare
insects (e.g. Melanophila (fire bugs; Vondran et al.
1995)) are sensitive to infrared radiation (750 nm–
300 lm). In addition, many insects are sensitive to
polarization (Land 1997). Colour-generating mechanisms
in the cuticle of extant insects and feathers fall into two
broad classes (with some overlap, see below) (Farrant
1997). Pigments are chemical compounds that are effi-
cient absorbers of specific wavelengths of light; light of
reflected wavelengths produces visible colour (Cromartie
1959). Structural colours are generated by constructive
inference when light is scattered in the visible part of
the electromagnetic spectrum by variations in tissue
nanostructure; such biophotonic nanostructures com-
prise materials of alternating high- and low-refractive
index and can be organized into one-, two- or three-
dimensional arrays (Prum and Torres 2003). Each of
these colour-generating mechanisms is discussed in
detail below.
Institutional abbreviations. BMNHC, Beijing Museum of Natu-
ral History, China; GDKE, Generaldirecktion Kulturelles Erbe,
Mainz, Germany; IVPP, Institute of Vertebrate Paleontology and
Paleoanthropology, Beijing; MCZ, Museum of Comparative
Zoology, Cambridge, MA, USA; MfN, Humboldt Museum f€ ur
Naturkunde, Berlin, Germany; NHMM, Naturhistorische
Museum Mainz, Germany; MNCN, Museo Nacional de Ciencias
Naturales, Madrid, Spain; NIGPAS, Nanjing Institute of Geology
and Paleontology, China; SMF, Forschungsinstitut und Natur-
museum Senckenberg, Frankfurt, Germany; UCM, University of
Colorado Museum of Natural History, Boulder, CO, USA;
UCMP, University College Berkeley Museum of Paleontology,
USA; YPM, Yale Peabody Museum of Natural History, New
Haven, CT, USA.
560 PALAEONTOLOGY, VOLUME 56

A B C D

E F G F

F I G . 4 . Scanning electron micrographs of modern (A–B) and fossil (C–H) feathers. A, eumelanosomes within a barbule of a satin
bowerbird (Ptilonorhynchus violaceus) feather. B, phaeomelanosomes within a barbule of a pigeon (Columba livia) feather. C, three-
dimensionally preserved eumelanosomes associated with feathers in a grebe (Podicipedidae) from the early Miocene of Libros, Spain.
MNCN-63817. D–F, feathers from the bird Confuciusornis from the Early Jurassic Jehol Group, China (modified from Zhang et al.
2010). IVPP-V13171. D, eumelanosomes preserved as external moulds. E, F, phaeomelanosomes preserved in three dimensions (E) and
as external moulds (F). G, rod-shaped feather degrading bacteria on a decaying contour feather from an extant finch (Taeniopygia gut-
tata). H, fossil feather from Paraprejica kelleri (Aves: Caprimulgiformes) from the middle Eocene of Messel, Germany, showing incom-
pletely degraded keratinous feather matrix. SMF MeV 1635a. All scale bars represent 2 lm.

GENERATION OF COLOUR IN INSECT
CUTICLE AND FEATHERS
Pigmentary colours
The mechanism by which pigments produce colour is
well understood. In most pigments, absorption of light
occurs in specific chemical regions (chromophores) that
comprise either conjugated p-systems (i.e. alternating sin-
gle and double bonds) or metal complexes (Shawkey and
Hill 2006). In each type of chromophore, empty electron
orbitals are available for electron excitation (due to shar-
ing of electrons between adjacent atoms); the energy dif-
ference between the ground- and excited state for a given
electron is equal to the frequency of light that is
absorbed.
Diverse pigments occur in insect cuticle and feathers.
Melanins, carotenoids and flavonoids are common in both
(Hill and McGraw 2006; Ghiradella 2009); pterins, ommo-
chromes, tetrapyrroles, bilins and quinones can also occur
in insect cuticle (Ghiradella 2009), and psittacofulvins and
porphyrins, in feathers (Hill and McGraw 2006). Melanins
are the most abundant and phylogenetically broadly dis-
tributed pigments in extant insects (Liu and Simon 2003)
and feathers (McGraw 2006; Ghiradella 2009; Stoddard
and Prum 2011), and they are the only class of pigments
in these taxa for which there is fossil evidence. Chemically,
melanins are large, inert polymers of dihydroxyindole and
dihydroxyindole carboxylic acid that crosslink strongly
with proteins (McGraw 2006); the precise structure is not
well understood (Li et al. 2012). In addition to their con-
tribution to inter- and intraspecific signalling, melanins
fulfil essential biological functions in metal scavenging,
radioprotection and photoprotection (Liu and Simon
2003; Liu et al. 2005); they also confer resistance to abra-
sion and immune attack, including bacterial degradation
(Goldstein et al. 2004; Nappi and Christensen 2005; Gun-
derson et al. 2008). In most insects, melanins are dissemi-
nated throughout the epi- and exocuticle and are
implicated in cuticle sclerotization (Andersen 2010). In
feathers, melanins occur within discrete membrane-bound
lysosome-related organelles termed melanosomes that are
embedded in the feather b-keratin matrix (Marks and
Seabra 2001; Fig. 4). Melanosomes are typically 470–
2000 nm long and vary in morphology according to the
chemical variant of melanin they contain: eumelanin
occurs in elongate, rod-shaped eumelanosomes (usually
900–1100 nm long; Fig. 4A), and phaeomelanin, in oblate
to spheroidal phaeomelanosomes (c. 470 nm long;
Fig. 4B). Eu- and phaeomelanosomes impart black, and
brown to rufous, tones in feathers, respectively; melano-
somes that impart grey tones are intermediate in morphol-
 MCNAMARA: FOSSIL COLOUR 561

A B C

D E F

FIG. 5. Biophotonic nanostructures in fossil insects. A–C, scanning (A) and transmission (B–C) electron micrographs of multilayer
reflectors in fossil beetles. A, fractured vertical section of cuticle from a jewel beetle (Coleoptera: Buprestidae) from the late Oligocene
of Enspel, Germany, showing laminated exocuticle (ex), epicuticular multilayer reflector (m) and vertical pore canals (arrow) traversing
exocuticle. GDKE ENS 2009 PE 5889. B, vertical section of cuticle from a leaf beetle from the middle Eocene of Messel, Germany,
showing laminated exocuticle (ex) and epicuticular multilayer reflector (m) characterized by alternating layers of high- and low-elec-
tron contrast. r, resin; s, sediment; t, trabecula. SMF MeI 15553. C, detail of epicuticular multilayer reflector (m) preserved in an
unidentified beetle from the early Miocene of Clarkia, USA. YPM 2010 P37b 005. D–G, scanning (D–E) and transmission (F–G) elec-
tron micrographs of multilayer reflectors from metallic moths from Messel; each of the nanostructures described below contribute to
the observed hue of the fossils. D, dorsal surface of cover scale showing longitudinal ridges flanked by short microribs and connected
by transverse cross-ribs. Internal laminae of the multilayer reflector are visible top left. E, dorsal surface of cover scale showing detail
of cross-ribs, microribs and perforations in the scale surface. F, vertical transverse section of two cover scales showing concave geome-
try of the multilayer reflector in between ridges (arrows). SMF MeI 11808 and MeI 12269. Scale bars represent: A–C, E–F, 1 lm; D,
5 lm, G, 500 nm.

ogy (Li et al. 2010). Melanosomes of different morphology
can occur within individual feathers (Li et al. 2010) and
can co-occur with other pigments (e.g. carotenoids; Lucas
and Stettenheim 1972; Hofmann et al. 2007). In Aves,
melanin-based plumage pigmentation (and its absence) is
evolutionarily primitive; carotenoids are considered to
have several independent evolutionary origins, and other
key pigments, for example psittacofulvins, unique origins
(Stoddard and Prum 2011).
Structural colours
Structural colours are the brightest and most intense
colours in nature (Vukusic and Sambles 2003; Parker
and Townley 2007) and are widespread in extant insects
and birds (Kinoshita and Yoshioka 2005). Such colours
function primarily in inter- and intraspecific visual
signalling (Parker 2000; Seago et al. 2009) and are usu-
ally associated with one or more striking optical effects,
for example iridescence (change in hue with observation
angle), opalescence and reflection that is metallic (highly
directional, near-100 per cent reflectance), ultraviolet or
polarized (Vukusic and Sambles 2003; Doucet and
Meadows 2009). Rare matte structural colours (with
probable functions in crypsis) are also known (Parker
et al. 1998a; Wilts et al. 2009). Biophotonic nanostruc-
tures in extant insects comprise three main classes: (1)
diffraction gratings, that is, arrays of parallel ridges or
slits, which usually generate weak spectral iridescence; (2)
two- or three-dimensional photonic crystals with hexago-
nal, cubic, diamond or gyroid lattices that generate a
spectrum of visual effects from dull matte colours to
bright opalescence; and (3) multilayer reflectors, that is,
alternating layers of high- and low-refractive index that
usually generate bright metallic iridescence (Parker et al.
562 PALAEONTOLOGY, VOLUME 56

cle (Neville and Caveney 1969) or endocuticle (Hinton

FIG. 6. Quadratic discriminant analysis of plumage colour
based on the morphology of feather melanosomes (from Li et al.
2010). Black, brown and grey dots represent data from extant
birds and numbers represent samples from the troodontid
Anchiornis huxleyi from the late Jurassic Taojishan Formation,
China. The geometry of melanosomes from the fossil feathers
clearly lies within the range exhibited in modern birds and, in
many samples, corresponds to distinct feather colours.
2001; Vukusic 2003; Seago et al. 2009). Multilayer reflec-
tors are the most common biophotonic nanostructure in
insects, where they can occur within the epicuticle
(Schultz and Rankin 1985; Kurachi et al. 2002), exocuti-
1973). Brighter colours are generated in multilayer reflec-
tors with more layers; mirror-like (near-100 per cent)
reflection is achieved where there are 10 or more high
refractive index layers (Land 1972). In lepidopteran
scales, colour-producing nanostructures in cover scales
are typically underlain by melanin-rich basal scales; the
melanin absorbs light transmitted through the biopho-
tonic array, thus preventing incoherent scattering and
enhancing the spectral purity of the observed colour
(Ghiradella 1998). The role of melanin in enhancing
structural colours in the cuticle of other insect groups is
unclear; in some beetles, melanin may be associated with
the high-index layers in multilayer reflectors (Schultz and
Rankin 1985) or may occur in a discrete layer underlying
multilayer reflectors (Parker et al. 1998a). The evolution-
ary origins of the various biophotonic nanostructures in
insects have been considered only for beetles (Seago et al.
2009) and some Lepidoptera (Tilley and Eliot 2002; Wilts
et al. 2009). Diffraction gratings and multilayer reflectors
are considered to have multiple evolutionary origins
(Seago et al. 2009); there is limited evidence that multi-
layer reflectors are evolutionarily primitive (Tilley and
Eliot 2002; Wilts et al. 2009), although this hypothesis is
controversial (Ingram and Parker 2008).
In feathers, structural colours originate via two distinct
mechanisms that are each localized to specific regions of

A B C

FIG. 7. Synchrotron rapid scanning X-ray fluorescence (SRS-XRF) mapping of the plumage of Confuciusornis sanctus from the Lower
Cretaceous Jehol Group, China (from Wogelius et al. 2011). A, optical image. IVPP MGSF 315B. B, SRS-XRF map of (A) showing
concentrations of Cu (red) in the plumage, Ca (blue) in the bones, and Zn (green) in the sedimentary matrix. C–E, SRS-XRF maps of
the region indicated in (A) for S (C), Ca (D) and Zn (E). Scale bar represents 20 mm.

the feather. Quasi-ordered three-dimensional nanostruc-
tures (with either a channel-type or microsphere architec-
ture) occur within the keratinous medulla of barb rami
(Shawkey et al. 2003; Shawkey and Hill 2006; Noh et al.
2010). Such keratin-air nanostructures generate bright
matte colours and are underlain by a thin layer of mela-
nosomes; as in lepidopteran scales (see above), the mela-
nin absorbs incoherently scattered light within the
structure and does not serve in colour production per se
(Shawkey and Hill 2006). Sheet-like arrays of melano-
somes within barbules act as thin-film reflectors and gen-
erate glossy black to highly iridescent colours (Doucet
et al. 2006; Yin et al. 2006; Yoshioka et al. 2007; Shawkey
et al. 2011); even slight organization can produce weakly
iridescent, glossy visual effects (Li et al. 2012). The precise
hue produced is a function of the thickness of the thin
(100–300 nm) keratin cortex that envelops the melano-
some layer(s) (Prum 2006). Quasi-ordered and thin-film
biophotonic nanostructures in birds are considered to
have multiple independent origins (Stoddard and Prum
2011).
FOSSIL EVIDENCE FOR COLOUR-
GENERATING MECHANISMS
Hand specimens and light microscopy
Fossil insects and feathers can manifest evidence for col-
our in hand specimen, for example metallic colours and/
or tonal (monochromatic) patterning. Metallic reflection
is evidence of structural colour (Parker et al. 1998a) and
is known in fossil insects from many Cenozoic localities
(Fig. 1; McNamara et al. 2012b and references therein).
Analysis of samples of cuticle from metallic fossil insects
using electron microscopy and mathematical modelling
confirms that the preserved colours in these specimens
are structural in origin (Parker and McKenzie 2003;
Tanaka et al. 2010; McNamara et al. 2012b). Barbules in
a single fossil feather from Messel (middle Eocene, Ger-
many) exhibit a silvery metallic sheen (Vinther et al.
2010); preserved ultrastructural evidence (see below)
strongly suggests that the feather is structurally coloured
(Vinther et al. 2010). The striking red-brown and violet
hues of the feather barbs and rami may reflect diage-
netic incorporation of metals, especially Fe, into feather
melanosomes (Vinther et al. 2010). Metallic hues may
also form during diagenesis of other organic tissues: fos-
sil graptolites associated with clay minerals can exhibit
blue colours (U. Farrell, pers. comm. 2012), possibly
due to light scattering from tectonically aligned clay
minerals. Whether or not preserved metallic colours are
structural in origin can be tested using electron micros-
copy and computer modelling (see below). Where the
MCNAMARA: FOSSIL COLOUR 563
colour-producing array is suspected to comprise a per-
meable array of air plus another material, however, an
expedient alternative is to immerse the fossil in media
of different refractive index; this alters the average
refractive index of the colour-producing array and thus
the visible hue (Mason 1927). This technique has been
successfully applied to metallic scales in fossil lepidopter-
ans from Messel, in which the nanostructure in the
scales comprises a chitin-air nanostructure. The tech-
nique could also be applied to metallic scales in other
fossil insect taxa (e.g. weevils) and in fossil feathers
where colour is generated by quasi-ordered nanostruc-
tures in the barbs.
Many fossil insects exhibit monochromatic patterning
expressed as varying tones of brown (Fig. 2) but the ori-
gins of this phenomenon have not been investigated. It
is conceivable that such patterning reflects original varia-
tions in cuticle thickness (Rasnitsyn and Quicke, 2002)
and/or pigment concentrations, in particular, eumelanin
(Vinther et al. 2008). Colour patterning can also occur
in fossil insects with metallic colours (e.g. Fig. 1C), but
has not been studied in detail. Chromatic variations in
metallic hue across a specimen presumably reflect varia-
tions in the thickness of the multilayer reflector; pattern-
ing consisting of alternating metallic and black regions
may reflect presence or absence of a multilayer reflector.
Interpretations of patterning in fossil insects are most
robust where specimens exhibit features that occur (and
have known visual functions) in extant insects, for exam-
ple symmetry systems (i.e. mirror images in paired tis-
sues such as insect wings), disruptive markings or
eyespots (Heads et al. 2005; Wang et al. 2010). Rare sub-
fossil and Miocene insect specimens exhibit patches of
dull red to yellow colouration when freshly exposed, but
these colours fade rapidly upon exposure to sunlight and
air; this process may reflect oxidation of partially
degraded carotenoid-based pigments (Rasnitsyn and
Quicke 2002).
Tonal variation is also known in fossil feathers, both
within individual feathers (Fig. 3A–C; Vinther et al. 2008;
Li et al. 2010; Barden et al. 2011; Wogelius et al. 2011;
Carney et al. 2012) and among feathers from an individ-
ual specimen (Fig. 3D; Li et al. 2010); this patterning can
relate, at least in part, to variation in melanin-based hues
(Vinther et al. 2008; Li et al. 2010, 2012; Barden et al.
2011; Wogelius et al. 2011; Carney et al. 2012). Patterning
within individual barbules of fossil feathers can closely
resemble melanin-based patterns in modern feathers and
can aid interpretation of feather microstructures (McKel-
lar et al. 2011). Survival of biological patterning can be
reasonably inferred in fossil feathers where the margins of
different colour bands match isochronic sections in pig-
mentary patterning in modern feathers (Vinther et al.
2008).
564 PALAEONTOLOGY, VOLUME 56
Fossils with metallic colours and colour patterns in
hand specimens are obvious targets for studies of fossil
colours, but monotonal fossils may also yield evidence
of colour. Indeed, many fossil feathers lack obvious
variations in tone but exhibit ultrastructural or chemi-
cal evidence of colour (see below). Further, recent taph-
onomic experiments using extant structurally coloured
insects demonstrate that ultrastructural evidence of
biophotonic nanostructures can endure the effects of
burial even where visible evidence of structural colour
is degraded (Fig. 8; McNamara et al. 2013a; see ‘The
fate of original colours’, below). Fossils that lack obvi-
ous colour thus have the potential to provide key
insights into the evolution of structural colour and
behaviour.
A B C
D E
F H
G ized remains of keratinophilic decay bacteria preserved in
F I G . 8 . Maturation experiments using the jewel beetle Chrys-
ochroa raja (Coleoptera: Buprestidae; modified from McNamara
et al. 2013a). A, untreated specimen. B–E, light micrographs of
cuticle from untreated (B) and experimentally decayed (C) spec-
imens, and specimens experimentally matured at 200°C, 500 bar
(D) and 270°C, 500 bar (E). F–G, transmission electron micro-
graphs of untreated (F) cuticles and cuticles matured at 270°C,
500 bar (G). H, reflectance spectra for cuticles in (A), (D) and
(E). Scale bars represent: A, 10 mm; B–E, 1 mm; F–G, 1 lm.
Electron microscopy
Electron microscopic imaging of preserved anatomical
detail is critical to identification of colour-producing
structures in fossil feathers (Fig. 4) and insects (Fig. 5).
Scanning electron microscopy (SEM) cannot distinguish
between multilayer reflectors and other laminated regions
of cuticle in extant and fossil insects (Fig. 5A). Transmis-
sion electron microscopy (TEM) of multilayer reflectors in
extant insects, however, reveals a characteristic ‘barcode’
pattern of alternating layers (each c. 100–150 nm thick) of
high- and low-electron contrast (Fig. 5B–C, F–G). The
high- and low-contrast layers are assumed to represent lay-
ers of high- and low-refractive index, but their precise
chemical composition and refractive index is notoriously
difficult to assess (Noyes et al. 2007). Identification of a
‘barcode’ pattern in TEM images of a laminated structure
in fossil insect cuticle is a strong indicator of a fossil mul-
tilayer reflector (Parker and McKenzie 2003; Tanaka et al.
2010; McNamara et al. 2011, 2012b), but unequivocal
determination of this requires mathematical modelling
(see below). SEM is also useful for determining the pres-
ence of other ultrastructures capable of producing or mod-
ifying colour. SEM imaging of structurally coloured scales
in fossil lepidopterans from Messel revealed evidence of a
fossil multilayer reflector and various colour-modifying ul-
trastructures (Fig. 5D–G); the latter produce visual effects
that are ecologically significant (McNamara et al. 2011).
SEM has the potential to reveal preserved remains of other
biophotonic nanostructures in fossil insects, for example
diffraction gratings and three-dimensional photonic crys-
tals. Diffraction gratings with a postulated anti-reflective
function are known from fossil flies hosted in amber (Par-
ker et al. 1998b; Tanaka et al. 2009) and have been
reported in arthropods from the Cambrian (Parker 1998),
but examples of three-dimensional photonic crystals have
not yet been reported in the fossil record.
Scanning electron microscopy imaging of many fossil
feathers reveals ovoid to rod-shaped microstructures that
can be closely spaced and aligned parallel to the barb or
barbule long axis (Fig. 4B–F; Vinther et al. 2008). Such
microstructures were originally interpreted as the fossil-
three dimensions via rapid replacement by authigenic
minerals (Wuttke 1983), but were reinterpreted recently
as the decay-resistant remains of feather melanosomes
(Vinther et al. 2008). Melanosomes and many bacteria
(e.g. Fig. 4G) are similar in gross morphology and size
(Davis and Briggs 1995; Vinther et al. 2008; Zhang et al.
2010; Knight et al. 2011); reinterpretation of the fossil
microstructures as preserved melanosomes is therefore
based on several arguments relating to their spatial orga-
nization and location within the feather. A melanosome
interpretation is supported where the fossil microstruc-

tures exhibit most or all of the following features: (1)
location within, or envelopment by, an organic matrix
(presumably the degraded remains of the feather keratin),
that is, the structures are internal to the feather and are
not external films of decay bacteria that grew on the
feather tissue during diagenesis (Zhang et al. 2010); (2)
preferential location, or at least high abundance, within
dark regions of fossil feathers (Vinther et al. 2008; Barden
et al. 2011); (3) dense packing, forming a discrete uni-
form surficial layer (e.g. Vinther et al. 2010) as in the
barbules of extant birds (Shawkey et al. 2006); and (4)
diagnostic nanoscale organizations, for example alignment
parallel to the barb long axis (Knight et al. 2011) that
cannot be generated by bacteria (Vinther et al. 2010; Li
et al. 2012). Despite increasing evidence for the preserva-
tion of melanosomes and melanin within theropods and
other fossil groups (Glass et al. 2012; Lindgren et al.
2012), interpretation of the fossil microstructures as pre-
served feather melanosomes (and thus survival of the pig-
ment melanin on geological timescales) is not universally
accepted on the basis of microstructure alone (Schweitzer
2011; Glass et al. 2012). Some authors have proposed that
the fossil microstructures represent melanosomes released
from the skin of the animals during decay (Lingham-
Soliar and Plodowski 2010) or the degraded remains of
structural collagen or keratin (Lingham-Soliar 2011).
Interpretations of fossil melanosomes may be more reli-
able where they are supported by chemical evidence of
melanin survival (Schweitzer 2011; Wogelius et al. 2011;
see below). Some fossil feathers retain three-dimensional
details of keratinous feather structures (e.g. Fig. 4H),
which are obvious targets for the recovery of nonmelanin
feather pigments and biophotonic nanostructures.
Modelling and statistics
Identification of biophotonic nanostructures in fossil
insects is contingent upon optical modelling of nanostruc-
tures within the cuticle. In extant insects, coherent scatter-
ing can be analysed using various techniques, in particular,
the matrix method of Macleod (1969) and applications of
the discrete 2D Fourier transform (Prum and Torres
2003). The matrix method is a powerful technique that
calculates the optical properties of laminar nanostructures;
average values for the thickness of each layer (measured
from TEM images), and estimates of each layer’s refractive
index, are used to generate a characteristic matrix for each
lamina (Macleod 1969). Matrices are then analysed using
purpose-built software, for example TFCalc (Software
Spectra, Inc., Portland, OR, USA). This approach is rou-
tinely used in thin-film optics and has been applied to bee-
tles from Messel (Parker and McKenzie 2003) and from the
Pleistocene of Japan (Tanaka et al. 2010). Predicted reflec-
MCNAMARA: FOSSIL COLOUR 565
tance spectra generated using this technique are compared
with observed reflectance spectra measured from the sur-
face of the fossil to assess whether the putative biophotonic
nanostructure in the fossil can produce the observed hue.
The 2D Fourier transform uses direct observations of
spatial variation in refractive index rather than idealized or
average values and can be applied to all types of biopho-
tonic nanostructure, not just laminar arrays (Prum and
Torres 2003); it has been used to analyse colour-producing
structures in extant insects, birds and mammals (Prum
and Torres 2003; Shawkey et al. 2009; Noh et al. 2010).
Digital TEM micrographs of cuticle are analysed using a
2D Fourier tool that is freely available (http://www.yale.
edu/eeb/prum/fourier.htm) and implemented in the
matrix algebra program MATLAB. The tool uses the distri-
bution of lighter and darker areas in the TEM image (and
therefore the distribution of materials of different refractive
index) to estimate the average refractive index of the struc-
ture. 2D fast Fourier transform analyses of spatial variation
in refractive index generate radial averages of Fourier
power spectra (useful for assessing whether a particular
structure can produce visible wavelengths) and predicted
reflectance spectra. This technique has been applied suc-
cessfully to cuticular nanostructures in fossil insects from
several localities (McNamara et al. 2011, 2012b).
Studies of the colouration of fossil feathers have used
statistical analysis of the morphology of fossil melano-
somes to predict colour within certain confidence inter-
vals (Clarke et al. 2010; Li et al. 2010, 2012; Carney et al.
2012). These analyses were based on a data set of melano-
somes from a phylogenetically diverse sample of extant
bird feathers with melanin pigmentation (Li et al. 2010,
2012; Clarke et al. 2010). The data set included parame-
ters such as long-axis, short-axis, long- and short-axis
skew, long- and short-axis variation, aspect ratio and
‘density’ (i.e. number of melanosomes per unit area).
Data on these aspects of the morphology and packing of
fossil melanosomes were compared with the data set of
modern samples using quadratic discriminant analysis, a
statistical technique that estimates the probability with
which unknown samples can be classified correctly using
data on known samples (Li et al. 2012). In each study,
forward stepwise analysis was used to determine which
melanosome parameters contributed significantly to the
analysis. Different combinations of parameters were sig-
nificant in different studies (aspect ratio and density in
the troodontid paravian Anchiornis (Fig. 6; Li et al.
2010); long-axis variation, short-axis skew, aspect ratio
and density in the fossil penguin Inkayacu (Clarke et al.
2010); and aspect ratio, long-axis, short-axis, long- and
short-axis variation, and aspect ratio skew in the paravian
Microraptor (Li et al. 2012)), but the importance of these
differences is unclear. The data from modern feathers
treated melanosomes from barb rami and barbules sepa-
566 PALAEONTOLOGY, VOLUME 56
rately (presumably to account for known intrafeather var-
iation in melanosome geometry; Prum 2006), but not all
analyses of fossil feathers made this distinction (e.g. Li
et al. 2010). The results of the statistical analyses were
used to predict the colours of fossil feathers from differ-
ent plumage regions with probabilities ranging from 56 to
100 per cent.
Chemistry
The chemistry of structurally coloured fossil insects is
incompletely resolved. Electron dispersive spectroscopy
(EDS) of structurally coloured cuticle in specimens from
Messel, Enspel (late Oligocene, Germany), Eckfeld (middle
Eocene, Germany), Clarkia (middle Miocene, USA) and
Randecker Maar (early Miocene, Germany) confirm that
the cuticle is invariably organically preserved (McNamara
et al. 2011, 2012b). The extent to which original biochemi-
cal components of the lipid-rich cuticle are preserved could
be determined using techniques such as pyrolysis–gas chro-
matography–mass spectrometry (py-GC-MS) and nuclear
magnetic resonance (NMR), which inform on macromo-
lecular complexes and proteinaceous moieties, respectively.
In contrast, several recent studies have investigated the
chemistry of fossil feathers (most of which are organically
preserved; Davis and Briggs 1995) using techniques that
provide insights into their elemental composition (EDS),
functional groups (Fourier transform infrared spectroscopy
(FTIR)), involatile macromolecular complexes (py-GC-
MS), organic free radicals (electron paramagnetic
resonance (EPR)), spatial distribution of elements
(synchrotron rapid scanning X-ray fluorescence (SRS-
XRF)) or local structure of specific metallic elements
(extended X-ray absorption fine structure (EXAFS) and
X-ray absorption near-edge structure (XANES)) (Barden
et al. 2011; Wogelius et al. 2011; Carney et al. 2012).
Combination of several such techniques is a powerful
approach to investigating the structural properties of
organic constituents, especially melanin and its degrada-
tion products, within fossils (Glass et al. 2012), and can
test interpretations of melanin fossilization based on
morphological evidence for preserved melanosomes
(Schweitzer 2011; Wogelius et al. 2011). EPR is a useful
technique for studies of eumelanin-based colouration in
fossils as eumelanin possess a unique free radical signature
(Glass et al. 2012). However, despite successful identifica-
tion of eumelanin and its derivatives in the ink sac of fossil
squid using this technique (Glass et al. 2012), application
of EPR to fossil feathers has met with only limited success;
analyses have demonstrated different free radical signatures
in fossil feathers and the surrounding sedimentary matrix,
but have not yielded diagnostic spectra for melanin
(Barden et al. 2011).
Recent SRS-XRF analyses of fossil feathers indicate that
certain trace elements (especially organic-bound Cu) in
fossil tissues rich in spheroidal and rod-shaped micro-
structures have melanin affinities and may act as biomar-
kers for melanin-derived compounds in fossils (Fig. 7;
Wogelius et al. 2011). This technique allows rapid, non-
destructive chemical mapping of entire fossil specimens
at concentrations in the ppm range (Wogelius et al.
2011). Some authors consider the resulting chemical data
superior to morphological data in studies of melanin-
based colour mechanisms in fossils as the effects of dia-
genesis on the geometry of melanosomes is poorly
resolved (Norell 2011; Wogelius et al. 2011; but see
McNamara et al. 2013b, and ‘The fate of original col-
ours’, below). Other analytical techniques that could be
applied to studies of fossil melanin in theropods include
ToF-SIMS, which allows simultaneous identification and
mapping of molecules and their structures at high spatial
resolution (Lindgren et al. 2012). Recent studies of fossil
squid and fish using ToF-SIMS confirm that chemical
evidence of melanin is restricted to micron-sized melano-
some-like bodies in tissues where melanin was probably a
major constituent in life (Glass et al. 2012; Lindgren
et al. 2012).
THE FATE OF ORIGINAL COLOURS
Morphological evidence of colour
Insects. The taphonomy of colour-producing nanostruc-
tures in insects is reasonably well understood. Multi-
layer reflectors in Cenozoic insects have a similar
preservation potential to other cuticular nanostructures
(McNamara et al. 2012b). The suite of ultrastructural
features preserved in fossil cuticles is therefore key to
assessing whether black colours are a taphonomic arte-
fact; the preservation of diverse cuticular ultrastructures,
but not biophotonic structures, indicates that biopho-
tonic nanostructures were originally absent. This does
not necessarily imply, however, that the cuticle was
black in vivo; nonmelanin pigments may have been
present originally, but visual evidence thereof is not
preserved.
Despite widespread preservation of multilayer reflectors
in metallic fossil beetles, original hues are not preserved
(McNamara et al. 2011, 2012b); observed reflectance
spectra of the fossils are redshifted from spectra pre-
dicted using the preserved biophotonic nanostructure.
This phenomenon was attributed to alteration of the
refractive index of the cuticle; changes in periodicity can
also effect colour change in multilayer reflectors (Adachi
2007), but cuticular features in the fossils lack clear evi-
dence of volume change, for example buckling, pull-apart

structures and desiccation cracks (McNamara et al.
2012b). In contrast to these results, high-pressure/high-
temperature maturation experiments using extant struc-
turally coloured beetles resulted in a blueshift in
observed hue (Fig. 8; McNamara et al. 2013a). The
experiments used the extant jewel beetle Chrysochroa
raja, which generates metallic colour using an epicuticu-
lar multilayer reflector; specimens were matured for
24 hours using various pressure–temperature regimes
(117 bar, 200°C; 250 bar, 200°C; 500 bar, 200°C; and
500 bar, 270°C). The hue of the beetles changed progres-
sively (decreasing wavelength) with increasing pressure.
This change resulted from alteration of both the refrac-
tive index and periodicity of the multilayer reflector; the
dimensions of the reflector and of various other cuticular
structures were altered without obvious distortion. The
colour change had two discrete components: a large
blueshift caused by a decrease in periodicity of the multi-
layer reflector, partly offset by a smaller redshift relating
to considerable alteration of the chemistry of the epicuti-
cle and, in turn, an increase in its refractive index. The
redshift is identical in magnitude and direction to the
discrepancy in wavelength between observed and pre-
dicted data for the fossil beetles, supporting the hypothe-
sis that the fossil redshift results from a change in
refractive index. The chemistry of structurally coloured
fossil insects has yet to be investigated comprehensively
(but see Parker and McKenzie 2003; Tanaka et al. 2010),
but it is clear that both chemical and morphological data
are critical to future attempts to reconstruct original
structural colours in fossil insects.
As with fossil beetle colours, the hues produced by
multilayer reflectors in fossil lepidopterans also alter dur-
ing diagenesis (McNamara et al. 2011); unlike the fossil
beetles, however, predicted colours for the lepidopterans
are blueshifted from preserved hues. This difference could
plausibly relate to differences in the chemistry of the bio-
photonic tissues in each taxon: in extant insects, beetle
epicuticle comprises lipid and protein (i.e. chitin is
absent; Neville 1975), whereas lepidopteran scales com-
prise predominantly chitin (Powell 2003). Regardless of
the extent to which original colours have been altered,
however, certain aspects of the visual ecology of structur-
ally coloured insects may be inferred using anatomical
evidence preserved in fossils. For instance, the colour-
producing nanostructures in the fossil moths described
above are associated with other ultrastructural features in
the scales that modify the visual signal (the inherent
iridescence and specular reflection of the multilayer
reflector are suppressed), implying a defensive function
for the colour (McNamara et al. 2011).
All fossil examples of structurally coloured insects
contain multilayer reflectors. This may be a function of
the relative abundance of different colour-producing
MCNAMARA: FOSSIL COLOUR 567
mechanisms in extant insects: multilayer reflectors are the
most common biophotonic nanostructure in animals
(Parker 2002), including beetles (Seago et al. 2009). Alter-
natively, the high abundance of fossil multilayer reflectors
may be taphonomic in origin. All published examples of
fossil multilayer reflectors are hosted within organic-rich
lacustrine sediments with significant volcaniclastic input
(McNamara et al. 2012a); pore waters from such sedi-
ments would be expected to have a slightly acidic pH.
Epicuticular lipids are insoluble in acidic media, and thus,
the composition and chemistry of host sediments may
influence the preservation potential of multilayer reflec-
tors. Maturation experiments on 3D photonic crystals
show that they have similar preservation potential to mul-
tilayer reflectors. The absence of 3D photonic crystals in
the fossil record (at least in biotas of Miocene age and
older) is thus considered to represent a real, evolutionary
absence (McNamara et al. 2013a). Critically, maturation
experiments also demonstrate that physical evidence of
colour-producing nanostructures survives in insects even
where visual evidence of colour is lost (McNamara et al.
2013a). Structural colour may thus have an extensive
cryptic fossil record in insect specimens that lack obvious
metallic colouration.
Feathers. Despite intense interest in the colour of fossil
feathers, the taphonomy of colour-producing mechanisms
in feathers has not been a focus of investigation. None-
theless, it is clear that the fidelity of preservation of fossil
feathers, and their melanosomes, varies considerably. Fos-
sil melanosomes can be preserved as three-dimensional
bodies (Fig. 4C, E) or as external moulds embedded in
amorphous organic material or diagenetic minerals
(Fig. 4D, F; Clarke et al. 2010; Li et al. 2010, 2012; Zhang
et al. 2010); both preservational modes can occur within
a single feather (Zhang et al. 2010). The nature of the
organic matrix surrounding some mouldic melanosomes
may represent degraded feather keratin (Zhang et al.
2010) or melanin (Clarke et al. 2010; Li et al. 2010). Fur-
ther, dark visual tones in fossil feathers can (Li et al.
2010), but do not always (Li et al. 2012), correspond to a
high abundance of melanosomes and do not correlate
with the mode of melanosome preservation (Li et al.
2010). Dark tones (i.e. a high absorbance of visible light)
commonly originate in organometal- or conjugated bonds
in modern pigments (Farrant 1997). The precise chemical
structure, and taphonomy, of the chromophore in fossil
feathers is, as yet, uncertain.
In structurally coloured fossil feathers, reconstructions
of original hue are precluded by degradation of the kera-
tin cortex that envelops the melanosomes in vivo; this
cortex is responsible for the exact hue produced by the
highly ordered melanosome array (Vinther et al. 2010; Li
et al. 2012). In other fossil feather examples, however
568 PALAEONTOLOGY, VOLUME 56
(i.e. those lacking structural colour), accurate predictions
of precise hue are contingent upon the geometry of mel-
ansomes (Li et al. 2010). Reconstructions of original
plumage colouration in fossil theropods have assumed
that the original geometry of melanosomes is preserved
in the fossils (Clarke et al. 2010; Li et al. 2010, 2012;
Knight et al. 2011). Fossil melanosomes, however, vary
in the mode of preservation: melanosomes preserved as
moulds and three-dimensional bodies from the same
feather region differ in size (Clarke et al. 2010) and yield
differing colour predictions (Li et al. 2010, 2012). Matu-
ration experiments on feathers from extant birds reveal
that melanosome geometry is altered by the effects of
elevated pressure and temperature (McNamara et al.
2013b). These experiments used melanosome-bearing
feathers from 12 extant taxa; feathers encompassed
diverse hues and melanosome types (eu- and phaeomela-
nosomes, solid and hollow melanosomes). The experi-
ments used two different pressure–temperature regimes
(200°C, 250 bar; 250 bar, 250°C) and lasted 24 hours;
melanosomes in all feathers altered progressively in
geometry (both long and short axes reduced in length)
between the 200°C, 250 bar experiment and that using
250°C, 250 bar. Survival of original melanosome geome-
tries in fossils is thus most likely where the host sedi-
ments experienced limited burial. Not all feather-
containing fossil deposits, however, meet this criterion
(McNamara et al. 2013b). Some studies of melanin-based
colouration in fossil feathers have considered that contri-
butions by other pigmentary and structural colouration
mechanisms to the visible hue in vivo would have been
masked by melanin (Carney et al. 2012; Li et al. 2012),
but this is likely only in feather regions with very abun-
dant melanosomes. Feathers in many extant birds con-
tain melanosomes but the visible hue derives from
nonmelanin pigments or biophotonic architectures
(McNamara et al. 2013b and references therein). Matura-
tion experiments on such feathers reveal that melano-
somes are retained in degraded feathers even where
visual evidence of all other colouration mechanisms has
degraded completely. Given this preferential preservation
of melanosomes, attempts to reconstruct colour in fossil
feathers should be integrated with anatomical and geo-
chemical data on the preservation of other pigments and
biophotonic structures.
Chemistry
Traces of original cuticular biomolecules, for example
chitin and amino acids, are preserved in subfossil bee-
tles with epicuticular multilayer reflectors (Tanaka et al.
2010). Less is known about the chemical fidelity with
which older structurally coloured insects are preserved.
Some authors have suggested that original organic
material has survived in metallic beetles from the mid-
dle Eocene of Messel, but this hypothesis is supported
only by bulk elemental analysis (Parker and McKenzie
2003). Where fossil cuticles contain an epicuticular
multilayer reflector, lipid extracts may be a useful proxy
for the chemistry of the epicuticle and thus of the col-
our-producing structure. Lipid extracts of thermally
matured cuticle from extant beetles analysed via py-GC-
MS are dominated by lipid–protein complexes. These
complexes are absent in fresh cuticle and represent
reaction products of functionalized epicuticular lipids
with proteinaceous moieties from the epi- or exocuticle
(McNamara et al. 2013a). The composition of structur-
ally coloured fossil cuticles has yet to be investigated
using techniques that inform on the structure of pre-
served components.
Unequivocal traces of melanin have been reported in
fossil squid (Glass et al. 2012) and fish (Lindgren et al.
2012) but have not been identified in fossil feathers (Bar-
den et al. 2011). Recent synchrotron-aided analyses using
X-ray fluorescence (XRF) show that distribution maps of
certain trace elements with a melanin affinity, for example
Cu, in fossil feathers may help reconstruct colour patterns
in fossil plumage (Wogelius et al. 2011). EXAFS and
XANES analyses demonstrate that Cu is present in fossils
in organometallic form, possibly derived from original
melanin (Wogelius et al. 2011). The spatial distribution
of trace elements in fossil specimens, however, may also
result (at least in part) from taphonomic modification of
original colouration signals. Trace metal concentrations
could increase in tissues as a result of adsorption by
microorganisms during decay (Hitchcock et al. 2009) or
chelation during later diagenesis (Shock and Koretsky
1993). Studies of melanin in fossil fish suggest that it may
be concentrated in the dark regions of fossils during dia-
genesis due to preferential degradation of more labile
organic molecules (Lindgren et al. 2012). This process
does not, however, preclude diagenetic migration of mela-
nin from source tissues. Diagnostic biomarkers for mela-
nin were recovered from sediment adjacent to the ink
sacs of fossil squid (Glass et al. 2012), although this may
reflect minor leakage of ink and/or intact melanosomes
from the ink sac during decay rather than later diagenesis.
Trace elements within fossil feathers could also derive
from endogenous sources other than melanin, for exam-
ple keratin–Cu complexes in feathers (Wogelius et al.
2011)), or from external, that is, sedimentary, sources.
Preservation of intact chemical moieties of the melanin
molecule may result from thermally induced polymeriza-
tion reactions during diagenesis (Glass et al. 2012). In situ
polymerization may also explain, at least in part, the ali-
phatic composition of some fossil feathers (Barden et al.
2011).

CONTROLS ON THE PRESERVATION
OF COLOUR
Patterns in the fossil record of structural colour in insects
relate to a hierarchy of taphonomic controls, including
decay, burial pressure, burial temperature, the nature of
diagenetic fluids, weathering and the mode of curation of
fossil specimens (McNamara et al. 2012a). These factors
are also likely to govern the preservation of colour in fos-
sil feathers, although the relative importance of different
taphonomic factors in preserving feather and insect col-
our may differ. Statistical analyses of the fidelity of the
preservation of structural colours in various insect taxa
from Cenozoic biotas revealed that age and taxonomic
compositions do not control taphonomy; age may, how-
ever, be more important in older material as it serves as a
proxy for more extensive or complex diagenesis.
Decay
Laboratory decay experiments on extant structurally col-
oured beetles at room temperatures and pressures did
not induce colour change (Fig. 8; McNamara et al.
2013a). Some fossil insect cuticles in which metallic col-
ours are poorly preserved or absent, however, exhibit evi-
dence for decay by fossil or modern microbes
(McNamara et al. 2012a); the latter have a particularly
negative impact on the fidelity of preservation. Decay
may also affect colours in fossil feathers, especially where
these are structurally coloured: the keratin cortex and
medulla (which house various colour-producing nano-
structures) are less resistant to decay than melanin (Gold-
stein et al. 2004). The high preservation potential of
melanin (Hollingworth and Barker 1991) does not, how-
ever, imply that it (or melanosomes) is immune to alter-
ation during decay. Laboratory decay experiments on
extant birds showed that decay-induced rupture of feath-
ers can liberate melanosomes from the keratin matrix,
obliterating original packing arrangements (McNamara
unpub. data).
Burial depth
The maximum depth to which fossil insects and feathers
are buried during diagenesis is an important control on
the preservation of colour. It determines the maximum
pressures and temperatures to which most fossils are
exposed and influences the fidelity of preservation of ana-
tomical features (McNamara et al. 2012a, 2013a); it may
also influence biomolecular preservation (H€ oss et al.
1996). Burial depth contributes to broad-scale inter-biota
patterns in the fossil record of structural colour in insects.
MCNAMARA: FOSSIL COLOUR 569
Structurally coloured fossils can be abundant in biotas
that experienced limited burial, for example Messel, Ens-
pel, Eckfeld, but absent in biotas buried to greater depths,
for example Green River (McNamara et al. 2012a). Matu-
ration experiments confirm that increasing pressure alters
insect structural colours, but the effect of pressure is sec-
ondary to that of temperature, which is the primary agent
of colour change during burial (McNamara et al. 2013a).
Structural colours alter progressively with increasing tem-
perature but are lost beyond a temperature threshold; the
value of this threshold temperature is likely to vary with
the chemistry of the tissue and host sediment. Feathers
experimentally treated to elevated pressures and tempera-
tures lose visual and ultrastructural evidence of all
colour-producing mechanisms save for melanosomes;
progressive loss of anatomical detail occurs in tandem
with loss of visual colour with increasing temperature
(McNamara et al. 2013b). As with insects, temperature
may be the primary determinant of colour (i.e. melano-
some) survival in fossils.
Diagenetic fluids and weathering
Other factors that influence biota-scale patterns in the fossil
record of colour (at least for structurally coloured fossil
insects) include the nature of diagenetic fluid flow and the
extent of weathering (McNamara et al. 2012a). Reactive
hydrothermal fluids can degrade insect cuticle and lead to
precipitation of authigenic minerals within colour-produc-
ing nanostructures, resulting in loss of visible colour
(McNamara et al. 2012a). Extensive oxidative degradation
of cuticles during weathering may also destroy structural
colours. The absence of structurally coloured insects in the
late Eocene of Florissant (USA), and their high abundance
in other Cenozoic biotas (i.e. Messel, Clarkia (early Mio-
cene, USA), Enspel and Eckfeld), has been attributed to
variations in the extent of Recent weathering among these
biotas (McNamara et al. 2012a). Host sediments at Floris-
sant are exposed subaerially, but those from the other local-
ities are located beneath the water table; sediments from
Messel comprise approximately 40 per cent water (Schaal
and Ziegler 1992). Variations in the extent of weathering
can also explain patterns in the fidelity of preservation of
colour within an individual biota. Specimens from Eckfeld
(middle Eocene, Germany) show a statistically significant
correlation between poor preservation of structural colours
and preservation in extensively weathered host sediment
(McNamara et al. 2012a).
Deep burial, exposure to diagenetic fluids, and subaer-
ial or subsurface weathering may also be responsible for
certain taphonomic features in fossil feathers. Melano-
somes are frequently preserved as external moulds, yet the
origin of this phenomenon is unclear. Melanin is resistant
570 PALAEONTOLOGY, VOLUME 56
to microbial and chemical attack (Goldstein et al. 2004),
but heat treatment can induce chemical changes in its
molecular structure, rendering the molecule soluble in
strongly oxidizing fluids and various acids and bases (Fox
1976). Degradation of three-dimensional fossil melano-
somes could therefore result from the effects of elevated
temperatures and reactive pore fluids during diagenesis,
or from oxidative weathering during exhumation and
exposure. Indeed, the last of these factors is considered to
be an important agent of colour loss in other pigmented
fossils (Hagdorn and Sandy 1998).
Mode of curation
The mode of curation of structurally coloured fossils can
affect also the long-term stability of the colour-producing
mechanism and the resulting hue after collection. Metallic
colours in insect specimens can degrade following dehy-
dration in air (Parker and McKenzie 2003; Schweizer
et al. 2006). Loss of colour via dehydration and microbial
degradation (Toporski et al. 2002) is usually, but not
always (McNamara et al. 2012a), prevented by storing
such specimens in liquid media, for example brine, etha-
nol or glycerine. Indeed, loss of nanostructural definition
and visible colour can occur after only several months’
storage in brine (McNamara et al. 2012a).
FUNCTIONAL AND EVOLUTIONARY
SIGNIFICANCE OF FOSSIL COLOURS
Identification of evidence of colour (and accurate recon-
structions of original colours) in fossils can inform on the
function of colour, especially visual signalling mecha-
nisms. Iridescent metallic colours in modern insects can
be cryptic in foliage but conspicuous in direct sunlight
and thus may function in both camouflage and sexual
selection, especially in environments characterized by
uneven dappled light, for example forest (Doucet and
Meadows 2009; Seago et al. 2009). Metallic colours in
fossil insects may have had similar functions, particularly
in specimens from palaeolakes surrounded by forest (e.g.
Messel (Schaal and Ziegler 1992), Clarkia (Smiley 1985))
and in specimens with original green hues. Many struc-
turally coloured fossil insects exhibit blue hues; given that
structural colours are blueshifted during fossilization
(McNamara et al. 2013a), at least some of these blue fos-
sil cuticles are likely to have been originally green. Some
fossil insects preserve anatomical evidence for modifica-
tion of iridescence to enhance defensive signals, that is,
camouflage and aposematism (Fig. 1E; McNamara et al.
2011); cryptic functions have been inferred from mono-
tonal colour patterns (Wang et al. 2010).
Striking colour patterns and glossy iridescence in vari-
ous feathered dinosaurs suggest that sexual display or
defence was important in the early evolution of plumage
and feather colour (Li et al. 2010, 2012). This is sup-
ported by variations in colour within individual penna-
ceous fossil feathers in Anchiornis, a basal paravian,
indicating that melanin-based intrafeather patterns
evolved before powered flight (Li et al. 2010). Some fossil
feathers exhibit evidence for the involvement of melanin
in nonvisual roles. Several authors have observed trends
in the intensity of dark tones within individual fossil
feathers and related these to the degree of melanization
and hence feather function. As in modern feathers (Lucas
and Stettenheim 1972; Hill and McGraw 2006), visual
tone can be pale in down feathers (McKellar et al. 2011)
and in basal regions of contour feathers (Li et al. 2010)
and darkest at the distal tip of contour feathers (Fig. 3B;
Clarke et al. 2010; Carney et al. 2012) and in distal bar-
bules (which overlap their proximal counterparts). Such
increased melanization has been suggested to confer
increased mechanical strength (Li et al. 2010; Carney
et al. 2012). Other fossil feathers exhibit dark tones in
proximal regions (e.g. Fig. 3C; Wogelius et al. 2011,
fig. 3A). Some authors have suggested that specific mela-
nosome geometries and configurations may affect the
material properties of fossil feathers (Clarke et al. 2010),
but these hypotheses are largely untested in modern
material.
Identification of colour-producing structures in some
fossil theropods has significant implications for our
understanding of the evolution of feathers. Discoveries of
melanosomes in integumentary filaments of the tail of
Sinosauropteryx (Zhang et al. 2010) refute claims that the
filaments are partially decayed dermal collagen fibres
(Lingham-Soliar 2003; Feduccia et al. 2005; Lingham-
Soliar et al. 2007) and support interpretations of the fila-
ments as feather homologues, that is, precursors of true
feathers. Although the function of colouration in such
structures is still unclear, the filaments themselves are
considered to occur in sufficient densities to have had
important nonvisual functions in thermoregulation and
protection (McKellar et al. 2011).
FUTURE DIRECTIONS
Certain aspects of the taphonomy of colour in insects and
feathers are poorly understood, not least that of pigmentary
colours in fossil insects. Some common insect pigments
(e.g. carotenoids and pterins) are also abundant in plants;
the degraded remains of plant-derived pigments are often
preserved in sedimentary organic matter and are the basis
of many studies of lake productivity (e.g. Romero-Viana
et al. 2009). It is therefore reasonable to hypothesize that

evidence of some insect pigments may survive in the fossil
record. Resolving the taphonomy of insect pigments will
require extensive chemical analysis of fossil insect remains
and taphonomic experiments. By providing insights into
the relative preservation potential of different insect pig-
ments, and into the nature of any diagnostic biomarkers
for such pigments when they decay, such experimental and
chemical studies are likely to prove critical to our under-
standing of the origins of monotonal patterning in fossil
insects and will test previous suggestions (Vinther et al.
2008) that such patterning reflects the distribution of eu-
melanin.
Loss of structural colour via oxidative weathering has
been invoked to explain inter- and intrabiota patterns in
the presence or absence of metallic colours, and even
within individual specimens, but the chemical processes
involved are uncertain. Chemical analysis of structurally
coloured cuticles at various stages of oxidative degrada-
tion could inform on this phenomenon and could also
help to explain patterns in the fidelity of preservation of
pigments in fossil insects and feathers.
The chemistry of melanin in fossil feathers is not fully
understood. Chemical techniques such as EPR and ToF-
SIMS have proved critical to our understanding of the
taphonomy of melanin in fossil squid (Glass et al. 2012)
and fish (Lindgren et al. 2012), but have not been applied
to fossil feathers. Doing so will test whether taxonomic or
tissue-related factors influence the chemical fidelity of
preservation of melanin in fossils. SRS-XRF analyses have
yielded intriguing insights into the elemental composition
of fossil fish, squid and, in particular, feathers (Wogelius
et al. 2011). The ability of this technique to resolve ques-
tions regarding the preservation of melanin in fossil feath-
ers could be tested by comparing data on the distribution
and abundance of various trace elements for specimens of
various taxa (including those that do not possess mela-
nin) from a single biota.
Other unresolved aspects of the taphonomy of colour
in feathers are also amenable to experimental testing.
Some authors have surmised that eumelanosomes and
phaeomelanosomes may differ in their resistance to dia-
genetic alteration (Clarke et al. 2010). The relative pres-
ervation potential of the different melanosome types
could be investigated via conventional decay- and matu-
ration experiments; decay experiments could also assess
whether autolytic and microbial decay processes affect
melanosome geometry. Morphological and chemical anal-
yses of degraded feather tissues could inform on the
taphonomic processes that influence the preservation of
melanosomes as external moulds; such studies could also
help resolve the nature of melanin (and its derivatives),
and the chromophore responsible for visual dark colour-
ation, in feathers preserving mouldic melanosomes.
Future studies of melanosome taphonomy should also
MCNAMARA: FOSSIL COLOUR 571
consider factors that may contribute to alteration of
melanosome geometry and packing arrangement in fos-
sils. Additional maturation experiments will constrain the
range of pressure–temperature conditions capable of
inducing morphological change and may allow the extent
of diagenetic alteration of melanosomes to be predicted.
The keratinous feather matrix is the basis of several col-
our-producing nanostructures, yet little is known about its
physical taphonomy. Recent maturation experiments using
structurally coloured feathers revealed that decay of quasi-
ordered nanostructures generates diagnostic ultrastructures
(McNamara et al. 2013b). Such textures could serve as a
proxy for the former presence of quasi-photonic nano-
structures in fossils. The chemical taphonomy of keratin
has been studied in detail in various fossil reptiles (Man-
ning et al. 2009; Edwards et al. 2011) and, to a lesser
extent, in fossil birds. Immunological evidence of keratin
has been reported in feathers and claw sheaths from Creta-
ceous theropods (Schweitzer et al. 1999a, b). Amide peaks
in FTIR spectra of fossil feathers may derive, in part, from
feather b-keratin (Barden et al. 2011). Keratin-chelated Cu
released during decay may bind to melanin during later
diagenesis, enhancing Cu concentrations in fossil feathers
(Wogelius et al. 2011). Sulphur associated with fossil feath-
ers may derive from feather keratin (Wogelius et al. 2011).
Future chemical analyses of fossil feathers will inform on
the taphonomy of the keratin molecule and will constrain
the extent to which it is linked to the taphonomy of feather
melanin and melanosomes.
CONCLUSIONS
Understanding the taphonomic processes that influence
the preservation of pigmentary and structural colour is of
fundamental importance to palaeobiologists interested in
the evolutionary history of colour and its functions. Fos-
sils with obvious colour and colour patterns, and those
lacking visible colour, have the potential to retain physical
and chemical evidence of colour-generating mechanisms.
Preservation of colour is controlled by a suite of tapho-
nomic factors that are, at present, not fully resolved; key
factors identified to date include the depth of burial and
the extent of hydrothermal alteration and weathering.
Additional taphonomic data from fossils and from
controlled laboratory experiments are critical in unravel-
ling the taphonomic history of colour in different taxa,
depositional environments and diagenetic regimes. This
will inform interpretations of original colour in fossils
and of the role of colour in visual signalling through
time.
Acknowledgements. My research on the taphonomy of colour in
fossils was funded by an Irish Research Council – Marie Curie
572 PALAEONTOLOGY, VOLUME 56
International Mobility Fellowship. I thank Derek Briggs and Pat-
rick Orr for their advice, collaboration and many fruitful discus-
sions of taphonomy. I also thank Mike Benton, Hui Cao, Daniel
Field, Neal Gupta, Stuart Kearns, Emma Locatelli, Laura Meyer,
Heeso Noh, Lin Qiu, Sonja Wedmann and Hong Yang for their
significant contributions to published and ongoing collaborative
studies. I am grateful to G€ unter Bechly, Susan Butts, Thomas
Engel, Larry Gall, David Grimaldi, Kristof Kristoffersen, Herbert
Lutz, Leonard Munsterman, Markus Poschmann, Michael Ras-
ser, Greg Watkins-Colwell and Michael Wuttke for access to fos-
sil and extant specimens, and to Daniel Field, Zhenting Jiang,
Robert Patalano, Barry Piekos, Vinod Saranathan, Elizabeth
Savrann, Jess Utrup, Robert Young and Shuang Zhang for assis-
tance with laboratory techniques. Thanks also to Ant
onia
Monteiro, Paul Nash, Jeffrey Oliver, Rick Prum, Jakob Vinther
and the members of the G&G Paleo group and the staff of the
Peabody Museum Entomology Division for their insights into
taphonomy, evolution and the colours of extant organisms.
Editor. Patrick Orr
REFERENCES
A D A C H I , E. 2007. Unexpected variability of millennium
green: structural color of Japanese jewel beetle resulted from
thermosensitive porous organic multilayer. Journal of Morphol-
ogy, 268, 826–829.
A N D E R S E N , S. O. 2010. Insect cuticular sclerotization: a
review. Insect Biochemistry and Molecular Biology, 40, 166–178.
B A R D E N , H. E., W O G E L I U S , R. A., L I , D., M A N N I N G ,
P. L., E D W A R D S , N. P. and V A N D O N G E N , B. E. 2011.
Morphological and geochemical evidence of eumelanin preser-
vation in the feathers of the Early Cretaceous bird, Gansus yu-
menensis. PLoS One, 6, e25494.
B E R G M A N N , U., M O R T O N , R. W., M A N N I N G , P. L.,
S E L L E R S , W. I., F A R R A R , S., H U N T L E Y , K. G.,
W O G E L I U S , R. A. and L A R S O N , P. 2010. Archaeopteryx
feathers and bone chemistry fully revealed via synchrotron
imaging. Proceedings of the National Academy of Sciences, 107,
9060–9065.
B L O D G E T T , R. B., B O U C O T , A. J. and F E R R I L L , B. A.
1983. A color-banded Beachia (Brachiopoda; Terebratulida)
from the Oriskany equivalent (mid-Early Devonian) of central
Alabama. Journal of Paleontology, 57, 865–869.
B O K O N Y , V., L I K E R , A., S Z E K E L Y , T. and K I S , J. 2003.
Melanin-based plumage coloration and flight displays in plo-
vers and allies. Proceedings of the Royal Society of London B:
Biological Sciences, 270, 2491–2497.
B U T L E R , M. J., G A R D I N E R , R. B. and D A Y , A. W. 2005.
Fungal melanin detection by the use of copper sulfide-silver.
Mycologia, 97, 312–319.
C A R N E Y , R. M., V I N T H E R , J., S H A W K E Y , M. D.,
D ’ A L B A , L. and A C K E R M A N N , J. 2012. New evidence
on the colour and nature of the isolated Archaeopteryx feather.
Nature Communications, 3, 637–643.
C L A R K E , J. A., K S E P K A , D. T., S A L A S - G I S M O N D I , R.,
A L T A M I R A N O , A. J., S H A W K E Y , M. D., D ’ A L B A , L.,
V I N T H E R , J., D E V R I E S , T. J. and B A B Y , P. 2010. Fossil
evidence for evolution of the shape and color of penguin
feathers. Science, 330, 954–957.
CE
S A R I N I , J. P. and I N S E R M 1996. Melanins and their pos-
sible roles through biological evolution. Advances in Space
Research, 18, 35–40.
C R O M A R T I E , R. I. T. 1959. Insect pigments. Annual Review
of Entomology, 4, 59–76.
D A V I S , P. G. and B R I G G S , D. E. G. 1995. Fossilization of
feathers. Geology, 23, 783–786.
D O U C E T , S. M. and M E A D O W S , M. G. 2009. Iridescence:
a functional perspective. Journal of the Royal Society Interface,
6, S115–S132.
- S H A W K E Y , M. D., H I L L , G. E. and M O N T G O M -
E R I E , R. 2006. Iridescent plumage in satin bowerbirds: struc-
ture, mechanisms and nanostructural predictors of individual
variation in colour. Journal of Experimental Biology, 209, 380–
390.
E D W A R D S , N. P., B A R D E N , H. E., V A N D O N G E N , B.
E., M A N N I N G , P. L., L A R S O N , P. L., B E R G M A N N ,
U., S E L L E R S , W. I. and W O G E L I U S , R. A. 2011. Infrared
mapping resolves soft tissue preservation in 50 million year-
old reptile skin. Proceedings of the Royal Society B, 278, 3209–
3218.
F A R R A N T , P. A. 1997. Color in nature: a visual and scientific
exploration. Blandford Press, London, 208 pp.
F E D U C C I A , A., L I N G H A M - S O L I A R , T. and H I N C H -
L I F F E , J. R. 2005. Do feathered dinosaurs exist? Testing the
hypothesis on neontological and paleontological evidence.
Journal of Morphology, 266, 125–166.
F O X , D. L. 1976. Animal biochromes and structural colours, Sec-
ond edition. University of California Press, Berkeley and Los
Angeles, California, 450 pp.
G H I R A D E L L A , H. 1998. Hairs, bristles, and scales. 637–645.
In L O C K E , M. (ed.). Microscopic anatomy of invertebrates,
Vol. 11A: Insecta. Wiley-Liss, New York, 496 pp.
-2009. Coloration. 213–220. In R E S H , V. H. and C A R D E,

R. T. (eds). Encyclopedia of insects, Second edition. Elsevier,
New York, 1024 pp.
G L A S S , K., I T O , S., W I L B Y , P. R., S O T A D , T., N A K A M -
U R A D , A., B O W E R S , C. R., V I N T H E R , J., D U T T A G ,
S., S U M M O N S , R., B R I G G S , D. E. G., W A K A M A T S U ,
K. and S I M O N , J. D. 2012. Direct chemical evidence for eu-
melanin pigment from the Jurassic period. Proceedings of the
National Academy of Sciences, 109, 10218–10223.
G O L D S T E I N , G., F L O R Y , K. R., B R O W N E , B. A., M A J -
I D , S., I C H I D A , J. M. and B U R T T , E. H. 2004. Bacterial
degradation of black and white feathers. Auk, 121, 656–659.
G R I M A L D I , D. and E N G E L , M. S. 2005. Evolution of the
insects. Cambridge University Press, Cambridge, 755 pp.
G U N D E R S O N , A. R., F R A M E , A. M., S W A D D L E , J. P.
and F O R S Y T H , M. H. 2008. Resistance of melanised feath-
ers to bacterial degradation: is it really so black and white?
Journal of Avian Biology, 39, 539–545.
H A G D O R N , H. and S A N D Y , M. R. 1998. Color banding in
the Triassic terebratulid brachiopod Coenothyris from the Mu-
schelkalk of Central Europe. Journal of Paleontology, 72, 11–
27.

H E A D S , S., M A R T I L L , D. M. and L O V E R I D G E , R. 2005.
An exceptionally preserved antlion (Insecta: Neuroptera) with
colour pattern preservation from the Cretaceous of Brazil. Pal-
aeontology, 48, 1409–1417.
H I L L , G. E. and M C G R A W , K. J. 2006. Bird coloration. vol.
1: Mechanisms and measurements. Harvard University Press,
Cambridge, MA, 589 pp.
H I N T O N , H. 1973. Natural deception. 97–159. In G R E G -
O R Y , R. L. and G O M B R I C H , E. H. (eds). Illusion in nat-
ure and art. Duckworth, London, 288 pp.
H I T C H C O C K , A. P., D Y N E S , J. J., L A W R E N C E , J. R.,
O B S T , M., S W E R H O N E , G. D. W., K O R B E R , D. R. and
L E P P A R D , G. G. 2009. Soft X-ray spectromicroscopy of
nickel sorption in a natural river biofilm. Geobiology, 7,
432–453.
H O F M A N N , C. M., M C G R A W , K. J., C R O N I N , T. W.
and O M L A N D , K. E. 2007. Melanin coloration in New
World orioles I: carotenoid masking and pigment dichroma-
tism in the orchard oriole complex. Journal of Avian Biology,
38, 163–171.
H O L L I N G W O R T H , N. T. J. and B A R K E R , M. J. 1991.
Colour pattern preservation in the fossil record: taphonomy
and diagenetic significance. 105–119. In D O N O V A N , S. K.
(ed.). The processes of fossilization. Columbia University Press,
New York, NJ, 303 pp.
HO € S S , M., D I L L I N G , A., C U R R A N T , A. and P A
€A€ B O , S.
1996. Molecular phylogeny of the extinct ground sloth Myl-
odon darwinii. Proceedings of the National Academy of Sciences,
93, 181–185.
I N G R A M , A. L. and P A R K E R , A. R. 2008. A review of the
diversity and evolution of photonic structures in butterflies,
incorporating the work of John Huxley (The Natural History
Museum, London from 1961 to 1990). Philosophical Transac-
tions of the Royal Society B, 363, 2465–2480.
K I N O S H I T A , S. and Y O S H I O K A , S. 2005. Structural colors
in biological systems: principles and applications. Osaka Univer-
sity Press, Osaka, 351 pp.
K L U G , C., S C H U L Z , H. and D E B A E T S , K. 2009. Red
Devonian trilobites with green eyes from Morocco and the
silicification of the trilobite exoskeleton. Acta Palaeontologica
Polonica, 54, 117–123.
K N I G H T , T. K., B I N G H A M , S., L E W I S , R. D. and S A V -
R D A , C. E. 2011. Feathers of the Ingersoll Shale, Eutaw For-
mation (Upper Cretaceous), Eastern Alabama: the largest
collection of feathers from North American Mesozoic Rocks.
Palaios, 26, 364–376.
K U R A C H I , M., T A K A K U , Y., K O M I Y A , Y. and H A R I Y -
A M A , T. 2002. The origin of extensive colour polymorphism
in Plateumaris sericea (Chrysomelidae, Coleoptera). Naturwis-
senschaften, 89, 295–298.
L A B A N D E I R A , C. 2005. Recent and exciting developments in
understanding fossil insects and their terrestrial relatives.
American Paleontologist, 13, 8–11.
L A N D , M. F. 1972. The physics and biology of animal
reflectors. Progress in Biophysical and Molecular Biology, 24,
75–106.
-1997. Visual acuity in insects. Annual Review of Entomol-
ogy, 42, 147–177.
MCNAMARA: FOSSIL COLOUR 573
L I , Q., G A O , K.-Q., V I N T H E R , J., S H A W K E Y , M. D.,
C L A R K E , J. A., D ’ A L B A , L., M E N G , Q., B R I G G S ,
D. E. G. and P R U M , R. O. 2010. Plumage color patterns of
an extinct dinosaur. Science, 327, 1369–1372.
- -M E N G , Q., C L A R K E , J. A., S H A W K E Y , M. D.,
D ’ A L B A , L., P E I , R., E L L I S O N , M., N O R E L L , M. A.
and V I N T H E R , J. 2012. Reconstruction of Microraptor and
the evolution of iridescent plumage. Science, 335, 1215–1219.
L I N D G R E N , J., U V D A L , P., S J O € V A L L , P., N I L S S O N ,
D. E., E N G D A H L , A., S C H U L T Z , B. P. and T H I E L , V.
2012. Molecular preservation of the pigment melanin in fossil
melanosomes. Nature Communications, 3, 824–831.
L I N G H A M - S O L I A R , T. 2003. The dinosaurian origin of
feathers: perspectives from dolphin (Cetacea) collagen fibres.
Naturwissenschaften, 90, 563–567.
-2011. The evolution of the feather: Sinosauropteryx, a col-
orful tail. Journal of Ornithology, 152, 567–577.
-and P L O D O W S K I G. 2010. The integument of Psittaco-
saurus from Liaoning Province, China: taphonomy, epidermal
patterns and color of a ceratopsian dinosaur. Naturwissens-
chaften, 97, 479–486.
- F E D U C C I A , A. and W A N G , X. 2007. A new Chinese
specimen indicates that ‘protofeathers’ in the early Cretaceous
theropod dinosaur Sinosauropteryx are degraded collagen fibres.
Proceedings of the Royal Society of London B, 274, 1823–1829.
L I U , Y. and S I M O N , J. D. 2003. Isolation and biophysical
studies of natural eumelanins: applications of imaging tech-
nologies and ultrafast spectroscopy. Pigment Cell Research, 16,
606–618.
- H O N G , L., W A K A M A T S U , K., I T O , S., A D H -
Y A R U , B., C H E N G , C.-Y., B O W E R S , C. R. and
S I M O N , J. D. 2005. Comparison of structural and chemical
properties of black and red human hair melanosomes. Photo-
chemistry and Photobiology, 81, 135–144.
L U C A S , A. M. and S T E T T E N H E I M , P. R. 1972. Avian
anatomy: integument. Agriculture handbook 362, US Depart-
ment of Agriculture, Washington, DC, 750 pp.
L U T Z , H. 1990. Systematische und pale€ okologische Untersuch-
ungen an Insekten aus dem Mittel-Eoz€an der Grube Messel bei
Darmstadt. Courier Forschungsinstitut Senckenberg, 124, 1–165.
M A C L E O D , H. A. 1969. Thin film optical filters. Adam Hilger,
London, 519 pp.
M A N N I N G , P. L., M O R R I S , P. M., M C M A H O N , A.,
J O N E S , E., G I Z E , A., M A C Q U A K E R , J. H. S., W O L F F ,
G., T H O M P S O N , A., M A R S H A L L , J., T A Y L O R , K. G.,
L Y S O N , T., G A S K E L L , S., R E A M T O N G , O., S E L L -
E R S , W. I., V A N D O N G E N , B., B U C K L E Y , M. and
W O G E L I U S , R. 2009. Mineralized soft-tissue structure and
chemistry in a mummified hadrosaur from the Hell Creek
Formation, North Dakota (USA). Proceedings of the Royal
Society B: Biological Sciences, 276, 3429–3437.
M A R K S , M. S. and S E A B R A , M. C. 2001. The melanosome:
membrane dynamics in black and white. National Review of
Molecular and Cellular Biology, 2, 738–748.
M A S O N , C. W. 1927. Structural colors in insects. II. Journal of
Physical Chemistry, 31, 321–354.
M C G R A W , K. 2003. Melanins, metals, and mate quality. Oi-
kos, 102, 402–406.
 574 PALAEONTOLOGY, VOLUME 56
-2006. Mechanics of melanin-based coloration. 243–294. In
H I L L , G. E. and M C G R A W , K. J. (eds). Bird coloration:
Vol. 1: mechanisms and measurements. Harvard University
Press, Cambridge, MA, 589 pp.
M C K E L L A R , R. C., C H A T T E R T O N , B. D. E., W O L F E ,
A. P. and C U R R I E , P. J. 2011. A diverse assemblage of Late
Cretaceous dinosaur and bird feathers from Canadian amber.
Science, 333, 1619–1622.
M C N A M A R A , M. E., B R I G G S , D. E. G., O R R , P. J.,
W E D M A N N , S., N O H , H. and C A O , H. 2011. Fossilized
biophotonic nanostructures reveal the original colors of 47-
million-year-old moths. PLoS Biology, 9, e1001200.
--- 2012a. The controls on the preservation of
structural color in fossil insects. Palaios 27, 443–454.
- - -N O H , H. and C A O , H. 2012b. The original
colours of fossil beetles. Proceedings of the Royal Society B,
279, 1114–1121.
- - -G U P T A , N. S., L O C A T E L L I , E. R., Q I U ,
L., Y A N G , H., W A N G , Z., N O H , H. and C A O , H. 2013a.
The fossil record of insect color illuminated by maturation
experiments. Geology, published online 20 February 2013. doi:
10.1130/G33836.1.
--- F I E L D , D. and W A N G , Z. 2013b. Experi-
mental maturation of feathers: implications for reconstructions
of fossil feather colour. Biology Letters, 9, published online
March 2013. doi: 10.1098/rsbl.2013.0184.
N A P P I , A. J. and C H R I S T E N S E N , B. M. 2005. Melanogen-
esis and associated cytotoxic reactions: applications to insect
innate immunity. Insect Biochemistry and Molecular Biology,
35, 443–459.
N E V I L L E , A. C. 1975. Biology of the arthropod cuticle.
Springer, New York, 458 pp.
- and C A V E N E Y S. 1969. Scarabaeid beetle exocuticle as
an optical analogue of cholesteric liquid crystals. Biological
Reviews, 44, 531–562.
N O H , H., L I E W , S. F., S A R A N A T H A N , V., M O C H R I E ,
S. G. J., P R U M , R. O., D U F R E S N E , E. R. and C A O , H.
2010. How noniridescent colors are generated by quasi-
ordered structures of bird feathers. Advanced Materials, 22,
2871–2880.
N O R E L L , M. A. 2011. Fossilized feathers. Science, 333, 1590–
1591.
N O Y E S , J. A., V U K U S I C , P. and H O O P E R , I. R. 2007.
Experimental method for reliably establishing the refractive
index of buprestid beetle exocuticle. Optics Express, 15, 4351–
4358.
P A R K E R , A. R. 1998. Colour in Burgess Shale animals and the
effect of light on evolution in the Cambrian. Proceedings of the
Royal Society B, 265, 965–971.
- 2000. 515 Million years of structural colour. Journal of
Optics A: Pure and Applied Optics, 2, R15–R28.
-2002. Natural photonic engineers. Materials Today, 5, 26–31.
-and M C K E N Z I E D. R. 2003. The cause of 50 million-
year-old colour. Biology Letters, 270, S151–S153.
- and T O W N L E Y H. E. 2007. Biomimetics of photonic
nanostructures. Nature Nanotechnology, 2, 347–353.
-M C K E N Z I E , D. R. and L A R G E , M. C. J. 1998a. Multi-
layer reflectors in animals using green and gold beetles as
contrasting examples. Journal of Experimental Biology, 201,
1307–1313.
-H E G E D U S , Z. and W A T T S , R. A. 1998b. Solar-absorb-
er antireflector on the eye of an Eocene fly (45 Ma). Proceed-
ings of the Royal Society B, 265, 811–815.
- M C P H E D R A N , R. C., M C K E N Z I E , D. R., B O T -
T E N , L. C. and N I C O R O V I C I , N.-A. P. 2001. Aphrodite’s
iridescence. Nature, 409, 36–37.
P O W E L L , J. A. 2003. Lepidoptera (Moths, Butterflies). 631–
663. In R E S H , V. H. and C A R D E,
R. T. (eds). Encyclopedia
of insects. Academic Press, Burlington, MA, 1024 pp.
P R U M , R. O. 2006. Anatomy, physics, and evolution of struc-
tural colors. 295–353. In H I L L , G. E. and M C G R A W , K. J.
(eds). Bird coloration. Vol 1: mechanisms and measurements.
Harvard University Press, Cambridge, MA, 589 pp.
-and T O R R E S R. H. 2003. A Fourier tool for the analysis
of coherent light scattering by bio-optical nanostructures. Inte-
grative and Comparative Biology, 43, 591–602.
R A S N I T S Y N , A. P. and Q U I C K E , D. L. J. 2002.History of
insects. Kluwer Academic Publishers, Dordrecht, 517 pp.
R I L E Y , P. A. 1997. Melanin. International Journal of Biochemis-
try and Cell Biology, 29, 1235–1239.
R O M E R O - V I A N A , L., K E E L Y , B. J., C A M A C H O , A.,
V I C E N T E , E. and M I R A C L E , M. R. 2009. Photoautotro-
phic community changes in Lagunillo del Tejo (Spain) in
response to lake level fluctuation: two centuries of sedimentary
pigment records. Organic Geochemistry, 40, 376–386.
S C H A A L , S. and Z I E G L E R , W. 1992. Messel: an insight into
the history of life and of the Earth. Clarendon Press, Oxford,
328 pp.
S C H U L T Z , T. D. and R A N K I N , M. A. 1985. The ultrastruc-
ture of the epicuticular interference reflectors of tiger beetles
(Cicindela). Journal of Experimental Biology, 117, 87–110.
S C H W E I T Z E R , M. H. 2011. Soft tissue preservation in terres-
trial Mesozoic vertebrates. Annual Reviews of Earth and Plane-
tary Sciences, 39, 187–216.
-W A T T , J. A., A V C I , R., F O R S T E R , C. A., K R A U S E ,
D. W., K N A P P , L., R O G E R S , R. R., B E E C H , I. and
M A R S H A L L , M. 1999a. Keratin immunoreactivity in the
Late Cretaceous bird Rahonavis ostromi. Journal of Vertebrate
Paleontology, 19, 712–722.
- - -K N A P P , L., C H I A P P E , L., N O R E L L , M.
and M A R S H A L L , M. 1999b. Beta-keratin specific immuno-
logical reactivity in feather-like structures of the Cretaceous
alvarezsaurid, Shuvuuia deserti. Journal of Experimental Zool-
ogy, 285, 146–157.
S C H W E I Z E R , M. K., W O O L L E R , M. J., T O P O R S K I , J.,
F O G E L , M. L. and S T E E L E , A. 2006. Examination of an Oli-
gocene lacustrine ecosystem using C and N stable isotopes. Pal-
aeogeography, Palaeoclimatology, Palaeoecology, 230, 335–351.
S E A G O , A. E., B R A D Y , P., V I G N E R O N , J.-P. and
S C H U L T Z , T. D. 2009. Gold bugs and beyond: a review of
iridescence and structural colour mechanisms in beetles (Cole-
optera). Journal of the Royal Society Interface, 6, S165–S184.
S H A W K E Y , M. D. and H I L L , G. E. 2006. Significance of a
basal melanin layer to production of non-iridescent structural
plumage color: evidence from an amelanotic Steller’s jay (Cya-
nocitta stelleri). Journal of Experimental Biology, 209, 1245–1250.

-E S T E S , A. M., S I E F F E R M A N , L. M. and H I L L , G. E.
2003. Nanostructure predicts intraspecific variation in ultravi-
olet-blue plumage colour. Proceedings of the Royal Society B,
270, 1455–1460.
-H A U B E R , M. E., E S T E P , L. K. and H I L L , G. E. 2006.
Evolutionary transitions and mechanisms of matte and irides-
cent plumage coloration in grackles and allies (Icteridae).
Journal of the Royal Society Interface, 3, 777–786.
-S A R A N A T H A N , V., P A
LSDO
T T I R , H., C R U M , J.,
E L L I S M A N , M., A U E R , M. and P R U M , R. O. 2009.
Electron tomography, three-dimensional Fourier analysis and
colour prediction of a three-dimensional amorphous biopho-
tonic nanostructure. Journal of the Royal Society Interface, 6,
S213–S220.
S H A W K E Y , M., M A I A , R. and D ’ A L B A , L. 2011. Proxi-
mate bases of silver colour in anhinga (Anhinga anhinga)
feathers. Journal of Morphology, 272, 1399–1407.
S H I , Y. and Y O K O Y A M A , S. 2003. Molecular analysis of the
evolutionary significance of ultraviolet vision in vertebrates.
Proceedings of the National Academy of Sciences, 100, 8308–
8313.
S H O C K , E. L. and K O R E T S K Y , C. M. 1993. Metal-organic
complexes in geochemical processes: calculation of standard
partial molal thermodynamic properties of aqueous acetate
complexes at high pressures and temperatures. Geochimica et
Cosmochimica Acta, 57, 4899–4922.
S M I L E Y , J. 1985. Late Cenozoic history of the Pacific northwest.
American Association for the Advancement of Science, San
Francisco, 417 pp.
S T O D D A R D , M. C. and P R U M , R. O. 2011. How colorful
are birds? Evolution of the avian plumage color gamut.
Behavioural Ecology, 22, 1042–1052.
T A N A K A , G., P A R K E R , A. R., S I V E T E R , D. J., M A E D A ,
H. and F U R U T A N I , M. 2009. An exceptionally well-pre-
served Eocene dolichopodid fly eye: function and evolutionary
significance. Proceedings of the Royal Society of London B, 276,
1015–1019.
- T A N I G U C H I , H., M A E D A , H. and N O M U R A , S.
2010. Original structural colour preserved in an ancient leaf
beetle. Geology, 38, 127–130.
T I L L E Y , R. J. D. and E L I O T , J. N. 2002. Scale microstructure
and its phylogenetic implications in lycaenid butterflies (Lepi-
doptera, Lycaenidae). Transactions of the Lepidopteran Society
of Japan, 53, 153–180.
T O P O R S K I , J. K. W., S T E E L E , A., W E S T A L L , F., A V C I ,
R., M A R T I L L , D. M. and M C K A Y , D. S. 2002. Morpho-
logic and spectral investigation of exceptionally well preserved
bacterial biofilms from the Oligocene Enspel formation, Ger-
many. Geochimica et Cosmochimica Acta, 66, 1773–1791.
T U R E K , V. 2009. Colour patterns in Early Devonian cephalo-
pods from the Barrandian area: taphonomy and taxonomy.
Acta Palaeontologica Polonica, 54, 491–502.
MCNAMARA: FOSSIL COLOUR 575
V I N T H E R , J., B R I G G S , D. E. G., P R U M , R. O. and
S A R A N A T H A N , V. 2008. The colour of fossil feathers.
Biology Letters, 4, 522–525.
- -C L A R K E , J., M A Y R , G. and P R U M , R. O. 2010.
Structural coloration in a fossil feather. Biology Letters, 6, 128–
131.
V O N D R A N , T., A P E L , K. H. and S C H M I T Z , H. 1995.
The infrared receptor of Melanophila acuminata De Geer
(Coleoptera: Buprestidae): ultrastructural study of a unique
insect thermoreceptor and its possible descent from a hair
mechanoreceptor. Tissue and Cell, 27, 645–658.
V U K U S I C , P. 2003. Natural coatings. 1–34. In K A I S E R , N.
and P U L K E R , H. (eds). Optical interference coatings.
Springer, Berlin, 500 pp.
-and S A M B L E S J. R. 2003. Photonic structures in biology.
Nature, 424, 852–855.
W A N G , Y., L I U , Z., W A N G , X., S H I H , C., Z H A O , Y.,
E N G E L , M. S. and R E N , D. 2010. Ancient pinnate leaf
mimesis among lacewings. Proceedings of the National Acad-
emy of Sciences, 107, 16212–16215.
W E D M A N N , S., P O S C H M A N N , M. and H O € RNSCHE-
M E Y E R , T. 2010. Fossil insects from the Late Oligocene
Enspel Lagerst€atte and their palaeobiogeographic and palaeo-
climatic significance. Palaeobiology and Palaeoenvironments,
90, 49–58.
W I L T S , B. D., L E E R T O U W E R , H. L. and S T A V E N G A ,
D. G. 2009. Imaging scatterometry and microspectropho-
tometry of lycaenid butterfly wing scales with perforated
multilayers. Journal of the Royal Society Interface, 6, S185–
S192.
W O G E L I U S , R. A., M A N N I N G , P. L., B A R D E N , H. E.,
E D W A R D S , N. P., W E B B , S. M., S E L L E R S , W. I.,
T A Y L O R , K. G., L A R S O N , P. L., D O D S O N , P., Y O U ,
H., D A - K I N G , L. and B E R G M A N N , U. 2011. Trace met-
als as biomarkers for eumelanin pigment in the fossil record.
Science, 333, 1622–1626.
W U T T K E , M. 1983. ‘Weichteil-Erhaltung’ durch lithifizierte
Mikroorganismen bei mittel-eoz€anen Vetebraten aus den
€
Olschiefern der ‘Grube Messel’ bei Darmstadt. Senckenbergiana
Lethaea, 64, 509–527.
Y I N , H., S H I , L., S H A , J., L I , Y., Q I N , Y., D O N G , B.,
M E Y E R , S., L I U , X., Z H A O , L. and Z I , J. 2006. Irides-
cence in the neck feathers of domestic pigeons. Physical
Review E, 74, 051916.
Y O S H I O K A , S., N A K A M U R A , E. and K I N O S H I T A , S.
2007. Origin of two-color iridescence in rock dove’s feather.
Journal of the Physical Society of Japan, 76, 013801 1–4.
Z H A N G , F., K E A R N S , S. L., O R R , P. J., B E N T O N , M. J.,
Z H O U , Z., J O H N S O N , D., X U , X. and W A N G , X.
2010. Fossilized melanosomes and the colour of Cretaceous
dinosaurs and birds. Nature, 463, 1075–1078.