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Pigmentation and color pattern diversity in Odonata
Genta Okude1,2 and Ryo Futahashi2
The order Odonata (dragonflies and damselflies) comprises
diurnal insects with well-developed vision, showing diverse
colors in adult wings and bodies. It is one of the most ancestral
winged insect groups. Because Odonata species use visual
cues to recognize each other, color patterns have been
investigated from ecological and evolutionary viewpoints. Here
we review the recent progress on molecular mechanisms of
pigmentation, especially focused on light-blue coloration.
Results from histology and pigment analysis showed that
ommochrome pigments on the proximal layer and pteridine
pigments on the distal layer of the epidermis are essential for
light-blue coloration. We also summarize genes involved in the
biosynthesis of three major insect pigments conserved across
insects and discuss that gene-functional analysis deserves
future studies.
Addresses
1
Department of Biological Sciences, Graduate School of Science, The
University of Tokyo, Bunkyo-ku, Tokyo, 113-0033, Japan
2
Bioproduction Research Institute, National Institute of Advanced
Industrial Science and Technology (AIST), Central 6, Tsukuba, Ibaraki,
305-8566, Japan
Corresponding authors: Okude, Genta (gentaokude@gmail.com),
Futahashi, Ryo (ryo-futahashi@aist.go.jp)
Current Opinion in Genetics and Development 2021, 69:14–20
This review comes from a themed issue on Developmental
mechanisms, patterning and evolution
Edited by Antónia Monteiro and Thomas Werner
For a complete overview see the Issue and the Editorial
Available online 20th January 2021
https://doi.org/10.1016/j.gde.2020.12.014
0959-437X/ã 2020 The Author(s). Published by Elsevier Ltd. This is an
open access article under the CC BY license (http://creativecommons.
org/licenses/by/4.0/).
Introduction
Insects account for over half of the described species and
exhibit a tremendous diversity of colors and patterns [1,2].
The order Odonata (dragonflies and damselflies) has
attracted attention in the study of insect color diversity
for the following reasons: (i) Odonata species show very
diverse colors in both adult wings and body [3–5], (ii)
Odonata species are diurnal insects with well-developed
vision [6,7], and their body color is essential for recogniz-
ing each other, (iii) unlike most other insects, Odonata
species drastically change their color from brownish dull-
colored aquatic larvae to colorful terrestrial adults without
pupal stage [4,8], (iv) many Odonata species have sexual
color dimorphism and change their color through sexual
Current Opinion in Genetics & Development 2021, 69:14–20
maturation [4,5,9–11], (v) some Odonata species exhibit
color polymorphism within the same sex, mostly in
females [4,12–14], and (vi) the order Odonata is one of
the most ancestral winged insects with metamorphosis
[8,15,16]; therefore the development, structure, and
molecular mechanisms of pigmentation in Odonata are
important to elucidate the evolution of insect color pat-
tern formation. Meanwhile, genes involved in pigmenta-
tion in Odonata remain largely unknown, partly due to the
difficulty of laboratory rearing and gene-functional anal-
ysis. Here we review the recent progress on the pigmen-
tation of Odonata with a particular focus on the light-blue
coloration and summarize genes involved in the biosyn-
thesis of three major insect pigments.
Pigments in Odonata
Among insect pigments, melanins, ommochromes, and
pteridines are present in most insects, including Odonata
[17]. In insects, melanins are present on the surface of the
cuticle, while ommochromes and pteridines are localized
in pigment granules within cells. Melanins are responsi-
ble for black and brown coloration and the hardness of the
cuticle across insects [18]. In Odonata, melanins are
involved in the darkening of the wings and epidermis
[5,19–21] and sometimes contribute to the production of
structural colors [20,22]. For example, males of the jew-
elwing damselfly Calopteryx japonica have a multilayer
structure in wing veins, which exhibits metallic blue
when melanin pigments accumulate during maturation
[5,20] (Figure 1a). In insects, NBAD-melanin and
NADA-melanin are involved in red and yellow pigmen-
tation [17,18], and the orange wing coloration of the wings
in the genus Mnais is attributed to melanin pigments [19]
(Figure 1b).
Ommochromes are pigments detected in the compound
eyes of most insects, and occasionally present as reddish
pigments in the wings and body [17,23]. In the reddish
Odonata species of the genera Sympetrum and Crocothemis,
two ommochrome pigments, xanthommatin and decar-
boxylated-xanthommatin, are the major pigments for
body color, and males turn from yellow to red during
maturation by redox reactions of the pigments [9]
(Figure 1c,d). Notably, in mature males of the genus
Sympetrum, red color turns darker at lower temperatures
[24,25] (Figure 1e), probably due to the migration of
ommochrome granules within the epidermal tissue,
although these temperature-dependent reversible color
changes are not prominent in Crocothemis [4] (Figure 1f).
Ommochrome pigments are also found in the wings, such
as in the red pterostigmas in the genus Mnais [19]
(Figure 1b, arrowhead).
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 Pigmentation in Odonata Okude and Futahashi 15

Figure 1

(a) (b) (c) (d)

(e) (f) (g)

(h) (i) (j) (k)

(l)

Current Opinion in Genetics & Development

Pigment-based or wax-based color pattern in Odonata. Sex and species names are provided for each image. (a,b) Melanin-based wing
coloration. White arrowhead in (b) indicates ommochrome-based pterostigma coloration. (c,d) Ommochrome-based body coloration. (e–g)
Temperature-dependent reversible color changes. After overnight treatment at 4 C (upper side) and after returning to 25 C for 1 hour (lower side).
(h-j) Pteridine-based yellow, green, or light-blue body coloration. (k) Wax-based body coloration. (l) Summary of pigments and waxes associated
with coloration of various Odonata species. Males of Calopteryx japonica, Calopteryx cornelia, Crocothemis servilia (mature and immature),
Ceriagrion melanurum, Paracercion hieroglyphicum, Lyriothemis pachygastra, and Orthetrum japonicum are shown.

Pteridine pigments are found on the light-yellow, green,
and blue regions of several dragonfly species and can emit
fluorescence under UV light [11,26,27,28] (Figure 1g–j).
As described below, pteridine pigments, together with
ommochrome pigments, are essential for the light-blue
coloration in several Odonata species [28,29–31].
Some Odonata species have UV-reflective whitish or bluish
waxes, originally defined as a supracuticular pigment [3], on
their body and/or wing surfaces [4,11,22,32,33] (Figure 1k).
Recently,uniquevery-long-chain methyl ketonesandalde-
hydes have been identified as the major wax components of
the abdomen of the Libellulidae Odonata species, and the
whitish color is attributed to the light-scattering fine struc-
tures produced by the self-assembly of the waxes [33].
It was also reported that carotenoids are present in various
Odonata species, but the components of carotenoids are
similar regardless of body color, suggesting that carote-
noids are not involved in the coloration of Odonata [34].
In Figure 1l, we summarized pigments and waxes associ-
ated with the coloration of various Odonata species.
Pigment synthesis genes in Odonata
Genes involved in pigment synthesis of melanins, ommo-
chromes, and pteridines have been identified primarily by

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16 Developmental mechanisms, patterning and evolution
Figure 2
(a) Melanins
Melanin (a), ommochrome (b), and pteridine (c) synthesis pathways in insects. Figure modified from Refs. [17,18,23,36]. Numbers in the
parenthesis indicate the number of homologous genes in Odonata. Asterisks (*) indicate differentially expressed genes among male and female
morphs of Iscunura damselflies [38,47].
analysis of Drosophila mutants [17,18,23]. A schematic
diagram of these pigment synthesis pathways is shown in
Figure 2. It should be noted that the melanin synthesis
pathway in insects differs markedly from that in verte-
brates by being dominated by dopamine-derived pig-
ments [18,35]. Ommochromes and pteridines are found
in the compound eyes in Drosophila and also contribute to
the coloration of other body parts in various insects. For
example, embryonic colors of water striders were identi-
fied as pteridines (erythropterin for red and xanthopterin
for yellow), and RNAi-mediated gene knockdown experi-
ments confirmed the conserved role of several genes in
pteridine synthesis and transport [36]. Here we sum-
marize the number of homologs of these genes in Odo-
nata based on the transcriptome and genomic information
[7,33,37–40]. Most genes can be identified as single
homologous genes in various Odonata species (Figure 2),
whereas there are several yellow and aaNAT family genes,
as reported in other insects [18]. The pathway of pteridine
synthesis has many similarities with vertebrates, suggest-
ing that studies in vertebrates may be helpful to under-
stand coloration in the order Odonata [41,42].
Pteridine-based light-blue coloration in
Odonata
Light-blue coloration is found across Odonata species and
families and has evolved more than 10 times indepen-
dently within the order Odonata [31]. Transmission
electron microscopic observations of the epidermis of
light-blue regions have identified two layers under the
Current Opinion in Genetics & Development 2021, 69:14–20
(b) Ommochromes (c) Pteridines
Current Opinion in Genetics & Development
transparent cuticle (Figure 3a) [28,29–31]. The distal
layer possesses numerous nanospheres arranged in a
specific manner, and this layer looks blueish by itself
when observed in reflected light [28]. In contrast, the
proximal layer possesses dark-colored granules, which
absorb stray light for fine-tuning the hue and brilliance
[28,29–31]. Simulations of reflectance spectra with an
optical finite-difference time-domain method suggest
that light-blue coloration can be produced by coherent
scattering of the pteridine-containing nanospheres in
the distal layer, and that the pigmentary absorption in
the proximal layer increases the saturation and brilliance
of the color [28]. It should be noted that the nano-
spheres in the distal layer are not mature in newly-
emerged adults, in which they are shiny-grayish in color
[28,29]. Many Ischnura species exhibit female color
polymorphisms [13,14]. The male-mimicking color
morph, called androchrome, shows a bluish color pattern,
whereas the female-specific color morph, called gyno-
chrome, often exhibits a reddish or brownish color pattern
(Figure 1i–j). Bluish regions of males and androchrome
females exhibit two epidermal layers: the distal layer with
pteridine nanospheres and the proximal layer with dark-
colored granules [28] (Figure 3a). In contrast, reddish or
brownish regions of gynochrome females have reddish
granules instead of dark-colored granules and a disor-
dered array of pteridine nanospheres intermingled with
reddish granules [28]. Some Odonata species exhibit
temperature-dependent reversible color changes from
light-blue to darker blue/gray [25,29] (Figure 1g). In
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 Pigmentation in Odonata Okude and Futahashi 17

Figure 3

(a) (b)

(c)

(d) (e)

Current Opinion in Genetics & Development

Coloration of a polymorphic damselfly Ischnura senegalensis (Coenagrionidae).

(a) Schematic illustration of epidermal coloration in a male and an androchrome female. (b) Schematic illustration of epidermal coloration in a
gynochrome female. (c) External and internal views of thoracic and abdominal epidermises after each treatment with different solvents. White
arrowheads indicate the black pigments in the proximal layer. Asterisks indicate the region where putative fat bodies turned whitish after organic
solvent treatment. (d) Dorsal views of RNAi phenotypes in the abdomen. MCO2: multicopper oxidase 2, bla: beta-lactamase (negative control). (e)
Dorsal views of RNAi phenotypes in the thorax. Gray arrowheads indicate unpigmented regions [44].

the lestid damselfly Austrolestes annulosus, dark-colored
granules in the proximal layer migrate to the distal layer at
low temperatures, darkening the blue color tone [29]. It
should be noted that temperature-dependent color
changes are not prominent in other Odonata species,
including I. senegalensis [30,31], and the effects sometimes
differ between markings in the same individual
(Figure 1g), suggesting that the mechanism of granule
migration differs among regions as well as species.
Pigment analysis in the blue-tailed damselfly
Ischnura senegalensis
The pigments in the dark-colored granules in the proxi-
mal layer of the light-blue regions have been presumed as
ommochrome pigments in several Odonata species
[29–31], while they are predicted to be melanins in
Ischnura elegans, based on the absorbance spectra. Because
the pigment group can be estimated from its solubility
[17], we examined the pigment composition of each

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18 Developmental mechanisms, patterning and evolution
epidermal layer of a closely related species, I. senegalensis,
by soaking body parts in different solvents (Okude et al.,
unpublished) (Figure 3c). The light-blue (male abdo-
men), green (male thorax), and reddish-brown (gyno-
chrome female thorax) colors were almost unaffected by
extraction with acetone-hexane and PBS-methanol, con-
firming a low contribution of carotenoids and flavonoids to
the body color. It should be noted that attached fat bodies
(observed from the internal view) become whitish in
organic solvents (Figure 3c, acetone-hexane). An acidic
methanol treatment dissolved the basal blackish color of
the epidermis (observed from the internal view), resulting
in a decolorization of blue or green (Figure 3c, 2% HCl in
methanol). An acidic methanol treatment also signifi-
cantly diminished the red color in gynochrome females
(Figure 3c, 2% HCl in methanol). These results suggest
that the dark-colored or reddish-brown granules in the
proximal layer of the epidermis contain ommochrome
pigments. Subsequent treatment with aqueous ammonia
turned the light-blue and green to transparent (Figure 3c,
aqueous ammonia), indicating the critical role of pteridine
pigments in these colors, as reported previously [28].
After the treatment with aqueous ammonia, the color of
the thorax of gynochrome females remained mostly intact
(Figure 3c, aqueous ammonia), suggesting the contribu-
tion of melanin pigments to the reddish-brown color,
which are insoluble in all these solvents.
We also performed a recently established electroporation-
mediated RNAi to inhibit the function of the multicopper
oxidase 2 (MCO2) gene, which is involved in the synthesis
of all melanin pigments [18,21,43,44] (Figure 2a). Sup-
pression of the MCO2 gene inhibited black and reddish-
brown coloration but had no effect on light-blue or green
coloration [44] (Figure 3d,e). These results, together with
the solubility results, indicate that both ommochromes
and pteridines are essential for light-blue and green
coloration, and both melanins and ommochromes are
important for the reddish-brown coloration (Figure 3a,
b). In the silkworm Bombyx mori and the red flour beetle
Tribolium castaneum, the MFS transporter encoded by the
red egg (re) gene, an insect homolog of the vertebrate
cysteine transporter MFSD12 [45], is required for dark-
colored ommochromes (ommins) [46], implying that the
re gene of Odonata may be involved in the production of
the dark-colored granules of light-blue regions.
Differentially expressed genes between
different color patterns in Odonata
With the spread of next-generation sequencers, studies
comparing gene expression among samples of different
color patterns have been reported in several Odonata
species [33,38,47,48]. Among male and two female
morphs of I. senegalensis, seven differentially expressed
genes were identified, including two melanin synthesis
genes, ebony and black, which are involved in the synthesis
of NBAD melanin [47] (Figure 2a). The higher
Current Opinion in Genetics & Development 2021, 69:14–20
expression of black and ebony genes in the thorax of
gynochrome females, as compared to males and andro-
chrome females, strongly suggests the contribution of
NBAD melanin in the reddish-brown color of the gyno-
chrome female thorax. Meanwhile, cinnabar and yellow
family genes were upregulated in gynochrome females in
I. elegans [38].
Notably, the sex-determining transcription factor double-
sex (dsx) was differentially expressed among female
morphs in I. senegalensis [47]. dsx is known to be involved
in sex determination in various insects [49,50] and sex-
limited polymorphism in Papilio swallowtail butterflies
[51–53]. In contrast, dsx was not differentially expressed
among female morphs in I. elegans, whereas doublesex-Mab
related 99B (dmrt99B) and several ecdysone synthesis
genes were differentially expressed [38,48]. The func-
tional analysis of these genes deserves future studies. It
should be noted that there are species differences
between I. senegalensis and I. elegans females in the pres-
ence or absence of light-blue regions on the female
abdomen. Because hybrids between these two species
can be produced [54], genetic analysis using hybrids may
be possible.
Other than Ischnura species, RNA-sequencing of Orthe-
trum albistylum revealed that three elongation of very long-
chain fatty acids (ELOVL) genes, members of the fatty acid
elongase gene family, were specifically expressed in the
wax-producing regions [33], suggesting that these
ELOVL genes are involved in production of the surface
wax of Odonata.
Conclusion and perspective
Here we review the recent progress on Odonata pigmen-
tation, especially focused on the light-blue coloration.
Comparative transcriptomics allow us to identify candi-
date genes for color pattern formation as well as genes
involved in pigment synthesis. However, gene-functional
analysis has rarely been reported in Odonata. To over-
come this problem, we have established a system for
breeding Odonata species in the laboratory [55], and a
method for electroporation-mediated RNAi [21,44]. We
hope that these techniques will accelerate to clarify the
molecular mechanisms underlying color pattern forma-
tion in Odonata.
Conflict of interest statement
Nothing declared.
Acknowledgements
We thank Minoru Moriyama and Takema Fukatsu for technical advises,
and Mizuko Osanai-Futahashi for helpful comments on the manuscript.
This work was partly supported by the JSPS KAKENHI Grant Numbers
JP18J21561 to GO and JP18H02491, JP18H04893, JP19H03287 and
JP20H04936 to RF.
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