Entomologia Experimentalis et Applicata 86: 125–133, 1998.

125

c 1998 Kluwer Academic Publishers. Printed in Belgium.

The determination of larval phase coloration in the African armyworm,

Spodoptera exempta and its consequences for thermoregulation and
protection from UV light

A. Gunn
School of Biological & Earth Sciences, Liverpool John Moores University, Byrom Street, Liverpool, L3 3AF, UK

Accepted: October 10, 1997

Key words: Spodoptera exempta, phase, coloration, thermoregulation, melanin, ultraviolet radiation

Abstract

Spodoptera exempta exhibits a density-dependent phase polyphenism in which caterpillars reared in isolation (phase
solitaria) tend to be green/brown and cryptic while those reared in groups (phase gregaria) are black and highly
visible. Differences in coloration between solitaria and gregaria phase larvae become apparent in the third instar
and are pronounced by the final instar. Larval rearing densities as low as two larvae per 250 ml container are
sufficient to induce gregaria coloration in 61% of the insects and at higher densities this is the only coloration
expressed. Larvae transferred from isolated to crowded conditions or vice versa tend to retain or adopt gregaria
phase coloration which suggests that this is highly advantageous. Black coloration is also induced in single larvae
reared with other species of caterpillar (S. littoralis). Isolated larvae reared at low temperature tend to become
melanic while crowded larvae reared at high temperature tend to be less melanised although they do not adopt
the solitaria phase coloration. These results suggest that phase coloration is determined by non species-specific
inter-larval contact, although it can be modulated by temperature. Under laboratory conditions, gregaria phase
larvae heat up faster than solitaria; this is partially a consequence of their black coloration but their smaller size
is a more important factor. The cuticles of the gregaria larvae absorb significantly more ultraviolet radiation than
those of solitaria but this does not confer any protection when larvae are irradiated at 254 nm.

Introduction the adult moths with consequent effects on fecundity

The African armyworm moth, Spodoptera exempta
Walker (Lepidoptera: Noctuidae) occurs throughout
tropical and southern Africa as well as southern Ara-
bia and the Far East (reviews by Haggis, 1984, 1986).
It is a serious but sporadic pest of grassland, maize
and a variety of other crops belonging to the family
Graminae.
In common with some other Noctuidae, the lar-
vae exhibit a density-dependent phase polyphenism
in which larvae reared under crowded conditions are
melanic and more active than those reared in isolation
(Faure, 1943; Matthee, 1946; Simmonds & Blaney,
1986). By analogy with locusts, the high- and low-
density forms are commonly referred to as gregaria
and solitaria respectively. Larval phase in S. exempta
has also been shown to affect lipid reserves present in
and flight capacity (Gunn & Gatehouse, 1987; Woo-
drow et al., 1987). Despite the dramatic differences
between the two phases, the role of polyphenism in the
ecology of S. exempta and its importance in the devel-
opment of outbreak populations has remained a matter
of debate (Rose, 1979; Gatehouse, 1987). This is prin-
cipally owing to the lack of field data on the solitaria
whose cryptic coloration has precluded observation.
The reasons for the occurrence of phase polyphen-
ism in some insects but not others is not known.
According to Rhoades (1985), insects exhibiting
density-dependent phase polyphenism are prone to
sudden outbreaks of high population levels, usually lay
their eggs in clusters, and their larvae are colonial or
semi-colonial at least part of the time. He suggests that
polyphenism enables these insects to adopt two altern-
ative foraging strategies to cope with plant defensive
126
chemicals. A wide range of secondary compounds have
been isolated from plants and many of these are thought
to be feeding deterrents (Coleman & Jones, 1991; Har-
borne, 1993). For instance, grasses (the main food
plants of S. exempta larvae) contain hydroxamic acids,
phenolics and polyphenolics, alkaloids and cyanogenic
glycosides, some of which increase in concentration in
response to plant damage sufficient to deter both ver-
tebrate and invertebrate herbivores (Vicari & Bazely,
1993). Alternative suggestions for the occurrence of
phase polyphenism are that high density populations
are at greater risk from disease and/or exhausting their
food supply, which would enhance selection of forms
capable of rapid development through the vulnerable
larval stage. All of these hypotheses assume that high-
density forms posses physiological and/or behaviour-
al adaptations which endow them with faster rates of
development than the low-density forms although both
laboratory and field data to support this are meagre.
This paper investigates factors determining the
expression of the most striking of the larval phase
characters, cuticular melanisation, in S. exempta and
whether this influences larval thermoregulation and
sensitivity to ultraviolet radiation.
Materials and methods
Insects. The insects were initially derived from a
laboratory colony held at the Natural Resources Insti-
tute, U.K and were maintained under a L14:D10
regime at 27  1  C and ambient humidity. The larvae
were reared either singly or in groups of ten in 250
ml clear plastic pots from the day of hatching unless
otherwise stated and fed fresh wheat seedlings ad lib..
Every experiment used a split-sib design in which all
the larvae were derived from one batch of eggs laid by a
single male/female cross and divided equally between
treatments. Unless otherwise stated all measurements
were made on larvae 2–3 days after they had moulted
to the final (fifth) instar.
Effect of larval density. Larvae were reared singly or
in groups of 2, 3, 4, 5, 15, or 30.
Effect of changing larval density during development.
(i) Larvae were initially reared in groups of ten but then
isolated immediately after moulting to the third or fifth
instar. (ii) Larvae were initially reared in isolation but
then kept in crowded conditions (i.e., in groups of ten
larvae) immediately after moulting to the third or fifth
instars.
Specificity of the phase response. Single S. exempta
larvae were reared in pots containing five S. littoralis
larvae of similar age.
Effect of high and low larval rearing temperature.
Larvae were reared from the second instar at a con-
stant temperature of either 35  C or 20  C.
Thermoregulation. Measurements of internal body
temperature were made on moribund final instar lar-
vae: the larvae were chilled (but not frozen) on crushed
ice, sufficiently to prevent them from recovering, then
used immediately. Active larvae were not used because
they struggled violently making it difficult to restrain
them at a fixed position under the heat source. Fur-
thermore, they usually vomited during the experiment
and this loss of body fluids affected the rates of tem-
perature change. Moribund larvae were weighed then
placed on a flat 6 cm thick block of white polystyrene
(as insulation) and a fine FM-S type thermistor inserted
up their anus until it was midway along the body. Once
the temperature of a larva had equilibrated with that of
the room (21  C) it was illuminated with a 60 W light
bulb, placed 15 cm cm vertically above, and the tem-
perature recorded every 30 s on a Grant SQ 32 logger
for 10 min. Direct irradiation over the spectral range of
the bulb (305–2800 nm) at 15 cm distance was 242.75
W m,2 as measured using a Mini Kipp Dome Solar-
imeter. This is equivalent to weak sunlight. The bulb
had a colour temperature of about 2300 K, with peak
spectral emmittance at a wavelength of about 1100 nm.
Therefore, the light used in the study was ’redder’ than
natural sunlight, which has a colour temperature of
about 6000 K. After 10 min irradiation, the temperat-
ure of the insects had begun to reequilibrate; they were
not irradiated for longer than this because preliminary
experiments indicated that there was a risk of larval
desiccation affecting the results. Non-rellective card-
board surrounds shielded the insects from air currents
during the experiment.
Ability of the cuticle to absorb ultraviolet radiation.
The cuticle of final instar larvae was removed from the
rest of the body tissue in lepidopteran saline, cleared
in 5% w/v potassium hydroxide (Long, 1953), washed
in distilled water, then attached to the outside of a
quartz cuvette (containing distilled water) by surface

tension, and placed in the light path of a scanning UV
spectrophotometer.
Sensitivity to UV radiation. Final instar larvae were
exposed to known levels of ultraviolet radiation
(254 nm) in a Spectronics Corporation XL-1500 UV
Crosslinker and their subsequent development recor-
ded.
Results
Larval coloration. Crowded larvae began to exhibit
dark coloration in their third instar and by the final
instar they all had black bodies with thin yellow lon-
gitudinal stripes and black head capsules (Tables 1 and
2). The black coloration was caused by dense deposits
of regularly-shaped melanin granules within the cuticle
which masked any underlying pigmentation in the body
tissues. Green coloration was primarily a result of pig-
mentation in the haemolymph and the ingested plant
material visible through the gut wall. Isolated larvae
were either green with a central brown dorsal stripe of
variable dimensions or various shades of pink or brown
with pale brown head capsules. Black larvae were not
recorded from isolated cultures in this study although
they have been found in previous experiments.
When two larvae were reared together, many of
them became darkly coloured or black and at higher
densities all the larvae were invariably black (Table 1).
There were no outwardly visible differences in the
degree of melanisation between caterpillars reared at
densities of greater than three larvae per container. Vir-
tually all single larvae reared with S. littoralis larvae
became black (Table 2).
Isolated larvae reared at 20  C had enhanced cutic-
ular melanisation and black head capsules, thereby
closely resembling the typical gregaria (Table 3).
The coloration of the crowded larvae did not change
markedly at low rearing temperature although some
larvae became almost pure black with no longitudinal
yellow stripes. At high rearing temperature, the col-
oration of isolated caterpillars did not change but the
degree of cuticular melanisation of the crowded larvae
was often substantially reduced. Some became grey
rather than black while many exhibited an olive green
colour quite unlike the typical solitaria green.
If formerly crowded larvae were isolated at the start
of their third instar none adopted the green solitaria
coloration although 20% turned dark brown (Table 2).
By contrast, the majority of formerly isolated larvae
127
which were crowded from their third instar became
black. There was no change in body coloration if the
larval rearing density was altered after the moult to the
final instar.
Thermoregulation. The black gregaria caterpillars
warmed up appreciably faster than the green solitaria
(Figure 1). However, the mean weight of the gregaria
caterpillars used in these experiments was significantly
lower than that of the solitaria (356:5  18:95 mg,
n = 31, cf. 493:1  32:13 mg, n = 20, ANOVA,
F1;50 = 4:77, P<0.05)) and analysis of covariance
demonstrated that although larval colour had a signi-
ficant effect on temperature excess (measured after 10
minutes) (F1;50 = 5:27, P=0.026) the influence of lar-
val mass (log10 ) was much more important (F1;50 =
15:58, P<0.001). The relationship between temperat-
ure excess after 10 minutes and larval mass is shown
in Figure 2. The equality of the two linear regression
lines was tested but no significant difference was found
(F1;48 = 0:48, P>0.05; slope of pooled within group
regression = 0.336)
Cuticular absorption of ultraviolet radiation. There
was no significant difference between the cuticles of
the two phases at 210 nm but those of gregaria had
significantly greater absorptions at 220 nm (20.5%,
ANOVA, F1;23 = 21:24, P<0.001), 240 nm (34.7%,
F1;23 = 61:35, P<0.001), 260 nm (38.3%, F1;23 =
75:18, P<0.001), and 280 nm (39.9%, F1;23 = 79:42,
P<0.001) (Table 4).
Susceptibility to UV radiation. There was no differ-
ence in the susceptibility of gregaria and solitaria to
254 nm ultraviolet radiation (Figure 3). The larvae
were able to detect ultraviolet radiation – as evidenced
by their increased locomotor activity when the source
was initially switched on – but they exhibited no extern-
al signs of distress (e.g., vomiting, curling) even at the
highest doses. Larvae continued to eat and move nor-
mally after treatment. Most deaths occurred at the time
of pupation – many larvae failing to produce, or only
partially producing, a pupal case.
Discussion
Previous work has shown that isolated S. exempta lar-
vae do not exhibit phase change in response to their
own frass or that of other larvae (A. Gunn, unpubl.) so
phase is, presumably, principally determined by inter-
128
Table 1. The effect of larval rearing density on the coloration of final instar
larvae of Spodoptera exempta

(n) Larval density

1 2 3 4 5 15 30
(50) (36) (18) (24) (30) (45) (30)

% green 40 14 0 0 0 0 0
% Pink/brown 60 25 0 0 0 0 0
% Black 0 61 100 100 100 100 100

Table 2. The effect of altering rearing density during development on the coloration of final instar larvae of Spodoptera
exempta

(n) Larval rearing conditions

Isolated Isolated Isolated 1 S. exempta + Crowded Crowded Crowded
until L3 until L5 5 S. littoralis until L3 until L5
(n) (21) (47) (15) (18) (50) (20) (36)

% green 95.2 33.3

% pink/light brown 4.8 46.7
% dark brown 8.5 1.3 20.0
% grey 25.5 11.1
% black 66.0 6.7 88.9 100 80.0 100

Figure 1. The effect of larval phase on weight related temperature excess of final instar larvae of Spodoptera exempta.
 129

Figure 2. The relationship between log10 temperature excess of final instar larvae of Spodoptera exempta after 10 minutes and log10 larval
= + = =
mass. The linear regression equations for the two phases are: gregaria y 0:2827x 0:1456, R2 0:1839; solitaria y 0:4018x 0:2005, ,
=
R2 0:3167.

Table 3. The effect of rearing temperature

on the coloration of final instar larvae of
1992). Larvae must be extremely sensitive to contact
Spodoptera exempta because even low larval densities invariably induced
the expression of black coloration. Despite this, high
Temperature ( C) densities of solitaria larvae have been recorded in the
20 27 35 field (Whellan, 1954; Langer & Rose, 1986) although
Isolated (n) (12) (14) (18) this may have been a consequence of the dense veget-
% green 0 7 61 ation restricting larval contact until late in develop-
% pink/brown 8 93 39 ment. The factor(s) responsible for phase change is
% black 92 0 0 clearly not species-specific. Non-specificity may arise
because S. exempta has to compete with other insect
Crowded (n) (34) (25) (38)
species for the same food supply and when this occurs
% olive green 0 0 61
% grey 0 0 18
black coloration is beneficial. A similar, though not so
% black 100 100 21
pronounced, effect is found when isolated Mythimna
separata larvae are reared amongst groups of M. loreyi
or Pieris brassicae (Iwao, 1962).
S. exempta caterpillars have a greater tendency to
larval contact as in Mamestra brassicae (Kazimirova, both adopt and retain the coloration of gregaria phase
130
Figure 3. The effect of larval phase on the susceptibility of final instar larvae of Spodoptera exempta to 254 nm ultraviolet radiation.
larvae than those of the solitaria after their rearing
conditions are altered. This, coupled with the early
expression of black coloration and their readiness to
become black at low population densities, suggests that
it offers important advantages. In fact, it would appear
that it is only when larval densities remain consistently
low throughout development that crypsis is beneficial.
Perhaps under these conditions local populations are
vulnerable to visually hunting predators while at high-
er larval densities the predators are swamped thereby
reducing their importance as a mortality factor.
It is often suggested that dark coloration in insects
performs a thermoregulatory function (e.g., Rhoades,
1985, Goulson, 1994, deJong et al., 1996). Cer-
tainly many caterpillars which bask are darkly col-
oured (Casey, 1976; Knapp & Casey, 1986; Stamp &
Bowers, 1990) and, in laboratory experiments, Sim-
monds & Blaney (1986) found gregaria phase larvae
of S. exempta to spend longer on the exposed upper
surface and edges of their food plant than solitaria.
However, field workers report that gregaria do not
exhibit basking behaviour (J. Klock, pers. comm; K.
Wilson, pers. comm.). This appears strange because
one might expect that a cryptic basking caterpillar
would be able to raise its body temperature to the level
of a non-basking black caterpillar without the associ-
ated risks of greater conspicuity. Unfortunately, field
data for solitaria are not available. The present res-
ults suggest that the dramatic phase-associated change
in coloration does not have as large an impact on the
thermoregulatory properties of S. exempta as might
be expected. However, because gregaria are usually
smaller than solitaria (Gunn & Gatehouse, 1987) they
will have a higher surface area to mass ratio and there-
fore heat up faster. In those species exhibiting density-
dependent phase variation, low-density forms are fre-
quently larger than high-density forms (Iwao, 1968)
and it is therefore important to consider size as well as

Table 4. The effect of larval phase on cuticular absorbtion of ultraviolet light
in final instar larvae of Spodoptera exempta. Results are expressed as means 
=
standard error and compared by ANOVA. n number of cuticles
Wavelength (nm) Absorbance (mean  s.e)
Isolated
=
(n 11)
210 4.01  0.128
220 2.59  0.083
240 1.67  0.051
260 1.44  0.045
280 1.36  0.045
coloration when determining thermoregulatory prop-
erties.
The fact that cuticular melanisation was enhanced
at low temperature, when it might be expected to
improve the absorption of radiant energy, and reduced
at high temperature where dark coloration might lead
to a risk of overheating, would appear to contradict
the above suggestion that larval coloration has a small
influence on thermoregulation. However, it is possible
that, at temperature extremes, the importance of colour
in thermoregulation become increasingly significant.
Because they bask and their wings provide a large
surface area to expose to the sunlight, adult butterflies
provide the most convincing evidence for the role of
colour in thermoregulation. For example, in Colias spp.
butterflies the proportion of black wing coloration has
a pronounced influence on raising body temperature
(Watt, 1968, 1969). However, in other invertebrates,
the evidence is sparse and frequently contradictory
(Casey, 1988). For example, Stower & Griffiths (1966)
were unable to find consistent differences in equilib-
rium body temperature or rate of increase in body
temperature between green and ’red-dark’ morphs of
Schistocerca gregaria. It should also be remembered
that some cryptic lepidopteran larvae bask (Sherman
& Watt, 1973) while many darkly coloured caterpillars
are secretive, avoid light or are nocturnal. For instance,
larvae of S. littoralis reared at high density are dark-
er than those reared in isolation but, at least during
the final instar when melanism is most pronounced
(Hodjat, 1970), both forms avoid the light and tend
to feed at night (Carter, 1984; A. Gunn, unpublished
observations). It is therefore unlikely that in this spe-
cies density-dependent melanisation is associated with
thermoregulation.
Many conspicuously coloured caterpillars are dis-
tasteful to insectivorous birds and mammals (Bowers,
131
F P
Crowded
=
(n 13)
4.13  0.070 0.73 ns
3.25  0.112 21.24 <0.001
2.56  0.094 61.35 <0.001
2.34  0.087 75.18 <0.001
2.25  0.084 79.42 <0.001
1993) but black S. exempta larvae are avidly consumed
by chickens, lizards, amphibians, and praying mantids
with no signs of any repellant effects (Gunn, unpub-
lished observations). Field workers have also repor-
ted that armyworm outbreaks attract large numbers of
birds (Brown, 1962). Black coloration in S. exempta is
therefore not aposematic.
Ultraviolet radiation presents a potential hazard for
any animal living in an exposed position, especially
in equatorial regions. Melanin has been linked with
protection against ultraviolet radiation (Kettlewell,
1973) so the demonstration that cuticular melanisa-
tion reduces the transmission of ultraviolet light, espe-
cially at the 260 and 280 nm wavelengths (the absorp-
tion peaks of nucleic acid bases and proteins), would
suggest that it may serve a protective function in S.
exempta. However, no differences were found between
the survival of green and black larvae subjected to
known levels of 254 nm radiation so this appears to be
unlikely.
These results are at first sight anomolous. Most
workers consider cuticular melanisation in inverteb-
rates to be associated with thermoregulation and/or
protection from ultraviolet radiation. However, in S.
exempta cuticular melanisation appears to have a sur-
prisingly small impact on these factors so it may
be performing other roles which are equally, if not
more, important. One recent suggestion is that density-
dependent melanisation may, amongst other things,
reflect an alteration in the immune status of an insect
(Wilson & Reeson, in press). In both Mythimna (cited
as Pseudaletia) separata, and S. exempta crowded
phase larvae are more resistant to virus infections
(Kunimi & Yamada, 1990; A. Reeson et al. in prep.
unpubl.). Melanisation is important in an insect’s non-
specific defence mechanism (Lai-Fook, 1966) against a
range of pathogens, including viruses (Washburn et al.,
132
1996). Furthermore, St Leger (1992) has suggested that
cuticular melanin may serve as a repellent preventing
fungal pathogens from invading the body. This is not
to suggest that cuticular melanisation in S. exempta, or
other caterpillars, never has more than a minor influ-
ence on thermoregulation but that its role(s) in insect
physiology are probably more complex than is com-
monly assumed.
Acknowledgements
I am very grateful to NRI who supplied the initial cul-
ture and to Andy Reeson and Ken Wilson for discussion
and allowing me to see their experimental data prior to
publication. The work was supported by a grant from
the Nuffield Trust.
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