Oecologia

Oecologia (Berl) (1981) 50:250-255

9 Springer-Verlag 1981

Field Analyses of Insect Heat Budgets:

Reflectance, Size and Heating Rates
P.G. Willmer and D.M. U n w i n
Department of Zoology, Downing St., Cambridge

Summary. This paper outlines simple techniques for determining

rates of heat gain and loss in relation to the weight and reflectance of insects caught in their natural habitats. In particular
the construction of a new 'reflectometer' is described. The results
thus obtained permit estimates of the relative importance of
size and colour in determining rates of heat exchange and temperature excesses, so allowing better predictions of heat budgets
for a given species.

Introduction
With a few exceptions, insects are classically regarded as ectothermic animals, whose body temperature (and hence activity)
is determined largely by the energy received. The thermal economy of an individual should thus be related to its behaviour
(the time spent in insolated zones, and orientation to incoming
radiation and to air currents), to its body form, and to ambient
conditions. However, literature analysing the actual temperatures
of insects in their natural habitats is sparse, so that discussions
are inevitably speculative and often involve extrapolations from
experiments with model insects (Parry 1951) or with a few insects
from laboratory cultures u n d e r simulated conditions (Digby
1955). A l t h o u g h the present d o c u m e n t a t i o n of insect heat budgets is so poor, there has been m u c h recent work on their thermal
ecology and energy balance (cf. Edney 1971 ; Casey 1976; Dreisig
1980; and review by M a y 1979), and though in some cases
this has included spot measurements of body temperatures m a n y
other studies have relied on assumptions a b o u t rates of heat
exchange, and the effects of size, shape or colour thereon, which
are poorly substantiated. There is thus a recognized need for
a realistic consideration of the effects of insect size (weight,
a n d / o r linear dimensions) and of colour (determined by emissivity or reflectance of the surfaces) on the rates o f heating and
cooling of insects, and on the temperature excesses which they
m a y attain in sunny conditions. This paper analyses such factors
for a range of c o m m o n insects. It describes a device, suitable
for field use, for rapid and accurate measurements of the reflectance properties of insect surfaces, which in c o m b i n a t i o n with
an equally adaptable microbalance can provide information
a b o u t the relative importance of reflectance and of size in determining the thermal properties of different species, thus giving
data u p o n which assumptions a b o u t temperature budgets may
more readily be based.

Methods
a) Field Techniques
Initial recordings of insect weights and refleetances were made from
fresh specimens, caught in their natural habitats while feeding on
Offprint requests to: P.G. Willmer

0029-8549/81/0050/0250/$01.20

flowers or resting in vegetation, in an area of sandy pine forest in

the Breckland area of Norfolk. Specimens which were damaged in
the net, or which were too small for accurate weighing with the available equipment ( < 2 mg) were discarded. All others were killed rapidly
with ethyl acetate, and weighed using a specially constructed field
microbalance (Unwin 1980). Their thoracic breadth was measured using a calibrated 7 x magnifying eyepiece, and their thoracic reflectance
determined (see below). (The dorsal surface of the thorax was used
for both linear measurements and for reflectance, as it overlies the
flight muscles and is thus the most crucial zone for thermal balance
and activity.) All the measurements required could be completed within
5 rain of the death of the insect. Many of the specimens were then
stored and allowed to dry, for subsequent more elaborate laboratory
tests of reflective properties.
Rates of heating and cooling, and temperature excesses, were recorded in the field in natural sunlight with a large number of the
freshly killed insects. These tests were all conducted during a sequence
of hot sunny days, at times when incoming radiation was high and
constant, and wind-speed very low. Extremely fine copper-constantan
thermocouples (0.081 mm wires) were used to minimise conduction
to and from the specimen, and these were connected to a portable
multi-channel indicator (Unwin 1980), each run involving a set of
three matched thermocouples. These were glued with a fine coating
of gum tragacanth to the insects' thoracic terga, or were inserted
through the cuticle to lie just within the underlying flight muscles;
the two techniques gave indistinguishable results. Insects were always
tested in pairs, with the third sensor recording air temperature. A
standard run involved successive 2-5 min periods in full sun and in
shade (provided by white reflective sheeting), the transitions being
repeated several times. Radiation was recorded with a thermopile solarimeter (Unwin 1980), and air temperature and humidity were periodically checked with a wet and dry bulb psychrometer (Unwin 1980).

b) Laboratory Tests and Equipment

The surface reflectance of insects was determined using a newly-designed device (" reflectometer"), incorporating a source which attempts
to mimic the effects of sunlight and a sensor with matched responses
sited to take account of both direct reflection and the diffuse reflections
from a microsculptured surface.
The construction of the reflectometer is shown in Fig. 1. The insect
is held against a small (2.8 mm) hole in a brass plate which is painted
matt black. The light source is a 2.5 volt 'lens-end' torch bulb, supplied
with 3 volts from a pair of torch batteries. "Over-running" the light
source in this way gives a 'whiter' output than that of a normal
tungsten lamp, and the light is thus a better approximation to sunlight.
Light is focussed onto the centre of the hole by a lens of about
20 mm focal length. Radiation reflected from the insect then illuminates the inner surface of a dome, made from half of a table-tennis
ball and painted matt white inside. The surface brightness of the inside
of the dome is measured by a pair of ORP12 cadmium sulphide photoconductive cells, connected to a meter circuit as in Fig. 1 B. To reduce
any effects of stray light from the source reaching the photo-conductive
cells directly, the lens system is mounted in a thin-walled metal tube,
matt black internally, fixed through the apex of the dome; and the
outside of the dome is also blackened to absorb and eliminate direct

251
Table 1. Reflectance and colour characteristics of a range of insects
Insect species

1 mA
~Lens

bulb

i Filtel

~
~176

..

Lens

tql

ORP'12 ' ~'

,I plate
Specimen

9v 9v

]
Fig. 1 A and B. Construction (A) and circuitry (B) of the reflectometer
input from the light source. The whole array may then be mounted
in a light tight box, with a small access-port for the specimens to
be tested.
The device may be calibrated with standard reflective surfaces
(Kodak), adjusting the 1 K variable resistance (a 10-turn helical potentiometer) to give an appropriate meter reading: the response is effectively linear. For general use, the spectral response of the instrument
with components as described is found to give a reasonable match
to that of the sun's radiation. But the overall response is skewed
somewhat towards the red end of the spectrum, and for applications
where this is important it could be corrected by the use of a filter
such as the Wratten 38. When used with small animals such as insects,
the exact spectral composition of the source is not likely to be crucial
(Digby 1955), and this precaution was omitted. However, some laboratory measurements of reflectance to different wavelengths were made
with selected insects, and for such purposes a range of filters can
be inserted as required between the bulb and the dome assembly
(Fig. 1 A). In some cases, this necessitated the use of a more powerful
fibre-optic light source and the substitution of a 50 gA meter for
the 1 mA meter shown in Fig. 1 B to improve sensitivity at low light
levels.
Once calibrated, the instrument was generally used to give a single
broad-band reflectance reading, by holding the thoracic tergum of
an insect against the hole in the sensor hemisphere and rotating the
animal gently to allow for any directional effects on reflectance. A
mean figure for percent reflectance could then be taken, though in
no case was the change in value of this parameter greater than 1.5%
during small movements of the thorax, except when the surface was
obviously striped; for such animals, the reflectance of the dominant
bands was determined.
For all specimens where a broad band reflectance reading was
obtained in the field immediately after death, the value subsequently
found in laboratory tests with the dried out animal was within 1%
of the original figure. Consideration of colour effects could therefore
be reasonably continued using some of these dried specimens and
the spectral properties of their thoracic surfaces were analysed using
a series of filters (Ilford) of known properties, covering the ultra-violet
to infra-red range. In every case the reflectometer calibration was
first checked with the standard surfaces.
Results

A. Reflectance and Colour

As an initial test of the efficacy of the reflectometer, a wide
range of thoracic reflectances of freshly-killed insects were recorded and compared with their subjectively determined
' c o l o u r ' . This information is given in Table 1, arranged as a
series of increasing reflectance. Darker insects had low reflectances (absorbing 90-98 % of incoming radiation), while the paler

Reflectance
%

Dorcus parallelopipedus
2
Chelonus sp.
2
Eristalis pertinax
2-3
Chloromyia formosa
2 3
Delia sp.
2-3
Chrysogaster hirtella
2 3
Alophora hemiptera
3
Poecilobothrus nobilitatus ~ 3
Hilara sp.
3
Arge gracilicornis
3
Dolichopus wahlbergi
3-4
Morellia aenescens
3-4
Delia sp.
3-4
Volucella bombylans
4
Poecilobothrus nobilitatus ~ 4
Melanostoma scalare
4
Cerceris arenaria
4
Dolichopus festivus
4
Calliphora vicina
4
Conops flavipes
4
Lonchaea sp.
4
Tenthredo acerrima
4 5
Lucilia caesar
4-5
Paravespula vulgaris
4-5
Dasyphora cyanella
4-5
Melangyna cincta
4 5
Necrophorus sp.
4 8
Psen equestris
5
Chrysotoxum cautum
5-6
Bombus terrestris
5-6
Helophilus pendulus
5-7
Bombus pascuorum
5-8
Hylemya strenua
6
Syrphus ribesii
6 7
Crabro cribrarius
7-8
Tachina fera
7-8
Psallis varians
7-9
Episyrphus balteatus
7-10
Lygocoris lucorum
8
Meliscaeva cinctella
8-10
Apis mellifera
8 10
Oxycera trilineata
9-10
Rhagio lineola
9-10
Macrotylus pakyulli
9-11
Cantharis livida
10
Xyphosia sp.
11
Ophion sp.
14
Scathophaga stercoraria
14
Chrysopa sp.
15
Opomyza sp.
16
Tricholauxania praeusta
17-19
Rhagonycha fulva
20-23

Dominant thoracic
colour
Black matt
Black matt
Dark brown
Blue-green metallic
Black matt
Black
Dark Brown-black
Blue-green metallic
Black
Dark blue metallic
Bronze metallic
Dark grey-green
Dark grey
Dark brown, paler hairs
Red-purple metallic
Black shiny
Black
Green metallic
Blue-green metallic
Black shiny
Black shiny
Black
Green metallic
Black
Green metallic
Dark bronze
Black & red
Black shiny
Brown
Brown, pale hairs
Grey & brown
Brown, pale hairs
Grey
Bronze
Black shiny
Grey-brown
Brown
Bronze
Green-brown
Yellow-bronze
Orange-brown
Pale green
Pale brown
Green
Yellow-brown
Pale orange-brown
Orange
Yellow-brown
Turquoise
Yellow-orange
Pale orange
Orange

forms, usually in the light brown/orange/yellow colour range,

reflected a higher percentage of the light and absorbed only
70-90%. The apparent anomalies in this scheme were the metallic
insects, which though appearing paler and shiny nevertheless
had very low reflectances. These results match those of Digby
(1955), obtained using different and more time-consuming techniques, and indicate that the reflectometer device does give meaningful results. They also suggest that since even very pale insects
still absorb at least 75% of received b r o a d - b a n d radiation, with
the majority absorbing or transmitting more than 90%, colour

252
2/,

og 2

Metallic

Non-metallic Insects

Insects

2C

el- Euglossa

{3.s)~

,o

18

iL

30
o 28
C26

~81 Xyphosia

91 Halictid

'I

A. Diptera (3mg)

2~

/'

22
20
18

38

:[

r'.

i~.[ Oxycera

(9.5)

'~1

9 9

36'

9v

3/,

/\

9 oo

30

:rv
8~-

oo

O c 32

9 o

t
~ Dolichopus

241" Rhagonycha

vo

":oo

"oO /~

26

o"too,o ,,oo
Wavelength

9
o

.~

2a

2~
22
20

:t

9 oo
%9
~

vVV

% % ~ 1 7 6 1 7 ~6

o%Oooooo

n..

**

vvVvvVvVTVv9

18
0

nm

Fig. 2. Reflectance at varying wavelengths of metallic and nonmetallic

insects. Figures in parentheses refer to the broad-band reflectance
values, and arrows indicate the apparent colour

in itself is unlikely to be a crucial factor in thermal balance

for most temperate insects. However, some desert beetles are
white in colour, or at least have white elytra, and a reflectance
of 74% has been recorded for such forms (Rficker 1933); the
calculations of Edney (1971) also indicate that very high values
may occur for white insects.
To further test the characteristics of the reflectometer, a range
of insects were subjected to lights of varying spectral properties.
The results of these analyses are shown in Fig. 2. For each
of the ' n o r m a l ' insects, whose colour arose from pigmentation
of the cuticle or its pubescence, the reflectometer gave entirely
predictable results, with peak reflectance corresponding to the
observed colour of the species (and usually with a high reflectance in the ultra-violet range also). But this clearly did not
apply for the metallic forms, where the reflectance at different
wavelengths showed no consistent relation with apparent colour ;
indeed there was often a trough of reflectance at the wavelength
corresponding to the colour seen by the human eye. This effect
presumably arises from the different methods of colour production in metallic insects (by scattering, diffraction or interference
effects: see Fox and Vevers 1960; Chapman 1969).

B. Temperature Changes of Insolated Insects

As examples of the temperature changes recorded in test insects,
three sets of paired runs are shown in Fig. 3, for small (A),

~.-- ---=.-.~
,~,~H'~..~.... -----",2 f; ~
2

Time

3
(m)

dae
(90-100 mg).

Fig. 3A-D. Thoracic temperatures versus time for insects in sunlight

(radiation strength 840 900 Watts/m2).) and in shade (2 min intervals;
hatched bars show shaded periods)9 The solid lines show air temperatures, and in each case solid symbols represent the less reflective insect
of a pair, A (o) Xyphosia sp., r = l l % , (y) Hilara sp., r=3%. B
(o) Rhagonychafulva, r=22%, (y) Cantharis livida, r=10%. C (o)
Crabro cribrarius, r=8%, (y) Cerceris arenaria, r=4.5%

medium (B) and large (C) insects. In each case, insects of similar
size and shape but different reflectauces were paired. Several
effects are apparent. The temperature excess (T.ex) clearly depends principally upon size, but is also progressively more dependent on reflectance as the size increases. Similarly the half-times
(T1/a) of temperature change increase for larger insects, (there
were no significant differences between half-times for heating
and cooling), and values again are modified by reflectance in
the upper size ranges.
Data of this kind from all the tests performed is summarised
in Table 2, where values of T.ex and T1/2 are shown together
with reflectance and size, for insects now arranged systematically.
Correlations of temperature parameters with weights and reflectances are shown in Fig. 4. Both temperature excess and T1/2
were very highly correlated with insect weight (4A and 4B),
and correlations with the recorded linear dimension (thoracic
breadth) were almost equally good (cf Table 2). On the other
hand the correlations of temperature changes with reflectance
were poor, and Fig. 4C and D give little indication of any predictable effect of colour on thermal changes. These findings
are again comparable with those of Digby (1955), who suggested
that colour probably contributed ' n o t more than some 25%'
to the temperature control of insects. At the most, Fig. 4 might
indicate that the more reflective insects do not show high values
of T.ex or T1/2, but even this apparent effect arises partly because all the very reflective insects tested were fairly small. In

253
Table 2. Size, reflectance and thermal characteristics of freshly-killed insects. The temperature excess in full sunlight, and the half-time to
r . ex/2
achieve this, are given together with the rate of heating K calculated as ~=c/z/z

* = pubescent insects ; ** =metallic insects

Weight
mg

Thorax
breadth
mm

r
%

Temp.
excess
~

T 1/2
s

K
~

9.7
9.0
3.6
21.0
34,0
9.5
42.0
105.0
82.0
110.0
118.0
39.0
18.0
75.5
48.0
47.0
20.0
6.0
16.5
14.5
15.0
3.5
8.5
7.0

2.4
1.8
1.1
2.4
2.7
1.7
3.0
4.6
3.2
4.8
5.0
3.2
2.5
4.2
3.2
3.0
1.9
1.4
2.0
1.8
1.9
1.2
1.7
1.5

2 3
9-10
3
7-10
6-7
4
2-3
2 3
5-7
5-6
4
4-5
4 5
4
7-8
3
3M,
6
2-3
3 4
14
4
16
17-19

1.4
1.8
0.5
2.6
4.0
1.6
4.0
10.0
7.2
9.0
12.1
3.5
2.2
6.2
5.4
6.0
3,7
1.4
3.2
3.0
2.7
0.8
1.0
0.7

6
14
18
19
7
14
64
44
52
60
20
15
53
26
20
9
10
10
18
12
-

0.117
0.113
0.130
0.105
0.114
0.143
0.078
0.081
0.086
0.100
0.087
0.073
0.058
0.104
0.150
0.206
0.160
0.150
0.075
0.042
-

52.0
5.0
17.5
90.0
102.0
117.0
93.0
155.0
190.0

2.5
1.2
2.2
3.4
3.6
3.9
4.0
5.9
6.8

4-5
2
14
7-8
4
4-5
8-10
5-8
5 6

8.2
1.8
1.4
7.9
10.3
11.5
7.5
12.0
14.0

18
14
40
32
44
46
85
115

0.228
0.050
0.098
0.161
0.131
0.104
0.071
0.061

27.0
26.0
130.0
220.0

2.7
2.5
5.9
10.0

20 23
10
4 8
2

3.2
4.4
11.6
16.5

27
17
58
115

0.065
0.126
0.100
0.072

2.5
4.0
3.8

1.0
1.7
1.3

9 11
8
7-9

0.3
0.4
0,3

9.0

2.0

0.7

11

0.032

Diptera
Chloromyia formosa **
Rhagio lineola
Poecilobothrus nobilitatus **
E pisyrphus balteatus
Syrphus ribesii
M e l a n o s t o m a scalare
Chrysogaster hirtella
Eristalis pertinax*
Helophilus pendulus*
Chrysotoxum cautum
Volucella bombylans *
D a s y p h o r a cyanella**
Lucilia caesar**
Calliphora vicina**
Tachina fera
Atophora hemiptera
Morellia aenescens
Hylemya strenua
Delia sp.
Delia sp.
Scathophaga stercoraria
Lonchaea sp.
Opomyza sp.
Tricholauxania praeusta

Hymenoptera
Tenthredo acerrima
Chelonus sp.
Ophion sp.
Crabro cribrarius
Cerceris arenaria
Paravespula vulgaris
Apis mellifera
Bombus p a s c u o r u m *
Bombus terrestris*

Coleoptera
R h a g o n y c h a fulva
Cantharis livida
Necrophorus sp.
Dorcus parallelopipedus

Herniptera
Macrotylus pakyulli
Lygocoris lucorum
Psallis varians

Neuroptera
Chrysopa sp.

fact, t h e p a r a m e t e r s o f size a n d reflectance are n o t t h e m s e l v e s

i n d e p e n d e n t v a r i a b l e s ; t h e c o r r e l a t i o n coefficient for the s a m p l e
u s e d w a s 0.29 ( n = 4 1 ; p < 0 . 1 0 ) .
T o s e p a r a t e t h e effects o f size
a n d c o l o u r t h e r e f o r e p r e s e n t s s o m e difficulties, as m u l t i v a r i a t e
r e g r e s s i o n a n a l y s i s is n o t strictly applicable. N e v e r t h e l e s s , a
r o u g h g u i d e to t h e relative c o n t r i b u t i o n s o f t h e t w o p a r a m e t e r s
c o u l d be a c h i e v e d b y s u c h a n a n a l y s i s ; this i n d i c a t e d t h a t c o l o u r
c o n t r i b u t e d o n l y a b o u t 10% to t h e d e t e r m i n a t i o n o f T . e x in

15

t h e s a m p l e o f insects u s e d , t h o u g h t h e figure rose to 2 2 % if

o n l y t h e larger ( > 80 m g ) insects were c o n s i d e r e d .
A f u r t h e r m e t h o d by w h i c h t h e c o n t r i b u t i o n o f reflectance
to t h e r m a l b a l a n c e c a n be e x a m i n e d is to u s e t h e v a l u e s o f
K c a l c u l a t e d in T a b l e 2 as t h e initial rate o f h e a t g a i n or loss
(T.ex/2/T1/2); this p a r a m e t e r h a s a relatively restricted r a n g e
o f values, a n d is i n d e p e n d e n t o f insect w e i g h t (r = 0.18). V a l u e s
o f K are p l o t t e d a g a i n s t reflectance in Fig. 5, a n d t h e c o r r e l a t i o n

254
o

,6

lO0

14

o
o

12

Tex

le

80

Oo

T1/2
o

oc

o
60

o~

S.

o
6

o
o

40

o o
o

r =0.971
4

o~
~,

o ~

io

'

'

100

150

Weight

200

2JO

16

5'o

,~,o

mg

1~o

200

mg

Weight

r =0.951

o ~176
o oO
o o

20

lOG
14

o
o

12

o
u

Texto

8C

TY2
o

oC

2
0

oo

o %0
40

o
o

o
o
oo

o
o

,
5

o o

o
o

S.

I
0

o
o

6C

2r

o
n

10

15

Reflectance

,
20

i
25

~
o

o~
~176

,
5

o
o

10

15

Reflectance

20

Fig. 4. Effects of weight and reflectance on temperature excess and half time of temperature change in insolated insects

is reasonably good; more reflective forms clearly gain and lose

heat less rapidly. Figure 5 also shows up the anomalies more
readily, in that both pubescent and metallic forms appeared
to change their temperatures more slowly than predicted. For
pubescent insects this can be attributed to the insulative effects
of the hairs and of trapped still air (Church 1960). But the
metallic forms appear to be genuine anomalies, and the use
of simple reflectance recordings for such insects may be inappropriate; their thermal characteristics are in every case consistent
with higher reflectances than those recorded.

.20

.15

Discussion

.IO

.05

O~LI

i
2

i
4

I~

[]

i
6

I
8
Reflectance

I
I0

I
12

i
14

i
16

l
!8

i
20

~o

Fig. 5. Rate of change of temperature (K) as a function of reflectance.

The regression line shown was calculated for normally pigmented insects (o) only. Metallic insects (|
and pubescent insects ([]) are
distinguished separately

This paper describes apparatus and techniques suitable for determining size, reflectance, and the heating effects of sunshine for
insects in natural conditions. Both the reflectometer and the
simple microbalance used have proved to be of practical value,
giving results when used in the field comparable to those of
earlier laboratory studies, and also showing up some of the
anomalies which occur. The present study is limited, though,
in that it ignores some of the factors which may be important
determinants of thermal ecology (see Willmer 1981). Of the environmental parameters, windspeed is probably the most crucial
variable which has been excluded from consideration here, and
this factor should be particularly important in altering the temperature excess for the smaller insects whose size lies in the
range where natural convection is supplanted by forced convec-

255
tion currents (Digby 1955). However, such small insects will
spend much of the time contained within the boundary layer
of still air around the surfaces on which they rest, so that macroclimatic windspeed may have little practical effect on the cooling
of many inactive insects. A second important effect which has
not been considered, since these tests involved recently dead
insects, is that of activity itself: 'metabolic heat'. Any active
insect produces heat internally in relation to the cube of its
linear dimensions, whereas radiative heat gain varies with L2;
hence metabolic heat is relatively less important for a small
insect. Larger insects therefore generally heat up more during
flight, whereas smaller forms show little change in body temperature when they take off or may even cool down slightly if evaporative and convective losses exceed the heat generated from the
flight muscles (Digby 1955).
Thus, although the range of factors controlling insect body
temperatures is large, and their interrelations are complex, a
knowledge of the two parameters of size and reflectance, in
conjunction with good microclimatic data, can be of considerable
predictive value in considering insect temperatures. The good
correlations of weight and linear dimension with thermal gains
(Fig. 4) accord with earlier studies, while the lesser but significant
effects of colour are more clearcut and predictable from the
present data, (controversies in the earlier literature on this issue
are reviewed by Edney 1971). Based on such data, simple predictions may be made about the thermal balance of insects. Large
insects will both attain, and maintain, high temperature excesses
fairly easily. If they are also dark in colour (low reflectance),
they will reach a high temperature more quickly, so that a large
dark insect will be at an advantage at dawn and dusk, when
it may more readily reach a temperature conducive to activity.
At greater radiation levels in the middle of the day smaller
insects will also be capable of activity, though they may be
restricted to sunlit patches while the larger forms extend their
range into the shade. At very high temperatures smaller insects
may be at a clear advantage, since the larger ones, with temperature excesses of up to 15-30 ~ C (present study, and see Heinrich
1975), may be in danger of overheating during prolonged exposure to direct insolation; whereas small insects, having lower
temperature excesses anyway, can also take advantage of very
local patches of shade more readily and will cool down more
quickly therein. When overheating is a real danger, as may be
the case in tropical or desert situations, insects may do better
if highly reflective, and this should be especially valuable for
insects of moderate or large size which are active during the
day. Hence large pale or shiny insects are relatively common
in the tropics, but rare in the temperate fauna, whereas large
desert insects are commonly either black (if crepuscular) or shiny
white (Hamilton 1975).
Clearly these predictions represent gross generalizations, and
there are obvious exceptions; in some cases these can be traced
to known physiological adaptations such as antifreezes (Salt
1969) or endothermie regulatory mechanisms (Heath et al. 1971 ;
Heinrich 1974; May 1979; Willmer 1981). But there are some
available studies, particularly where these consider related insects
of similar shape, which accord well with the predictions outlined
here; those of Heinrich (1976), Schlising (1970) and Willmer
and Corbet (1981), all on bees or wasps, are suitable examples.
Studies of all insect types active at particular sites through a
day (Willmer in prep) also support these suggestions. Furthermore, the few available studies of different eolour forms of a
single species (eg Watt 1968; Cena and Clark 1972) provide
good evidence for the thermal effects of colour in isolation from
other factors, as do the known cases of colour change within

an individual in response to temperature (Key and Day 1954;

O'Farrell 1963). The use of the described field techniques to
measure these crucial parameters should therefore prove useful
in further studies of insect activity cycles in the field, and of
the hygrothermal considerations which underlie then.
Acknowledgements. We would like to thank F.S. Gilbert for helpful

discussions about this manuscript, and C.P. Ellington and Dr. W.B.
Amos for loans of equipment. PGW is also grateful to the Childerhouse
family for permission to work on their land and use their facilities,
and to New Hall, Cambridge for continued financial support.

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