Activity Patterns, Body Temperature and Thermal Ecology in Two Desert Caterpillars (

Lepidoptera: Sphingidae)
Author(s): Timothy M. Casey
Source: Ecology, Vol. 57, No. 3 (May, 1976), pp. 485-497
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1936433
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Ecology (1976) 57: pp. 485-497

ACTIVITY PATTERNS, BODY TEMPERATURE AND THERMAL

ECOLOGY IN TWO DESERT CATERPILLARS
(LEPIDOPTERA: SPHINGIDAE)"
TIMOTHY

M.

CASEY2

Department of Biology, University of California, Los Angeles, California 90024 USA

A bstract. The activity patterns, body temperatures, microhabitat characteristics and
feeding behavior of larvae of two sphinx moths which inhabit the Mojave desert were
investigated. Hyles lineata occurred in association with and fed upon a variety of desert
annual plants; Manduca sexta was associated with only one plant species, Datura metalloides,
a perennial. Body temperatures of H. lineata were relatively constant over a wide range of
air temperatures;body temperatures of M. sexta approximated the air temperature for most
of the day. Elevated Tb was associated with exposure of the larvae to direct solar radiation
and high ground temperatures. Hvles lineata regulated Tb by orienting with respect to solar
radiation and exploiting thermal heterogeneity in its microhabitat. Manduca sexta was shielded
from solar radiation and thermal reradiation by its foodplant and never voluntarily spent time
on the ground. Feeding and locomotor activity in both species were reduced during periods
of both high and low air temperatures but were not related to time of day. Upper lethal
temperaturein saturated air was 450C for both species. Feeding rates were strongly dependent
on body temperaturein both species. Hyles lineata is a generalist, utilizing a variety of energy
resources; the occurrence of M. sexta in the desert may depend on the presence of a single
plant species, the jimson weed.
Key

words:

Activity

patterns;

body

temperature;

California;

feeding;

Hyles lineata;

Manduca sexta; Mojave Desert; Sphingidae; thermoregulation.

INTRODUCTION

The deserts of the world, despite extreme and fluctuating temperatures and aridity, support a rich and
diverse arthropod fauna. Although a few desert
arthropods are specialized to xeric habitats, the survival of most results from an enhancement of already
existing adaptations for heat tolerance and water
economy rather than a new and special suite of adaptations peculiar to xeric forms (Cloudsley-Thompson
1964, Edney 1967).
The success of arthropods in occupying deserts
is also due to their ability to find and utilize more
equable microenvironments which allow the proper
functioning of their physiological machinery. Behavioral responses that prevent overheating have been
reported for several desert insects. Cicadas (Heath
1967, Heath and Wilkin 1970) and tenebrionid
beetles (Edney 1971) exert partial control over their
body temperatures by shuttling between sunny and
shady areas of their habitat. Harsh climatic extremes
may be avoided by burrowing (Cloudsley-Thompson
1970), and activity patterns may be modified in response to high environmental temperatures (Hadley
and Williams 1968, Holm and Edney 1973).
The white-lined sphinx moth, Hyles (Celerio)
lineata, and the tobacco hornworm, Manduca sexta,
1 Manuscript received 2 June 1975; accepted 2 September 1975.
2 Present address:
Naval Arctic Research Lab, University of Alaska, Barrow, Alaska 99723, USA.

are abundant at certain times of the year throughout

the Mojave desert in the southwestern United States.
Hyles lineata is a cosmopolitan species which experiences periodic population fluctuations in Old and
New World deserts (Grant 1937). Manduca sexta is
a New World species, usually occurring in mesic habitats where it is a major pest on tobacco and tomato
plants (Madden and Chamberlin 1945).
Although lepidopterans are quite numerous in
deserts (Edney 1974), there is little information on
the thermal relations of lepidopteran larvae in deserts.
Caterpillars should be particularly vulnerable to heat
stress because they have limited mobility and do
not burrow. The purpose of this study is to examine
the behavior of caterpillars of H. lineata and M.
sexta in the desert with particular reference to t-heir
activity patterns and responses to ameliorate heat
stress.
MATERIALS AND METHODS

Behavior of fourth and fifth instar larvae of H.

lineata and M. sexta was observed in the Mojave
desert near Lovejoy Butte and near Palmdale, Los
Angeles County, California.
During periods of observation, air temperature
(Tair) and relative humidity were continuously monitored with a shielded, ventilated hygrothermograph
which was calibrated against a U.S. Bureau of Standards thermometer and a sling psychrometer. Relative humidity in the vegetation was measured with a

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486

TIMOTHY M. CASEY

~B

Ecology, Vol. 57, No. 3

)A~~~~~~

t~~~~~~~~~~~~

4.'

I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

NI
occurin

pln

on whc

M.sxai

on

nteMjv

eet

FIG. 1. (A) The white-lined sphinx caterpillar, Hyles (formerly Celerio) linzeafa. (B) The habitat of H. lineata
in May. (C) The tobacco homnworm,Maniduca sexia. (D) Jimson weed, Datura mnetalloides, the only naturally
occurring plant on which M. sexta is found in the Mojave Desert.

thermistor psychrometer. Ground temperature was

measured with a calibrated thermistor.
A silicon cell pyranometer with a spectral response
of 0.35-1.15 u measured solar and sky radiation.
Maximum incident solar radiation at various times
of the day was measured by holding the instrument
perpendicular to the sun's rays. The instrument was
inverted to measure radiation from the ground. Due
to the spectral characteristics of the instrument, this
measurement represents reflected solar radiation only
because thermal radiation occurs at longer wavelengths.
Body surface temperature (Tb) of fifth instar
larvae was measured to the nearest 0.2?C with a calibrated thermistor within 3 s of capture. The caterpillars were held by the head and Tb was measured
on the dorsum of the metathorax. Tests showed that
T,, thus recorded were not significantly different from
values obtained by puncturing the body wall with the
thermistor. Body weight of caterpillars ranged from
2 to 6 g in H. lineata and from 2 to 14 g in M. sexta.

Caterpillars were logged as active if they were

feeding or moving. Activity was gauged at hourly
intervals for H. lineata by observing caterpillars in a
habitat area
50 m2. Caterpillars of M. sexta were
collected and placed on three large jimson weeds
(Datura inetalloides) in the desert at least 24 h before their behavior was recorded. The position of the
caterpillars (horizontal or vertical) on the vegetation
and on the ground was recorded at hourly intervals
and their orientation with respect to solar radiation
was also noted. At night, their activity was observed
under red light. Caterpillars showed no response to
red light but abruptly stopped their activity in response to light from a flashlight.
Mandibular movements of feeding larvae were
timed with a stopwatch and Tb was measured immediately thereafter.
To measure upper lethal temperature a chamber
was maintained at the desired temperature by immersing it in a water bath. The chamber was lined with
moist paper towels to maintain a saturated humidity.

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Late Spring 1976

THERMAL

ECOLOGY IN DESERT CATERPILLARS

L4-

487

1.4

A
Q2 -

1.2 !

E
N

Q~
s

/
0.4 -

0
a:
00,4

OA

0.2

Lo
C40
02

/3

C%( DI

0800

/.

1000

1200

1400

1600

time

60

50

50
30

40

25
40

30

301

20~

30

20

110

0600

0800

1000

time

1200

1400

1600

FIG. 2. Microclimate measurementsat Lovejoy Butte

field site during a day in May 1973. (A) Shaded
circles = solar and sky radiation, unshaded circles =
reflected radiation from the ground. (B) Shaded circles
= air temperature in the shade, unshaded circles ground temperature.
In the chamber Tair was continuously monitored with
a copper-constantan thermocouple connected to a recording potentiometer. Caterpillars were dropped
into a small chamber through a hole in the lid. Once
the larvae were inside the hole was plugged with
cotton. A few minutes were allowed for Tb to approximate Tair. Third instar larvae were used in
these experiments to prevent long equilibration periods.
RESULTS

Local distribution and microclimate

Hyles lineata.--This
species (Fig. IA) is extremely abundant in the Mojave Desert in late spring
in certain years and almost completely absent in other
years. Its abundance in arid regions is correlated with
the magnitude of winter rains which directly affects
the abundance and species diversity of the annual
vegetation (Grant 1937). Lovejoy Butte in mid-May
(Fig. 1B) supports a wide variety of low-growing
annuals which provide little shelter from solar radiation and thermal reradiation from the ground.

20
0600

0800

1000

1200

hime

I
1400

I
1600

20

FIG. 3. Microclimate measurements in Palmdale,

California during a day in July. Symbols same as Fig. 2.
Mean daily maximum and minimum air temperatures for the month of May in the study area are
27?C and 9?C, respectively (Mathias et al. 1968).
Microclimate measurements at the Lovejoy Butte
field site during a day in May 1973 are plotted in Fig.
2.
Manduca sexta.
The occurrence of M. sexta
caterpillars (Fig. IC) in the desert is coincident with
the flowering of jimson weed (Fig. 1D). The caterpillars are present from about June to late September
and their population size is about the same each
summer. Jimson weed grows in disturbed habitat,
most often along roadsides. Its distribution in the
desert is quite irregular and individual plants do not
usually grow in close proximity to each other. The
caterpillars occur in any area of the desert where the
jimson weed grows. Usually plants are occupied by
1-10 caterpillars although up to 60 eggs may occur
on a single plant.
Mean daily maximum and minimum air temperatures in Palmdale in July are 36.8?C and 17.3?C,
respectively (Mathias et al. 1968). During the day,
ground temperatures beneath jimson weeds were
similar to Tair even though ground temperatures of

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488

TIMOTHY M. CASEY

Ecology, Vol. 57, No. 3

100

:00

80

80
0

c
O 60
c
4

60

*
~00 0

040

0
O

Ole

20

-00

800

00:00

25

Time

1D
3

1240

0~~~~~~~~~~~

20

0800

160

80

T;

B 5220

t ?

*
0

20

45

120

0800

160

25Time

0
80

2o-60
0

*60
0

00

40

00

40~

0~~~~~~~~~~~~~~~~~~~~~~
20

00 0

0
0

15

20

25

30

35

40

Air temperature (?C)

FIG. 4.

-2

15

20

20

25

30

Air temDerature

35

40

(?C)

Location of caterpillars of H. lineata in the habitat as a function of the time of day and Tair. (A & B)

Percent of population on the ground. (C & D) Percent of population vertical on vegetation. Unshaded circles
represent a cool day (max Tair = 280C); shaded circles represent a hot day (max Tair = 36.20C). Minimum of
28 animals counted for each point.
up to 60?C were recorded only 1 m away from the
plants (Fig. 3). The large surface area of the leaves
shielded the caterpillars from direct solar radiation,
particularly at midday, and modified the effect of
wind. Relative humidity of air beneath the bush was
similar to that of air outside it.
Feeding behavior and foodplants
Hyles lineata.
Caterpillars fed either while on
the vegetation or while on the ground. In early May
most of them fed on evening primrose, Oenothera
breviceps, consuming flowers, leaves, and tender distal portions of the stems. By 13 May, they had eaten
most of the 0. breviceps in the area, and were feeding
on at least five other desert annuals including buckwheat (Eriogonum), desert dandelion (Malacothrix),
a four o'clock (Mirabilis bigelovii), and sand verbena
(Abronia). By 1 June, all larvae were gone from the
Lovejoy Butte area.
The larvae had no typical feeding posture or position on the foodplants. They occurred on the stems,
petioles, secondary veins, and margins of leaves, and
on the ground. They often dropped off the plant
even though much edible tissue remained. These

caterpillars usually moved to another plant and resumed feeding.

Manduca sexta.--Caterpillars
were found only
on jimson weed, and only on the underside of the
leaves. The caterpillars anchored to the base of the
leaf by their prolegs, and during feeding they grasped
the leaf margin with their thoracic legs (feeding behavior of the species has been analyzed by Heinrich
1971). While feeding they are always at least
partially shaded from solar radiation.
During inactive periods between feeding bouts,
the larvae clung to the leaf's midrib by the thoracic
legs as well as by the abdominal prolegs and were
completely shielded from sunlight.
Orientation and location
Hyles lineata.--The
location of caterpillars in
the habitat varied at different times of the day.
Throughout the night they remained on the foodplants but shortly after sunrise the ground warmed
up and many caterpillars moved to the ground. The
proportion of caterpillars on the ground decreased
during midday (Fig. 4A). During the afternoon the
caterpillars again moved from their position on the

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Late Spring 1976

THERMAL

ECOLOGY IN DESERT CATERPILLARS

100 _

489

45

A
0

40
-

.5

35

50

*0

C6

800

~~~~~ar0

z
-

25
0600

0800

1200

1000

1400

1600

1ime

FIG. 5. The proportion of M. sexta oriented perpendicular to solar radiation at various times of the day.
Sample size for each point = 25.
foodplant to the ground. The proportion of caterpillars on the ground during the day was inversely
related to the air temperature (r = 0.75, p < 0.01,
Fig. 4B).
The orientation of the larvae to solar radiation
varied during the day and between different days.
Early in the morning they oriented perpendicular to
the rays of the sun. At midday their orientation
varied with the air temperature. When Tair was
50% of the caterpillars were horizontal
28?C,
(with respect to the ground). When Tair was 36?C,
87% of the caterpillars were oriented parallel to
solar radiation (Fig. 4C).
Manduca sexta.
Caterpillars remained on the
jimson weed at all times of the day and night. Early
in the morning they were located on the periphery of
the plant. The proportion of caterpillars oriented
perpendicular to solar radiation was greatest in the
morning, decreased during midday, and remained low
in the afternoon (Fig. 5). At about 1000 the larvae
moved from the outer parts of the plant to its interior
where most were shaded from direct solar radiation
for the rest of the day.
If a plant was completely denuded of edible tissue
th- caterpillars usually dropped to the ground and
wandered away. At midday those caterpillars that
remained on denuded Datura plants either oriented
parallel to sunlight or utilized shade provided by the
large bare stems. A caterpillar which had dropped
50
off a denuded bush in the morning was found
m away on some dried grass at noon, oriented parallel
to sunlight. This was the only instance in three summers of observation that a caterpillar of M. sexta
was found in the desert on a plant other than jimson
weed.
Body temperature
Body temperature was not
Hyles lineata.
closely coupled to air temperature during the day in
this species. In the morning shortly after sunrise
caterpillars basked and elevated their body temperatures. Thirty minutes after sunrise mean Tb of 12

0.

20.

1 200

1000

0800

a,L

30

1600

1400

I
1800

*
*

25

T01

35

0500

0700

0900

1100

1300

1500

1700

tiMe

FIG. 6. Body temperature of H. lineata caterpillars

during the day on (A) a warm day and (B) on a cool
day. Shaded circles represent caterpillars on the ground;
unshaded circles represent caterpillars on vegetation.

basking caterpillars was 3.6?C above Tair (SE =0.66,

n =12; Fig. 6) while mean Tb of caterpillars forced
to remain in the shade at that time was 1 .6?C below
n =11 )
(SE =0.39,
'air
The orientation and location of the caterpillars
in the habitat affected body temperature. Animals
on the ground usually had higher body temperatures
than those on the vegetation. At midday the difference between Tb and Tair (AT) was related to the
orientation of the larvae. On a hot day (Fig. 6A)
when most larvae were oriented parallel to sunlight,
the mean Tb was only 1.5?C above Tair (SE - 0.46,
n =15).
On a day when Tair was moderate at midday Fig. 6B ) most caterpillars were perpendicular
to direct sunlight and mean Tb was 8.1 ?C above
Tair. The largest temperature excess recorded was
20?C for a caterpillar on the ground. Mean AT decreased at all temperatures above 20?C (Fig. 7).
After sunset, the body temperatures of caterpillars
were not significantly above the air tempertature.
temperatures were simiManduca sexta.--Body
lar to the air temperature during the day (Fig. 8 and
Fig. 9). In the morning the AT was greatest (x
3.50C, n - 15) and this decreased as Tair increased

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TIMOTHY

490

40

A
A

40

Ecology, Vol. 57, No. 3

M. CASEY
-

M.sexta

350

30

2? 30 0.

25

20

/&

CO 20

15
I

10/o
I

FC%Fo
-o

10

15

* 10H-

16 0

11

0
160

10

200%

I
40

FIG. 8. Body temperature of caterpillars of M. sexta

in the field at various times of the day. Mean values are
plotted and vertical lines represent ? 2 X SE.

5
0
0

I
30

Tair (0C)

10

2
12

20

2
22
1

ing. Feeding decreased during the early morning; by

0400 it had stopped completely and was not resumed
15
20
25
30
35
10
until after sunrise.
Air temperature (0C)
To determine the minimum temperature for feeding, caterpillars were placed in a constant-temperature
FIG. 7. The relation of body temperature to air
temperaturein H. lineata caterpillars. (A) Mean values cabinet. The temperature was raised in small increare plotted and vertical lines represent ? 2 X SE. Dashed ments and the behavior of the animals was noted
line is least squares regression for values of air temperature from 200C to 360C. (B) Mean temperature (Table 1). Minimum Tb for feeding was 14.6?C
excess vs. Ta.r. Numbers above points indicate sample and at least half the larvae fed when Tb was 15.8?C.
size.
In general, the animals were capable of movement at
air temperatures 2-3?C below the temperature at
which
they began to feed. Below air temperatures of
during the day. Body temperature of most cater12.7?C
the animals did not move or respond to
pillars was above the air temperature although Tb of
stimulation.
tactile
small fifth instar larvae which were completely
Manduca sexta.
This species is more sedentary
shielded from solar radiation was often slightly below
H.
than
lineata.
moved slowly to differCaterpillars
Tair. At night Tb was at or slightly below the air
ent
of
and rested frequently
portions
the
foodplant
temperature.
between feeding bouts. Usually there was little or
no response to tactile stimulation even when the
Activity patterns
caterpillars were lifted off the plant. Occasionally
of caterpillars during they jerked the anterior portion of the body but they
Hyles lineata.--Activity
the day varied with the air temperature. At moderate rarely regurgitated or attempted to bite. The proair temperatures the larvae fed continuously during portion of caterpillars feeding at any one time during
the day but at midday when air temperature exceeded
the day in the field was lower than in H. lineata.
Activity was temperature-dependent. There was a
34?C the fraction of caterpillars feeding sharply decreased (Fig. 10 and Fig. 11). Feeding and general marked decrease in the proportion of feeding animals
activity during the day was intense with few rest for about 3 h during the hot part of the day and a
periods between bouts of feeding. Larvae were majority of the animals fed during the rest of the 24-h
highly irritable and responded to tactile stimulation period. The response of activity to air temperature ih
by violently jerking the anterior half of the body M. sexta and H. lineata was similar (Fig. 11). Caterfrom side to side, regurgitating, and attempting to pillars of M. sexta collected in the desert and placed
in a constant temperature of 23?C under a light cycle
bite.
of 12 h light and 12 h dark fed continuously during
After sunset, all larvae crawled onto the vegetation
and continued to feed throughout much of the even- the 24-h period (Fig. 12).
0

0 l

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Late Spring 1976

THERMAL

ECOLOGY IN DESERT CATERPILLARS

491

Lethal temperatures

40 -

17

12

12

12

17
35

17

7
aoir

16
-30

17

The upper lethal temperature for 30-min exposures

of third instar larvae of both species in saturated air
is about 45?C (Table 3). After exposure to 45?C
~~~~~~~~~~~~~~~~~~~~~~~12
the animals lost all motor control and did not respond
to tactile stimulation. Although some coordination
returned when they were placed at lower temperatures, all caterpillars died within 24 h.
DISCUSSION

1200

1000

0800

1400

1600

Body temperature

1800

time

In nature, body temperatures of insects often differ

from air temperature (see, for example, Wellington
1950). Except during periods when flight muscles
are utilized, endogenous heat production by insects is
of negligible importance in determining their body
temperature (Gunn 1942, Church 1960, Heinrich
1974). Solar radiation is the main source of exogenous heat responsible for elevated body temperatures
in nonflying insects (Parry 1951, Shepard 1958),
with high substrate temperatures and thermal radiation being particularly important to insects which are
active on the ground (Hadley 1971, Edney 1971).
Cooling in most cases occurs by convection (Digby
1955, Stower and Griffiths 1966, Edney 1971, Hadley 1971). Tsetse flies (Edney and Barrass 1962)
and sawfly larvae (Seymour 1974) may rely on evaporative cooling for short periods at high air temperatures.
Field measurements of Tb of lepidopteran larvae
indicate that heat responsible for elevated body temperatures is exogenous. Spruce budworms (Choristoneura fumiferana) exposed to full sunlight showed
a mean AT of 7?C, but when partially or completely
shaded had a mean AT of 2.1?C (Shepard 1958).
Temperature excesses of 8-13?C were found in tent
caterpillars in sunshine but these animals had mean
body temperatures within 1?C of air temperature
while they were in the shade or at night (Wellington
1950).

FIG. 9. The relation of body temperature to air

temperature in caterpillars of M. sexta. Sample size
varies from 12 to 17. Mean values are plotted and
vertical lines represent + 2 X SE.

In the laboratory, caterpillars of M. sexta were

capable of movement at temperatures of 8 0C and
above (Table 2). The first sign of feeding occurred
at Tair of 10.4?C. At least half the caterpillars fed
when the temperature reached 140C.
Rates of feeding
Biting rates of the caterpillars in the field were
measured as an index to the rate of feeding. Biting
rates of both species are strongly correlated with body
temperature (p < 0.001; Fig. 13), with those of M.
sexta being slightly higher at a given Tb than those
of H. lineata. The actual feeding rate of caterpillars
depends not only on the biting rates of the caterpillars
but also on the amount of time that they spend feeding. Within each species the proportion of animals
active is similar over a wide range of air temperatures
(Fig. 11). It is therefore assumed that the time spent
feeding over this range of temperatures is independent of Tair. Consequently, the actual feeding
rate should be proportional to body temperature up
to the point where activity is voluntarily reduced by
the caterpillars.

1. Feeding in H. lineata as a function of air temperature.Animals were cooled below temperatureat which
feeding occurs. Temperature in the chamber was raised in small increments. Larvae were allowed five minutes
to equilibrate, after which they were checked at 1-min intervals for 5 consecutive minutes

TABLE

No. feeding larvae/No. nonfeeding larvae

Minutes
Notes

10.0

0/22

0/22

0/22

0/22

0/22

Larvae dormant

11.2

0/22

0/22

0/22

0/22

0/22

Larvae dormant

12.3
14.6
15.7

1/21
1/21
7/15

1/21
1/21
6/16

1/21
1/21
6/16

1/21
1/21
7/15

1/21
1/21
6/16

Four larvae moving sluggishly

Eight larvae moving-not feeding
Movement of larvae coordinated

Tair

(OC)

16.8

11/11

11/11

11/11

11/11

11/11

Most animals moving

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492

TIMOTHY M. CASEY

loo

100

Ecology, Vol. 57, No. 3

A
0

H. lineato

80

80
60

60

000

H. lineata
0

40

40

00

20 _
20

a)
<

.)

80 _B

0-

_o100_

60 _

0
~0

00
0o

00

o0
0

M.
sexto
0

0~~~~~~~

oO

80

40 _
20

M. sexta

0~0

40
20

60 -

l0

40 -

20
30
Air temperature (?C)

40

FIG. 11. The relation of activity in (A) H. lineata

and (B) M. sexta in the desert to air temperature.Sample
size for each point same as in Fig. 10.

20

0600

1000

1400

1800

2200

0200

0600

time

FIG. 10. Activity of hornworms in the desert at

various times of the day (unshaded circles) and night

(shaded circles).

(A) Activity of H. lineata on (C) a

cool day (max. Tair = 280C), (W) on a warm day

(max. Tair = 36.20C), and at night. (B) Activity of
M. sexta in July. Minimum sample size for H. lineata

during the day = 30, at night = 20. Minimum sample

size for M. sexta = 20.

In this study both microclimate and behavior affected caterpillar body temperature. Elevated body
temperatures occurred only when caterpillars were
exposed to solar radiation, high ground temperatures,
or both. At night or when the caterpillars were in
the shade, however, their Tb was closely related to
the air temperature.
The thick foliage of the jimson weed protected M.
sexta from both solar radiation and ground radiation.
The movement of caterpillars from the periphery to
the interior of the plant at midday also tended to
stabilize Tb close to Tair. These caterpillars fed while
effectively avoiding the major heat sources in the
desert.
Caterpillars of H. lineata exploit the thermal
heterogeneity of their microhabitat to maintain a relatively constant Tb. Changes in orientation account
for a tenfold difference in the body surface area ex-

posed to sunlight. In addition, the rate of convective

heat exchange is inversely proportional to an object's
length in the direction of the wind (Bartlett and
Gates 1967, Smith and Miller 1973). Convective
heat loss is maximized when caterpillars are vertical
(with respect to the ground) and minimized when
they are horizontal with their long axis parallel to
the direction of the wind. Consequently, orientation
can be an important means of regulating Tb by varying both the rates of radiative heat gain and convective heat loss.
The thermoregulatory orienting behavior of H.
lineata is similar to that of locusts in the desert
(Fraenkel 1929). Attempts to classify locust orientation into taxes and kineses by Fraenkel and Gunn
(1961) are somewhat confusing because this behavior
is mediated by temperature as well as by light
(Fraenkel 1930). Furthermore, it is difficult to
classify this behavior in caterpillars because while
they are orienting, other types of behavior such as
feeding or movement to and from the ground may
occur simultaneously. The behavior is sufficiently
complex, containing elements of both taxes and
kineses, that such a dichotomous classification would
seem to be oversimplified and arbitrary.
Behavioral thermoregulation is, of course, not
unique to the insects. Such behavior occurs in am-

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THERMAL

Late Spring 1976

ECOLOGY IN DESERT CATERPILLARS

100

0
0

80

0o000

H. lineatg
M. sext

0
0

493

~ 60

40 1

20
0000

I 1

0400

0800

00
0

V
0
(L)
0

1200

1600

2000

~~

(j)

Time

FIG. 12. Activity of M. sexta caterpillars at constant

air temperature. Bar indicates 12-h light-12-h dark
photoperiod utilized. Sample size = 20.

*0

34

20

30

20

10

40

Tb (0C)
FIG. 13. Biting rate of hornworms as a function of
body temperature.

Activity patterns
Activity patterns of lepidopteran larvae are entrained by photoperiod (see review by Beck 1968)
although other environmental stimuli, notably air

temperature, may modify such entrainment. In larvae

of Holisdata argentata (Lepidoptera: Arctiidae) the
ambient temperature determines whether feeding is
diurnal or nocturnal (Edwards 1964). Activity patterns of desert arthropods, both daily and seasonal,
may be modified as a result of temperature (Hadley
and Williams 1968, Holm and Edney 1973). In
addition, Young (1972) suggests that predator pressure is important in the determination of activity patterns of Morpho caterpillars (Lepidoptera: Nymphalidae).
Field data from the present study indicate that
the larvae of H. lineata and M. sexta show little
change in activity with changing light levels over a
24-h period but that both high and low temperature
will modify activity. The season in which these
species occur in the desert determines the periodicity
of their activity. Hyles lineata is inactive in the
early morning because in the spring, air temperatures
are usually below the level at which activity occurs

Feeding in M. sexta as a function of air temperature.

No. feeding larvae/No.

000

ofbo

phibians (Lillywhite 1972, Seymour 1973) and is

quite common among the reptiles (Cowles and Bogert
1944, Heath 1965, Bartholomew 1966, DeWitt 1971,
White 1973). In these groups the behavior is qualitatively similar. When cool, animals maximize surface
exposed to solar radiation and when heat stressed they
minimize surface exposed to solar radiation or seek
shelter. In amphibians, this behavior increases the
rate of development (Lillywhite et al. 1973). Since
the biting rate of caterpillars is directly related to
TI, (Fig. 13), any elevation of Tb above Tair should
result in an increased rate of food intake. In caterpillars such as M. sexta in which Tb approximates
Tair, the rate of development increases directly with
air temperature (Madden and Chamberlin 1945,
T. M. Casey, personal observation). In Colias caterpillars (Lepidoptera: Pieridae) the rates of development and feeding increase with body temperature.
However, they decline when caterpillar temperatures
rise above the preferred body temperature (Sherman
and Watt 1973). This may be due to activity, for in
H. lineata and M. sexta the amount of time spent
35?C.
feeding decreased when T1) exceeded

2.

~ ~

00

TABLE

00

2400

Method same as in Table 1

nonfeeding larvae

Minutes
3

Notes

7.1

0/19

0/19

0/19

0/19

0/19

8.2
9.3
10.4
11.5
12.7
13.9

0/19
0/19
2/17
5/14
7/12
10/9

0/19
0/19
2/17
5/14
6/13
10/9

0/19
0/19
2/17
5/14
7/12
9/10

0/19
0/19
3/16
5/14
7/12
10/9

0/19
0/19
2/17
5/14
7/12
11/8

All larvae dormant, lying on side

with no righting response
Larvae upright, sluggish movement
Larvae upright, sluggish movement
Locomotion poorly coordinated
Majority of larvae moving
All animals moving
All moving; majority feeding

Tair (?C)

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494

TIMOTHY M. CASEY

3. Upper lethal temperatureof desert hornworms.

Third instar larvae were exposed to various air temperatures in saturated air for thirty minutes. n = 4
at each temperature

TABLE

Temperature
(OC)

Percentage alive 24 h after exposure

H. lineata
M. sexta

42
44

100
100

45

50

46

100
100
0

(Table 1). In May, air temperatures during the day

are hot enough to restrict activity only occasionally.
Manduca sexta occurs in the desert later in the summer where high midday temperatures regularly reduce
their activity, but air temperatures at night remain
moderate so that activity during the night may be
continuous.
The capacity to feed whenever the air temperature
is not extreme should be advantageous to both species
of caterpillars because they must process large quantities of food. For example, a caterpillar of M. sexta
processes about 1400 cm2 of tobacco leaves (85%
of which occurs in the fifth instar) in completing its
development (Jones and Thurston 1970). Manduca
sexta larvae have been observed feeding both night
and day in Kentucky tobacco fields although studies
of their activity patterns are lacking (Dr. R. Thurston, personal communication). The larvae of H.
lineata are somewhat smaller than those of M. sexta,
and they probably do not process as much food to
complete their development. Nevertheless, the temporal limitations of the food source make it advantageous to feed whenever possible.
Feeding behavior
It is impossible to say with certainty whether differences in the observed feeding behaviors of H.
lineata and M. sexta are due to genetic differences
or to the environment because their foodplants are so
different. However, the behavior of these caterpillars
in mesic habitats suggests that the differences are
species specific. Orientation for temperature regulation is unlikely in M. sexta because they feed while
attached to the underside of leaves. Hyles lineata
actively orients with respect to solar radiation while
feeding and this type of temperature regulation is
particularly effective in relation to the open growth
form of its foodplants. Eliot and Soule (1902) reported that in Vermont H. lineata fed at midday while
exposed to direct solar radiation, noting that such
feeding behaivor was atypical for sphingid caterpillars.
The two species of caterpillars also differ with regard to inactive periods. A black color phase of H.

Ecology, Vol. 57, No. 3

lineata basks on bare ground (Forbes 1948). In the

desert, H. lineata is exposed to sunlight and often
spends time on the ground during inactive periods.
Manduca sexta remains sheltered when inactive, however, and never leaves its foodplant while edible tissue remains (see McFadden 1968).
Ecological and evolutionary considerations
Although both H. lineata and M. sexta are common in the same habitat, they are active at different
times. Different factors will determine their distribution and abundance and they exhibit a number of
behavioral differences (Table 4).
The feeding behavior of M. sexta insures that the
entire edible portion of each leaf is consumed (Heinrich 1971). Economical harvesting of the foodplant
is critical to M. sexta caterpillars because jimson
weed is sparsely distributed in the desert and the
larvae must often complete their development from
food provided by a single plant.
Feeding behavior of H. lineata maximizes the rate
at which nutrients are harvested from its environment. Behavioral temperature regulation allows the
caterpillars to maintain body temperatures at which
feeding rate is highest. Such behavior is adaptive
because although the desert annuals are very abundant, they are present for only a short time. Increased
rates of feeding should shorten the duration of the
larval stage of the life cycle, helping insure that the
caterpillars complete their development before their
food source dries up.
Birds prey upon both species of caterpillars (Thurston and Prachuabmoh 1971, Stewart 1969, Miller
and Stebbins 1964) and the striking differences in
feeding behavior and coloration between these species
may reflect different evolutionary responses to predation. Manduca sexta is cryptically colored and by
occupying the underside of leaves, the larvae remain
inconspicuous (Fig. 1A; see Wigglesworth 1972, fig.
12 and fig. 21).
Hyles lineata is brightly colored and conspicuous.
The caterpillars varied from bright yellow to jet
black. The importance of such coloration in this
genus has been the subject of much controversy. Cott
(1940) infers from the larvae's irritating habit of
regurgitation when disturbed that Hyles euphorbiae
larvae exhibit warning coloration. Beddard (1895)
argues that the conspicuousness of Hyles galii is of no
advantage because the caterpillars have been shown
to be palatable to predators.
Development of different color patterns in caterpillars within a population may be related to food
quality and to more subtle intraspecific interactions
(Schneider 1973). Since caterpillars of H. lineata
regulate body temperature by basking, the coloration
of H. lineata larvae, particularly the darker forms,

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Late Spring 1976

THERMAL ECOLOGY IN DESERT CATERPILLARS

TABLE

4.

Comparison of aspects of the biology of two desert caterpillars

Hyles lineata

Distribution
Abundance
(Mojave Desert)
Abundance
(mesic areas)
Foodplants in desert
Microhabitat
Coloration
Thermoregulation
Feeding time
(final instar)
Feeding behavior
Foodplants
(distribution)

495

Manduca sexta

Cosmopolitan; mesic and xeric

habitats (Grant 1937).
Very abundant in April and May
but absent the rest of the year.
Population size varies drastically
from year to year.
Late spring to early autumn
(Dickerson 1901).
Polyphagous; at least six plants
eaten at field site.
Plants provide no shelter from

New World; most abundant in mesic

habitat (Madden and Chamberlin 1945).
Present in low numbers from June to
late September. Population size similar
at Palmdale field site during summers
of 1972, 1973, 1974.
Same as in the desert.

sunlight; Tair moderate.

Tair often high.

Various, conspicuous.
Well developed.
Continuous when Tair not extreme.
Independent of photoperiod.
Mobile; feeds while on ground and
at midday while exposed to sunlight.
Extremely abundant in space but not
in time; unpredictable (year to year).

Cryptic.
Poorly developed.
Same as H. lineata.

may have thermoregulatory significance. Darker

forms should be able to attain a higher Tb and potentially spend a greater portion of the day at preferred body temperature (for further discussion, see
Hamilton 1971, Edney 1974). Further data are
needed to substantiate this hypothesis.
In deserts, many years may intervene between outbreaks of H. lineata (Grant 1937) and their value as
a food source may be too unpredictable to allow
predators to key on them as a major food source.
During outbreaks the caterpillars are so abundant
that predator populations are likely to be saturated.
Temperature regulation could convey a selective advantage to H. lineata by decreasing the larval stage
duration and therefore the amount of time that the
caterpillars are exposed to predators.
Larvae of M. sexta utilize jimson weed for nutrients, energy, water, and predator protection via
camouflage. In addition, even though M. sexta is
present in the desert at the hottest time of the year,
the foodplant provides a more equable microenvironment for the larvae. The specificity of M. sexta for
solanaceous plants is chemical and is shown not only
by the oviposition behavior of the adults (Yamamoto
et al. 1969), but also by the feeding preferences of
the caterpillars (Yamamoto and Fraenkel 1960,
Waldbauer and Fraenkel 1961). In the Mojave
desert jimson weed is the only noncultivated solanaceous plant on which M. sexta has been collected. It is
therefore probable that the occurrence of M. sexta
in the Mojave desert depends on the presence of
jimson weed.
Hyles lineata is preadapted to the desert because it

Found only on jimson weed.

Plants provide shelter from sunlight;

Sedentary; never feeds while on ground.

Feeds from underside of leaf (shaded).
Not abundant, present from May to October; predictable (year to year).

can feed on any of several foodplants. In years of

rich blooms of desert annuals, H. lineata adults move
into the habitat in great numbers, laying eggs. Large
numbers of larvae utilize the resource as long as it is
available. When the plants dry up, emerging adults
probably migrate out of the desert into more mesic
habitats. This species is quite mobile and in the Old
World regularly migrates north as the seasons progress (Grant 1937), and there are indications that
the North American subspecies may also exhibit such
a migratory pattern (Corfe 1938). An opportunistic
response to temporary local bursts of primary productivity coupled with a nonspecific food preference
and behavior which maximizes feeding rates, and a
highly mobile adult stage contribute to the abundance
and ubiquity of H. lineata.
ACKNOWLEDGMENTS

I am grateful to George A. Bartholomew for his

guidance throughout the course of this study, and particularly for his editorial comments and criticisms. I also
thank E. B. Edney, F. Engelmann, K. A. Nagy, and F. N.
White, for reading the manuscriptand providing stimulating discussions.
I am indebted to my wife, Kathy, for her kind assistance in all phases of this work and to C. Henne, Associate in Entomology, Los Angeles County Museum of
Natural History, for many kindnesses during the course
of my fieldwork.
Financial support for this study from National Science
Foundation grant GB32947 (administered by George A.
Bartholomew) is gratefully acknowledged.
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496

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