Feeding Behavior 357

ENDOCRINE CONTROL OF use at the molting periods are the characteristics of the larval stages.
FAT BODY METABOLISM During the prepupal period, mass quantities of reserve material are
accumulated in the fat body cells. Lysis of fat body cells in higher
Neuroendocrine secretions from brain and ganglia, ecdysteroids
dipteran species at metamorphosis results in the discharge of stored
(molting hormones), JHs, and the myriad corpora cardiaca neuro-
reserves into the hemolymph. However, as the new adult fat body
secretions affect the metabolic state of the adipocytes. These endo-
cells are formed, nutrients are reabsorbed. In contrast, in lepidop-
crine secretions are strongly influenced by stimuli from internal and
teran species in which cell remodeling occurs, a status quo prevails.
external environments, and they function to coordinate and integrate
Fat body cells in adults are depleted of reserve materials, which are
crucial metabolic activities involved in molting, growth, metamor-
used for locomotion and reproduction.
phosis, and reproduction. The fat body is a target tissue for endo-
crine regulation as is illustrated shortly. Stored glycogen and proteins
are mobilized during the molting process to form the newly synthe-
See Also the Following Articles
sized cuticular chitin–protein complex. The blood level of trehalose Excretion ■ Juvenile Hormone ■ Metabolism ■ Vitellogenesis
is regulated by a corpora cardiaca neurohormone. The adipokinetic
hormone from the corpora cardiaca stimulates the adipocytes to Further Reading
release diglycerides and the accompanied lipoprotein carrier, and
Akiduki, G., and Imanishi, S. (2007). Establishment of a lipid accumulation
F
enhances lipid oxidation to fuel flight in favor of carbohydrate oxi-
model in an insect cell line. Arch. Insect Biochem. Physiol. 66, 109–121.
dation. Synthesis and release of vitellogenins by the female fat body Dean, R. L., Locke, M., and Collins, J. (1985). Structure of the fat body. In
cells are usually under the control of JH, although in certain insect “Comprehensive Insect Physiology, Biochemistry, and Pharmacology” (G.
species the molting hormone is also involved. A. Kerkut, and L. I. Gilbert, eds.), Vol. 3, pp. 155–210. Pergamon Press,
Oxford, U.K.
Haunerland, N. H., and Shirk, P. D. (1995). Regional and functional differen-
FAT BODY DURING DEVELOPMENT AND
tiation in the insect fat body. Annu. Rev. Entomol. 40, 121–145.
METAMORPHOSIS Keeley, L. L. (1985). Physiology and biochemistry of the fat body. In
During the period of metamorphosis, the fat body tissue undergoes “Comprehensive Insect Physiology, Biochemistry, and Pharmacology” (G. A.
extensive morphological, histological, biochemical, and organizational Kerkut, and L. I. Gilbert, eds.), Vol. 3, pp. 211–248. Pergamon Press,
changes. These processes are triggered by the molting hormone on the Oxford, U.K.
Law, J. H., and Wells, M. A. (1989). Insects as biochemical models. J. Biol.
background presence of extremely low levels of the JH. Such altera-
Chem. 264, 16355–16638.
tions have been thoroughly studied in dipterans and lepidopterans.
Two major strategies for transforming the larval fat body into an adult
tissue exist: (1) the histolytic pathway, in which the larval fat body adi-
pocytes in dipteran species are completely histolyzed and the adult new
tissue is formed from undifferentiated stem cells, and (2) the remod-
eling pathway, in which adipocytes in the larval stage of lepidopteran Feeding Behavior
insects dissociate at metamorphosis into individual cells before being
reassociated into the adult new tissue. In certain holometabolous insect R. F. Chapman†
species, a combination of the two processes takes place. University of Arizona
Dynamic exchanges of nutrients between fat body cells and the

T
hemolymph compartments are evident throughout the life cycle of he most extensive studies of insect feeding behavior and its reg-
holometabolous insects (Fig. 1). Buildup of reserves and their partial ulation have focused on two insects with completely different
feeding habits: the adult black blow fly (Phormia regina), which
is a fluid feeder, and the final nymphal stage of the migratory locust
Adipocytes Hemolymph (Locusta migratoria), a grass-feeding insect. Although their feeding
Stored reserves habits are different, there are many common features in their feeding
Larval stages
behavior patterns and the mechanisms that control their feeding behav-
ior. It is likely, though unproven, that similar principles are involved in
other insects, and this article focuses on these two insects, making refer-
ence to other species and other feeding habits where data are available.
Prepupal period V. G. Dethier initiated and guided the earlier work on the blow fly, sum-
marized later by Stoffolano. The studies on the locust were initiated by
R. F. Chapman and developed by E. A. Bernays and S. J. Simpson.
Cell lysis*
At metamorphosis
Cell remodeling
WHY INSECTS HAVE DISCRETE MEALS
Although casual observation may give the impression that insects
feed nonstop, many insects eat in discrete meals separated by peri-
Adult stage ods of nonfeeding (Figs. 1A and 1B). This is most extreme in species
that feed on vertebrate blood: larval Rhodnius prolixus take a single
FIGURE 1 Exchange of stored reserves between fat body cells meal in each developmental stage, and adult female mosquitoes usu-
and hemolymph during the life cycle of holometabolous insects. ally have a single blood meal associated with each vitellogenic cycle.
Asterisk indicates that later, as stem cells are differentiated into adult
†
fat body cells, a buildup of reserves occurs. Deceased May 2, 2003.
 358 Feeding Behavior
(A) Manduca
(B) Locusta
0 1 2 3
Time (hours)
FIGURE 1 Patterns of meals taken by a caterpillar and a locust.
Periods of feeding are shown by black bars. Notice that short pauses
sometimes occur within meals, but the intervals between meals are
F markedly longer. (A) A final stage larva of M. sexta feeding on tobacco
leaves. [Reproduced, with permission, from Reynolds, S. E., Yeomans,
M. R., and Timmins, W. A. (1986). The feeding behaviour of caterpillars
on tobacco and artificial diet. Physiol. Entomol. 11, 39–51.] (B) A final
stage nymph of L. migratoria feeding on wheat. [Reproduced, with per-
mission, from Blaney, W. M., Chapman, R. F., and Wilson, A. (1973).
The pattern of feeding of Locusta migratoria. Acrida 2, 119–137.]
Nectar-feeding insects and phytophagous insects also feed in discrete
meals, but the degree to which this is true in other insects has not
been investigated. For predatory insects, a single prey item is com-
monly not sufficient to produce satiation, and it is likely that a “meal”
would involve several prey, just as a “meal” for a nectar-feeding
insect involves foraging from a number of flowers because no single
flower contains sufficient nectar to produce satiation.
The underlying causes of this behavior are probably both physi-
ological and ecological. Energy is expended in acquiring food and ini-
tiating digestion so that, when food is first ingested, there is a net loss
of energy. Subsequently, as food is digested and absorbed, there is a
gain in resources and energy, but as the process continues the rate of
gain declines and the net gain plateaus as digestion and absorption are
completed (Fig. 2). Consequently, there is an optimal period for which
Start of End of
feeding feeding
Optimal
retention time
Gain of
energy and
nutrients
Loss of
energy Time
FIGURE 2 Nutrient and energy returns associated with eating a
discrete meal. At first, the insect expends energy in obtaining its food.
As the food is digested, nutrients are absorbed increasingly rapidly
but as the nutrients are removed from the food the nutrient return
decreases. To optimize the rate of nutrient return, most insects have
discrete meals, with intervals between meals that approximate to peak
rates of return. [Reproduced, with permission, from Sibly, R. M.,
and Calow, P. (1986). “Physiological Ecology of Animals.” Blackwell
Science, Oxford.]
an insect should retain food in its gut before replacing it with newer,
undigested food. Thus, it is advantageous for the insect to eat a discrete
amount of food (a meal) and process it before taking more food.
The risk of predation has almost certainly had a major role in shap-
ing feeding behavior. Predation risks are much higher during feeding
presumably because of the movements made by the insect that can be
detected visually or mechanically by potential predators. For example,
genista caterpillars, Uresiphita reversalis, feed for only about 3% of
the day, yet 80% of predation by anthocorid bugs occurs during this
period. Similarly, although tobacco hornworm caterpillars (Manduca
4
sexta) on tobacco plants in a greenhouse fed for only about 7% of the
time, 20% of predation occurred during this period.
Grasshoppers typically move away from a feeding site following a
meal, sometimes backing down into the mass of vegetation and remain-
ing unmoving and hidden until the time to feed again approaches. The
caterpillars of U. reversalis move into silken shelters between meals.
Most blood-sucking insects leave the host as soon as they are replete,
usually moving to shaded places where they are inconspicuous, and
it is probably true that most insects move away from the immediate
area of feeding where food-related cues might reveal their presence to
predators.
As a consequence of feeding in discrete, relatively short meals, the
time spent feeding by most insects is only a small proportion of the
available time; for most of the remainder they remain inactive and
presumably minimize predation risks. Blood-sucking insects, which
commonly ingest more than their own weight of food in a single meal,
feed for less than 1% of the time; nectar-feeding butterflies and flies
(feeding on unlimited supplies of nectar in the laboratory) feed for
up to 14% of the time and this is also true for grasshoppers, both in
the laboratory and in the field. All these insects have part of the gut
modified for temporary food storage. Final-stage caterpillars of the
tobacco hornworm spend about 35% of the time feeding in the field.
In grasshoppers, the reduction in activity after feeding is controlled,
at least partly, by a hormone released from the corpora cardiaca at
the end of a meal. Hormonal release is induced by distension of the
crop at this time.
Phloem-feeding homopterans appear to differ from most other
insects. Planthoppers and aphids do not have discrete meals and ingest
food more or less continuously. The phloem provides a continuous sup-
ply of sugars and free amino acids, requiring little or no digestion, so
the availability of nutrients for absorption remains virtually unchanged
over time. Under these circumstances the physiological necessity of
eating discrete meals is eliminated. In these insects, the act of feed-
ing is not associated with obvious body movements because once the
feeding tube is plugged into a phloem sieve tube, the insect remains
in one place for hours. This probably applies to xylem-feeding insects,
which also need to process very large amounts of fluid because of the
low concentrations of nutrients in xylem. Filter-feeding aquatic insects,
such as some mosquito larvae, also probably feed continuously.
THE START OF FEEDING
As the time from the previous meal (the intermeal interval) gets
longer, the likelihood that the insect will respond to food stimuli
increases. A locust starts to move again and so the likelihood of
encountering food is increased. Other factors, not related to the food,
may also further increase the probability of feeding. In a locust, a
sudden increase in light intensity or the act of defecation may have
such effects. Conversely, an encounter with a highly unpalatable food
source may delay the start of feeding and careless movements by an
observer may have a similar effect. Simpson demonstrated that, in
 Feeding Behavior
the migratory locust, there was in addition a tendency for meals to
begin with some pattern of regularity which, in his observations, had a
period varying from 12 to 16.5 min in different individuals. This does
not mean that feeding or some other activity occurred every 15 min,
but when it did so it was usually at some multiple of 15 min from
a set time, which he determined to be lights-on in his experiments
(Fig. 3). There is now evidence for similar rhythms in the caterpillars
of an arctiid moth, Grammia geneura, and the sphingid M. sexta. The
evidence for the latter is based on field observations, and the rhythm
had a period of 3–4 min. The discovery of these rhythms was depend-
ent on detailed, long-term observations on individual insects. Such
sets of observations are rare, and the extent to which similar rhythms
occur in other insects is not known because observations are lacking.
(A)
Lights
Feed
on
Oscillation with
14.5 min period
(B)
30
Frequency
20
10
0
6 4 2 0 2 4 6
Minutes before Peak of Minutes after
peak oscillation peak
FIGURE 3 Oscillation underlying the feeding behavior of the
migratory locust, L. migratoria. (A) Feeding record of an individual
during a 12-h light phase. Notice that each meal begins close to the
peak of a 14.5-min oscillation. (B) Times at which feeding started rel-
ative to the peak of the oscillation for eight insects on one 12-h day.
[Reproduced, with permission, from Simpson, S. J. (1981). An oscil-
lation underlying feeding and a number of other behaviours in fifth-
instar Locusta migratoria nymphs. Physiol. Entomol. 6, 315–324.]
The effects of these varying factors on feeding can be accounted
for by an, as yet hypothetical, excitatory state in the central nervous
system, first postulated for the blow fly and subsequently elaborated
for the migratory locust (Fig. 4). Only when the central excitatory
state exceeds a certain threshold can feeding occur, but feeding is not
an automatic consequence of reaching the threshold; it is a proba-
bilistic event. At the end of a meal, the central excitatory state is
assumed to be depressed below threshold. As time since the previous
meal increases, so does the level of the central excitatory state so that
it approaches and ultimately exceeds threshold. Rhythmic changes
in the central excitatory state are presumed to account for the basic
rhythmicity of feeding, and other events, such as defecation, may tem-
porarily elevate it, while others (disturbance) may depress it.
359
Level of feeding excitation
Feed Nibble
Feeding
threshold
0 60 120 180 240
Time (min)
FIGURE 4 Model of the control of feeding in a locust eating F
wheat. Similar principles are believed to apply to other insects. The
irregular line shows the level of feeding excitation (the central exci-
Lights
off tatory state). When this exceeds a threshold, the insect feeds. Notice
that after a meal, the excitatory state declines sharply. Subsequently
it rises slowly and the level oscillates with a period of about 15 min.
Defecation (upwardly pointing arrows) causes a sudden rise in exci-
tation. If this causes excitation to exceed the threshold, the insect
may feed. Biting the food (oblique arrows) releases juices from the
food and phagostimulants cause a sharp rise in the central excitatory
state. [Reproduced, with permission, from Simpson (1995).]
THE SIZE OF A MEAL
The size of a meal, assuming the food supply to be unlimited,
depends on the net phagostimulatory effects of the food and the nutri-
ent requirements of the insect. Net phagostimulatory effect is the bal-
ance between nutrient components of the food that stimulate taste
receptors leading to feeding and any other factors, such as the presence
of toxic compounds or undue hardness, that tend to inhibit feeding. For
many insects, sugars are major phagostimulants and higher concentra-
tions in the food result in larger meals. Amino acids may also influence
the phagostimulatory input. Plant secondary compounds, such as alka-
loids, are often feeding deterrents even for insects, such as the blow fly,
whose food does not normally contain them. Inorganic salts at higher
concentrations are also deterrent although at low concentrations they
may stimulate feeding. Phagostimulatory compounds do not just switch
on feeding that then continues until the insect is replete; their con-
tinued input is necessary for feeding to continue and the behavior of
some insects appears to reflect this. Chemosensory receptors usually
adapt within a few seconds if continually stimulated, but the palpation
behavior of grasshoppers and caterpillars appears to permit a continual
flow of information by bringing the receptors into contact with the food
for frequent very brief periods. The sensilla on the palp tips of a locust
make about 10 contacts per second and each contact may be only 10–
20 ms. As a result they remain largely unadapted.
Although continual stimulation is important to maintain feeding,
meal size seems to be determined by the level of phagostimulation
when the insect first bites into its food and releases the internal flu-
ids containing a mixture of stimulating chemicals. This was demon-
strated by an experiment in which the mouthpart chemoreceptors of
locusts were stimulated with a highly phagostimulatory solution that
they were not allowed to ingest. These insects subsequently ate larger
meals than others stimulated with water alone, despite the fact that
during the meal the receptors of both sets of insects were equally
 360 Feeding Behavior
stimulated. Events before feeding started determined how much was
eaten. Comparable experiments have shown that distasteful com-
pounds can reduce meal size. Such experiments are interpreted as
reflecting changes in the central excitatory state. A high concentration
of phagostimulant is believed to elevate the central excitatory state
well above threshold and feeding continues, provided some level of
input is maintained, until the excitatory state declines to threshold.
Thus, the higher the initial level, the longer it takes the excitatory
state to reach threshold and the larger is the meal. It is supposed that
a high level of deterrent compounds would inhibit feeding by depress-
ing the level of the central excitatory state below threshold.
The elevated level of the central excitatory state is also believed to
account for the “dances” of flies and the palpation behavior of locusts
and grasshoppers following loss of contact with food early in a meal.
F When a fly loses contact with a drop of sugar, it moves in an irregu-
lar path with frequent turns as if “searching” for the food. The more
concentrated the solution, the more frequent the turns (Fig. 5A).
(A)
(B)
FIGURE 5 Searching for food after loss of contact during a meal.
(A) Blow flies dance when they lose contact with a drop of sugar.
After experiencing more concentrated sugar solutions the high rate
of turning is much more sustained. [Reprinted, with permission,
from Dethier, V. G. (1957). Communication by insects: Physiology
of dancing. Science 125, 331–336. Copyright 1957. American
Association for the Advancement of Science.] (B) The migratory
locust palpates when it loses contact with a blade of grass. Each bar
represents the percentage of time palpating after losing contact with
the food at a different stage of the meal. Soon after the start of a
meal it palpates for most of the time, but toward the end of a meal
it is less persistent. The open bar represents the time palpating just
before starting to feed. [Reprinted, with permission, from Bernays,
E. A., and Chapman, R. F. (1974). The regulation of food intake by
acridids. In “Experimental Analysis of Insect Behaviour” (L. Barton
Browne, ed.). Springer–Verlag, Berlin. Copyright Springer–Verlag.]
If a locust loses contact with its food, it palpates vigorously and such
behavior lasts longer if loss of contact occurs earlier in a meal. Toward
the end of a meal, loss of contact with the food results in only a lim-
ited period of palpation (Fig. 5B). The so-called searching behavior of
other insects, such as that described for coccinellid larvae when feed-
ing on aphids and temporarily losing contact with the prey, probably
has a similar basis.
THE END OF A MEAL
The end of a meal is ultimately determined by the degree of dis-
tension of the part of the gut in which the food is temporarily stored.
In grass-feeding grasshoppers and nectar-feeding flies, this temporary
store is the crop (part of the foregut). In R. prolixus, feeding on ver-
tebrate blood, the food is stored in the anterior midgut. In each case,
the degree of distension is monitored by some form of stretch recep-
tor. In locusts, these receptors are multipolar cells on the wall of the
foregut, and receptors on the most anterior part of the foregut, which
is the last part to fill; these receptors are responsible for inhibiting fur-
ther feeding. R. prolixus has chordotonal organs in the body wall. The
input from these stretch receptors has an inhibitory effect on feeding,
presumably by leading to a decline in the level of the central excita-
tory state to below threshold. If food quality and the nutritional and
feeding status of the insect are constant, stretch receptor input deter-
mines that an insect ingests a similar amount of food at each meal.
CHANGES IN FEEDING BEHAVIOR
The pattern of feeding changes with the age of the insect, its pre-
vious experience, and its nutritional needs. Phytophagous insects
in general tend to eat greater amounts in the middle of a develop-
mental stage and more in the light than in the dark. The average
meal size taken by the final-stage nymph of the migratory locust, for
example, increases from about 50 mg on the day of molting to almost
100 mg 4 days later, whereas the average interval between meals
declined from 82 to 71 min. At night, the insects take fewer meals
even though the temperature may be constant.
Some phytophagous insects become less selective if they expe-
rience a long period without food and this has given rise to some
confusion in the literature. For experimental purposes, it is often
convenient to use insects that feed readily when presented with food.
This is achieved by depriving them of food, often for 24-h periods.
However, because such insects are less selective than insects with
continual access to food, grasshoppers, for example, were generally
considered to be unselective in their choice of foods. More critical
observations, however, show that this is not accurate. With increasing
periods of food deprivation, several grasshoppers have been shown
to accept a wider range of food plants. It is probable that this accept-
ance of previously unacceptable plants reflects a need for water
rather than for other specific nutrients, although this hypothesis has
not been thoroughly investigated. It is, however, clear that a well-
hydrated locust actively moves away from wet filter paper, whereas
a dehydrated one attempts to eat it. Similarly, dehydrated flies drink
water, whereas hydrated ones do not.
The tendency of grasshoppers and caterpillars, and probably other
insects, that are deprived to sample food that would otherwise be
rejected can play a major part in the subsequent acceptance of food.
This becomes possible because taste receptors that initially signaled
rejection because of some distasteful component of the food become
habituated and are no longer stimulated by the distasteful compound.
At the same time, detoxifying enzymes are probably mobilized within
 Feeding Behavior 361

the insect, providing it with the capacity to minimize any harmful
effects that the compound might have.
The nutritional requirements of insects vary through life and this
is reflected by changes in their feeding behavior. During larval devel-
opment, the amount of food consumed is usually maximal in the mid-
dle of each developmental stage, falling to zero for a period before
each molt. Changes also occur in adults in relation to somatic devel-
opment and, in females of many species, in relation to egg develop-
ment. This variation is illustrated for adult red locusts (Nomadacris
septemfasciata) in Fig. 6. When the insect first becomes an adult the
cuticle is soft and the flight muscles are poorly developed. During
this teneral period, both sexes feed actively during the day (Fig. 6A).
Subsequently, the insects enter reproductive diapause and feeding is
reduced to a single meal each day (Fig. 6B). During the reproduc-
tive period, females eat much more than males (Fig. 6D).
Among some adult flies and grasshoppers, there is good evidence
that mature females change their feeding behavior to acquire protein
for the synthesis of vitellogenin. This is most obvious in blood-sucking
flies, such as mosquitoes and tabanids, females of which use nectar as a
flight fuel, but vertebrate blood as their primary protein source. Males
of these same species feed only on nectar. This is also true of blow flies.
Mature female grasshoppers, given the opportunity, tend to select food
with a higher protein level than do males or immature females. Thus,
they tend to eat the seed heads of developing grain rather than foliage.

(A) (B)
F

(C) (D)

FIGURE 6 Variation in feeding behavior during adult life. The red locust, N. septemfasciata, in the field. (A) April. Soon after becoming
adult both sexes feed for most of the day. This is a period of somatic growth when flight muscles and cuticle become fully developed. (B)
September. Despite moderately high temperatures during the day, very little feeding occurs until late afternoon. The insects are in repro-
ductive diapause. (C) October. Little feeding occurs in the middle of the day, perhaps because of the high temperature. The insects are
beginning to become sexually mature. (D) December. Females eat much more than males during the period of egg development. All these
samples were taken from the same generation and population of insects, which live for about 9 months as adults. Each graph shows the
amount of food in the foreguts of a sample of insects taken at each time point over a 24-h period; 100% would indicate that all the locusts
were full, 0% that they were all empty. When the temperature is 30°C or above, the foregut becomes more than half empty within an hour,
so that crop fullness above 50% during the day indicates recent feeding. At 25°C and below, the food takes several hours to leave the foregut
so that night time values largely reflect feeding before dark. [Reproduced, with permission, from Chapman, R. F. (1957). Observations on
the feeding of adults of the red locust. Br. J. Anim. Behav. 5, 60–75.]
 362
Under laboratory conditions, when fed on artificial diets, locusts and
caterpillars are able to select from the foods with different amounts of
proteins and carbohydrates to maintain an appropriate balance of the
two classes of compound. Locusts can make the adjustment from one
meal to the next, with an interval of less than an hour between meals.
The extent to which insects can fine-tune their nutritional balance
when feeding on natural food with much smaller deficiencies of protein
or carbohydrate has yet to be demonstrated.
FEEDING BEHAVIOR UNDER
NATURAL CONDITIONS
In the field, feeding behavior is determined to a large extent by
environmental factors, although relatively few extensive studies have
been carried out. Temperature has a major effect on feeding behav-
F ior, as it does on other insect activities, with little feeding occurring
at low or at very high temperatures (Fig. 6A and 6C). The effects of
temperature are most obvious in insects living under extreme condi-
tions of low or high temperature. For Gynaephora groenlandica cat-
erpillars living within the Arctic Circle, feeding is possible only when
the insect has raised its body temperature by basking. As a result,
most feeding occurs in a relatively narrow window of time around
noon each day, when the sun is highest in the sky (Fig. 7).
100
Basking
75
% of larvae
Movement
50 of sun
25
Feeding
0 burning sensation. The term “fire ant” actually applies to a group of
Midnight 06.00 Noon 18.00 Midnight
Time of day
FIGURE 7 Feeding is limited by temperature. Most caterpillars of
G. groenlandica, living within the Arctic Circle, feed during a 2-h win-
dow when the sun is at its zenith. For most of the time, the insects
bask to raise their body temperatures, enabling them to feed effi-
ciently; during feeding their temperature falls rapidly. [Reproduced,
with permission, from Kukal, O., Heinrich, B., and Duman, J. G.
(1988). Behavioural thermoregulation in the freeze-tolerant Arctic
caterpillar, Gynaephora groenlandica. J. Exp. Biol. 138, 181–193.
Copyright Company of Biologists.]
Darkness also tends to reduce feeding. For many visually forag-
ing insects, finding food at night is impossible, although night-flying
moths obtain nectar only during darkness and some blood-sucking
insects, such as mosquitoes, feed most actively at night or in the cre-
puscular periods. These insects locate their host primarily by odor,
although night-blooming flowers often also present conspicuous tar-
gets because of their size and whiteness.
Biotic factors may also have a profound effect. For example, a cat-
erpillar of M. sexta that has defended itself from the attack of a tachi-
nid fly does not feed for some time after it has successfully repelled
the attacker.
Fire Ants
See Also the Following Articles
Blood Sucking ■ Mouthparts ■ Salivary Glands
Further Reading
Bernays, E. A. (1997). Feeding by lepidopteran larvae is dangerous. Physiol.
Entomol. 22, 121–123.
Bernays, E. A., and Simpson, S. J. (1995). Control of food intake. Adv. Insect
Physiol. 16, 59–118.
Chapman, R. F., and de Boer, G. (1995). In “Regulatory Mechanisms of
Insect Feeding” Chapman & Hall, New York.
Dethier, V. G. (1976). “The Hungry Fly.” Harvard University Press,
Cambridge, MA.
Simpson, S. J. (1995). Regulation of a meal: Chewing insects. In “Regulatory
Mechanisms of Insect Feeding” (R. F. Chapman, and G. de Boer, eds.),
pp. 137–156. Chapman & Hall, New York.
Simpson, S. J., Raubenheimer, D., and Chambers, P. G. (1995). The mecha-
nisms of nutritional homeostasis. In “Regulatory Mechanisms of Insect
Feeding” (R. F. Chapman, and G. de Boer, eds.), pp. 251–278. Chapman &
Hall, New York.
Stoffolano, J. G. (1995). Regulation of a carbohydrate meal in the adult
Diptera, Lepidoptera, and Hymenoptera. In “Regulatory Mechanisms
of Insect Feeding” (R. F. Chapman, and G. de Boer, eds.), pp. 210–247.
Chapman & Hall, New York.
Fire Ants
Les Greenberg
University of California, Riverside
M
ost residents of the southeastern United States are very
familiar with fire ants. These reddish brown ants are well
known for their aggressiveness and stings that produce a
New World ant species in the genus Solenopsis. Many people refer
to them as “red ants,” although this term is also used to refer to the
larger red harvester ants found in desert climates. In Spanish, the fire
ant is sometimes called hormiga colorada or hormiga de fuego, and
in Portuguese it is formiga de fogo. In North America, there are four
native fire ant species, two introduced species and two hybrid forms.
The two imported species in the United States are the red imported
fire ant, Solenopsis invicta, and the black imported fire ant, S. rich-
teri. S. invicta has spread throughout the southeastern part of the
country, whereas S. richteri is restricted to northeastern Mississippi,
northern Alabama, and northwestern Georgia. Seventeen fire ant
species are currently described from South America.
ORIGIN AND SPREAD
There seems little doubt that the most important fire ant pest,
S. invicta, traveled from South America to Mobile, Alabama, in ship
ballast between 1930 and 1940. It spread in all directions from there,
limited only by cold winters or desert drought conditions. S. richteri
may have arrived earlier (perhaps in 1919), only to be largely dis-
placed by S. invicta. The latter is currently established in 12 states
(Florida, Georgia, Alabama, Mississippi, Louisiana, Texas, North
Carolina, South Carolina, Arkansas, Tennessee, Oklahoma, and
California) and Puerto Rico.