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Elements of Behavior

Behavior is what animals do. It can be defined more precisely as an internally directed system
of adaptive activities that facilitate survival and reproduction. Ethology is the scientific study of
animal behavior -- particularly when that behavior occurs in the context of an animal's natural
environment. Ethologists strive to observe, record, and analyze each species' behavioral
repertoire in order to understand the roles of development, ecology, physiology, and evolution in
shaping that behavior. Insects have always been popular as subjects for behavioral research because, in
comparison to vertebrates, they have relatively simple nervous systems, they exhibit discrete responses to
external stimuli, and they are more conducive to ethical experimentation. Although they may lack the
capacity for intelligence or forethought, they still display a fascinating array of adaptive behavior that is a
source of wonder and amazement to anyone who takes the time to study it.

Any behavior we can observe by watching an animal is overt behavior. In insects, this usually includes
responses to external stimuli as well as spontaneous activities that are related to the animal's internal
(physiological) needs. Ethologists use the term "drive" ("hunger drive", "sex drive", etc.) to describe
motivational urges that compel animals to behave as they do. Insects also appear to have internal "drives"
for dispersal or migration as well as "drives" to complete stages in development such as constructing a nest
or spinning a cocoon.
In general, overt behavior may be classified as innate, learned, or complex. Many people use the term
"instinctive behavior" as a synonym for innate behavior. Although both terms refer to natural, inborn patterns
of behavior, some ethologists avoid the word "instinct" because in common English usage it often includes
the connotation of acquired aptitudes or talents, as in: "She has good business instincts." Purists avoid any
confusion in meaning by using the adjective "innate" for behaviors that are
acquired through inheritance, and "learned" for behaviors that are acquired
through experience. Complex behavior is a blend of innate and learned
components.

Innate Behavior

Innate behavior is genetically programmed. Individuals inherit a suite of

behaviors (often called an ethogram) just as they inherit physical traits such
as body color and wing venation. In general, innate behaviors will always be:
1. Heritable -- encoded in DNA and passed from generation to
generation
2. Intrinsic -- present in animals raised in isolation from others
3. Stereotypic -- performed in the same way each time by each
individual
4. Inflexible -- not modified by development or experience
5. Consummate -- fully developed or expressed at first performance
Since innate behavior is encoded in DNA, it is subject to genetic change through mutation, recombination,
and natural selection. Just like physical traits, innate behaviors are phylogenetic adaptations that have an
evolutionary history.

Comparative study of similar species often sheds light on the selective pressures that drive evolutionary
changes in behavior. It may also help explain the origin of some very unusual behavior. One species of
dance fly, for example, has a courtship ritual in which a male gives a ball of silk to a female. She unravels
the ball while he mates with her. By itself, this curious behavior seems truely bizarre. But a study of
courtship in other dance flies reveals that males use a nuptial gift as a way to divert a female's aggressive
behavior long enough for insemination to occur. In "primitive" species, the nuptial gift is an item of prey that
the female consumes during copulation. In more "advanced" species, males wrap the prey insilk, thus
buying a little extra time for copulation. In the species where males offer just a ball of silk, they are exploiting
the female's innate response to the stimulus of a nuptial gift. Just another example of "selfish genes" at
work!

Just because an insect's behavior is innate does not necessarily mean it is simple. Over time, natural
selection can lead to surprisingly intricate and sophisticated behavior such as the dance language of honey
bees or the courtship rituals of dance flies. These behaviors may appear purposeful and intelligent, but they
are merely the product of millions of years of genetic refinement through natural selection.
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In general, innate behaviors are viewed as "programmed"

responses to external stimuli. They usually fit into one of the
following categories:

Reflex. The most basic unit of innate behavior is a simple

reflex arc. This is a neural pathway that may involve as few as
two neurons: a sensory neuron detects a stimulus and is linked
with a motor neuron that sets off a response in an effector cell (such as a muscle or a gland cell). More
commonly, reflex arcs also include an association neuron spliced between the sensory and motor
neurons. The association neuron also synapses with other neurons to relay information to the brain and
other parts of the body. Most insects have simple "startle" reflexes triggered by small disturbances as well
as more comprehensive "escape" reflexes triggered by larger disturbances.

Orientation Behaviors are coordinated movements (walking, flying, swimming, etc.) that occur in response
to an external stimulus. These behaviors have adaptive value for survival by helping the insect locate (or
avoid) the source of a stimulus. Orientation behaviors can be viewed as elements in a neural hierarchy. The
simplest behaviors involve input from only a single sensory receptor whereas more advanced behaviors
require bilateral input from a pair of receptors.

Kinesis is a change in the speed of movement (orthokinesis) or a change in the

rate of turning (klinokinesis) which is directly proportional to the intensity of a
stimulus. Input from only a single sensory receptor is necessary. A kinesis is
non-directed orientation, that is, the animal exhibits a "random walk". The
change in speed or rate of turning increases the probability of locating the
stimulus but does not guarantee it.

Taxis is a movement directly toward (positive) or away from (negative) a stimulus. A klinotaxis involves side-
to-side motions of the head or body with successive comparison of stimulus intensity as the animal moves
forward. A tropotaxis requires bilateral input from paired sensory receptors such that the signal is equalized
in both receptors. Stimulus intensity increases with movement toward the source and decreases with
movement away from the source. A prefix may also be used to designate the type of stimulus involved (i.e.
phototaxis=light; geotaxis=gravity; thigmotaxis=contact with other objects
The dorsal light reaction is a special case (telotaxis) in which movement occurs at a constant 90 angle to
a light source. By keeping the sun directly overhead, a flying or swimming insect can insure that it travels
parallel to the ground (or water surface). A diving beetle, for example, can be fooled into swimming upside
down in an aquarium that is lit from below. The dorsal light reaction also explains why moths tend to circle a
street lamp at night.
The light compass reaction is another special case (menotaxis) in which insects (honey bees, for example)
fly away from the nest site at a fixed angle (x) to the sun and return at the supplementary angle (180-x) --
all without ever taking a class in geometry!

Fixed Action Pattern (FAP) is a sequence of coordinated movements that are

performed together as a "unit" without interruption. Each FAP is triggered by a unique
stimulus variously known as a sign stimulus, a key stimulus, or a releaser. A praying
mantis striking at prey is a typical example. The releaser for this FAP is any movement
by a small (prey-sized) object within striking distance. Once initiated, the mantis cannot
change direction in mid-strike or abort the mission if the prey escapes. Other common
examples of FAPs include courtship displays, hunting or food gathering, nest-building
activities, and attack or escape movements. Unlike simple reflexes, FAPs may involve a
whole-body response and often require a threshold level of internal readiness (drive).

A fixed action pattern is rarely triggered by the "big picture" (Gestalt) in an environmental
context. Instead, the sign stimulus is usually a highly specific signal that is consistently encountered at an
appropriate time. Thus a male fruit fly will perform a courtship display for a pheromone-impregnated cork
even though the cork doesn't look, taste, feel, or act like a female fruit fly. Sometimes, itis possible to find or
create a supernormal stimulus, essentially an exaggerated signal that produces a more vigorous or more
sustained response than a normal releaser. Apple growers, for example, hang large red spheres coated with
stick-um in their orchards to catch adults of the apple maggot (Rhagoletis pomonella). The red spheres are
a key stimulus for oviposition, and to female apple maggots, size matters!
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Performing one FAP may lead an insect to encounter the releaser for a second FAP and that in turn may
lead to the releaser for a third FAP, etc. This type of behavioral cascade is common in insects. Niko
Tinbergen, one of the "fathers" of modern ethology, demonstrated that hunting behavior in a predatory wasp
proceeds through a stepwise series of three FAPs. The first releaser is visual: movement of a prey-sized
object triggers down-wind pursuit of the prey. At that point, prey odor is a releaser that triggers catching
behavior. Finally, tactile cues from the prey release stinging and egg laying behavior.

When a fixed action pattern fulfills or satisfies a physiological drive, it may be known as a consummatory
act. Taking a blood meal, for example, satisfies a mosquito's hunger drive. Any behavior that increases an
individual's probability of encountering the releaser for a consummatory act is often called an appetative
behavior. Thus, chirping by a cricket may be regarded as an appetative behavior because it increases the
chances of finding a mate and satisfying the sex drive.

Learned Behavior

Learning can be defined as a persistent change in behavior that occurs as a result of experience. Since a
newborn nymph or larva has no prior experience, its first behaviors will be entirely innate. Each individual
starts life with a "clean slate": it acquires new skills and knowledge through trial and error, observation of
other individuals, or memory of past events. In general, learned behaviors will always be:
1. Nonheritable -- acquired only through observation or experience
2. Extrinsic -- absent in animals raised in isolation from others
3. Permutable -- pattern or sequence may change over time
4. Adaptable -- capable of modification to suit changing conditions
5. Progressive -- subject to improvement or refinement through practice

Although insects have relatively simple nervous systems and are not able to master college-level physics,
they have demonstrated the ability to "learn" in each of the following ways:

Habituation is learning to "ignore" stimuli that are

unimportant, irrelevant, or repetitive. For example, a puff of
air on the cerci of a cockroach will cause the animal to
scamper away. But repeating the same stimulus over and
over will lead to a decrease in the response and eventually
to no response at all. In some insect populations,
widespread use of sex pheromone will disrupt mating
behavior. By making everything in the world smell like a
virgin female, males become habituated to the odor and stop responding to the signal. If a female
cannot attract a mate, she will not produce any offspring.

Classical Conditioning is learning to associate one stimulus with another, unrelated

stimulus. Honey bees, for example, learn to associate floral colors and fragrances with the presence
of nectar. They can be "trained" to collect sugar water from colored dishes on a feeding table. If a
blue dish with pure water sits next to a yellow dish with sugar water, worker bees will quickly learn to
associate "yellow" with "food" (even if the dishes are moved around). When solutions in the two
dishes are suddenly swapped (sugar to blue and water to yellow), the bees will ignore blue and
continue to forage at yellow until they eventually "learn" (by trial and error) to look for the blue dish.

Instrumental Learning depends on the animal's ability to remember the outcome of past events and
modify future behavior accordingly. Good consequences (positive feedback) reinforce the behavior
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and increase its likelihood of occurrence in the future. Bad consequences (negative feedback) have
the opposite effect. Cockroaches learning to run through a simple maze to find food is a simple
example of instrumental learning (also known as operant conditioning).

Latent Learning involves memory of patterns or events when there is no apparent reward or
punishment associated with the behavior. A sand wasp, for example, learns the location of her nest
site by taking a short reconnaissance flight each time she leaves the nest. She remembers the
pattern of surrounding landmarks to help her find the nest when she returns. Likewise, worker ants
can remember a series of landmarks along a trail and follow them (in reverse order) back home to
the nest site. Honey bees also show latent learning when they follow the waggle dance of a forager
and then use that information to find the reported nectar source.

Imprinting is a special case of programmed learning that occurs early in life and only within a short
time-window known as the "critical period". During this brief interval, the animal acquires an indelible
memory of certain salient stimuli in its "home" environment (taste of the host plant, smell of the nest
site, etc.). This memory is retained throughout life and recalled later when needed. Fruit fly larvae,
for example, will imprint on the taste and smell of their food. If reared on a diet that contains apple
extract, adult females will show a strong preference for apples when they eventually search for a
place to lay their own eggs. Not just any stimulus will do. Imprinting is apparently regulated by an
innate "neural template" that restricts what can be remembered.

Complex Behavior

Ethologists are often careful to distinguish between learned and innate behaviors, but in reality the two are at
extreme opposite ends of a single continuum. Most overt behavior is neither 100% innate nor 100%
learned.

Sometimes innate behaviors may be modified (or modulated) through practice and experience.

In locusts, for example, the ability to fly is innate, but an older, experienced individual consumes less
energy (per unit time) than a novice flier. This suggests that the older insect has "learned" to fly more
efficiently. Similarly, learned behaviors may incorporate or depend upon elements of innate
behavior. Indeed, the ability to learn, to associate, or to remember is almost certainly an innate feature of the
insect's nervous system. Schematically, it may be useful to think of a box that represents the boundaries of
an animal's ethogram. All behavior must occur inside the physiological limits of this box (e.g. a beetle larva
does not have wings, therefore it cannot fly). Within the box, a set of innate behaviors can be simplistically
represented by straight lines. By following a zigzag route, an insect can use only innate behavior to get from
point "A" to point "B". But a learned behavior, superimposed on this innate grid, might provide a
"shortcut" that is more useful or more efficient. As in the locust example above, the innate ability to fly may
be refined and improved through experience.
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Periodic Behavior

Insects, like most other living

organisms, exhibit cycles of activity that
are synchronized with periodic events in
their environment. Hatching, molting,
foraging for food, courtship, and nest
building are all examples of common
behaviors that may recur at regular
intervals. When these periods of activity
are repeated at about the same time
each day, an animal is said to exhibit
diel periodicity (a daily activity cycle).
An actograph is a device that entomologists use to record
movement (or other activity) of a caged insect. Data from an
actograph is usually displayed by a chart recorder - much like
a seismograph registers earthquakes. Mouse over each term
highlighted at the left to see a corresponding pattern of daily
actograph data.

Diurnal insects are active during

daylight hours
Nocturnal insects are active at
night
Crepuscular insects are active
at dawn and dusk

Two additional sub-categories exist to

differentiate dawn and dusk periods:

Matinal behavior occurs only

near dawn
Vespertine behavior occurs only near dusk

In addition to daily rhythms, many insects also exhibit periodic behaviors with much longer cycles:

Circalunar rhythms are synchronized with the phases of the moon and have a cycle of about 28
days. Many insect species tend to molt or emerge as adults during the dark phase of the moon as
an adaptation to evade predators.
Circannual rhythms are synchronized with seasons of the year and have a cycle of about 12
months. Examples include molting in periodical cicadas and migratory flights in monarch butterflies.

Entrainment

Environmental cues such as temperature, light intensity, or location of the sun provide the information
animals need to synchronize their behavior with the earth's daily, monthly, or yearly cycles. The process of
establishing and maintaining this synchrony is commonly known as entrainment. In insects, entrainment of
periodic behavior usually occurs in one of two ways:

1. Exogenous entrainment involves direct response to environmental cues that "trigger" the onset and
termination of a particular behavior. In some species of crickets, for example, males begin chirping
at nightfall in response to low light intensity and they continue until the light returns at dawn. These
crickets will also chirp during a solar eclipse or any time they are covered by a box that keeps out the
light. They will not chirp at all if they are exposed to constant illumination. As the name implies,
exogenous behavior is regulated by forces "outside" the organism. An experimenter can control
when crickets chirp simply by changing the light intensity in their environment.
2. Endogenous entrainment involves an internal biological clock that "measures" the passage of time
and sends "start" and "stop" signals to the nervous system. These periodic behaviors are often
called circadian rhythms. In a cricket species where chirping behavior is controlled endogenously,
the males will always start to chirp around dusk, regardless of light intensity. They will not be fooled
into chirping during a solar eclipse, or when covered by a box, and they will start chirping around
dusk even if the lights are always on. Start and stop signals are generated from "inside" the
organism and cannot be arbitrarily changed by an experimentor.
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Even with this internal chronometer, an animal's circadian rhythms must still be synchronized with the
environment. As day lengths change throughout the year, insects have to adjust their activity cycles to
compensate. In fact, a slight adjustment is made each day in response to a cue (usually light or
temperature) from the environment. This signal, which serves as a daily "reset" mechanism for the biological
clock, is called a zeitgeber - derived from the German word for "time" (zeit) and the verb "to give" (geben).

An actograph of an entrained insect shows a constant pattern of diurnal activity under a regimen of 8
hours light and 16 hours dark. When held in constant light (or constant dark), the same insect shows
a phase shift in behavior as its biological clock becomes "free-running."

In the complete absence of any environmental cues, a biological clock becomes "free-running". Its
endogenous period is usually between 24.5 and 25.0 hours, so the animal's behavior tends to occur a little
later each day as the internal clock gradually drifts out of phase with the environment. Humans experience
this same phenomenon in sleep/wake cycles when housed under conditions of constant light or constant
dark. We also experience "jet lag" when we travel to a distant time zone. In both humans and insects, it
takes several days for the body to adjust to a new photoperiod regime.

Recent discoveries in insect genetics and physiology have revealed that there are different biological clocks
for different rhythms. The clock that controls when a fly lays an egg may not be the same as the clock that
controls when the fly forages for food. Biological clocks are emergent properties of individual cells and may
be widespread throughout the body. Although they regulate different functions, they all appear to have a
common genetic origin. In fruit flies (Drosophila spp.), the clock mechanism consists of four regulatory
proteins that interact through a negative feedback system. The clock cycle begins when two of these
proteins (named JRK and CYC) bind together in the nucleus where they "activate" genes that produce two
other proteins (PER and TIM). As the levels of PER and TIM accumulate over timethey inactivate the genes
that produce JRK and CYC. This, in turn, leads to reduced production of PER and TIM and, eventually,
renewed production of JRK and CYC to complete the cycle. Normally, this system completes one cycle
every 24 hours. Mutations in the genes that encode these four clock proteins can change the duration of the
cycle or completely disable the clock mechanism.

Periodic behavior is certainly not unique to insects -- it occurs, to some extent, in nearly every living
organism on earth. To a biologist, this fact suggests that there are strong selective pressures acting to
preserve the cells' genetic mechanisms for periodicity. For insects, daily periods of inactivity serve as an
opportunity to conserve energy and resources. Many mosquito species, for instance, seek a humid, shady
habitat during the day to avoid dehydration. Likewise, honeybees (Apis mellifera) return to the hive at dusk
and pass the night in a quiescent state that has been described as "sleep". For other species (e.g.
cockroaches and stoneflies) circadian rhythms in emergence and/or molting may reduce the threat of
predation. And finally, synchronized periods of activity within a species can expedite the timing of courtship
behavior and reduce the metabolic costs of reproduction. It's all a matter of timing!
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Insect Communication

Communication may be defined as any exchange of information between

individuals. For members of the Human species, it is an essential part of all social
interactions. We communicate through speech, written language, sign language, body
language, Braille, Morse code, and many other cultural and technological
inventions. There is even some evidence that we use body odor in
communication. Our brains are uniquely adapted for symbolic communication, but
most of our "language" skills are acquired through learning.

Insects also have many ways to communicate but, unlike humans, their "language" is almost entirely
innate. Each individual is born with a distinctive "vocabulary" that is shared only with other members of its
own species. Learning plays little or no role in the ability to produce these signals or to understand them.

An act of communication is not always overt or obvious. No physical entity passes from one individual to
another, so it is not always possible to know when exchange of information occurs. The situation is
analogous to an alien from another planet who comes to Earth and observes human behavior. Without
knowing our language and customs, the alien would be unlikely to recognize a black arm band, a coy wink,
or a "yellow ribbon 'round the old oak tree" as forms of human communication. We are in much the same
predicament when we study insect communication. The only way to distinguish communicative behavior
from non-communicative behavior is by looking for evidence of a change in the behavior (or sometimes,
physiology) of another individual.

For experimental purposes, ethologists (scientists who study animal behavior) often define communication
as:

An action or condition on the part of one organism that alters the behavior of another organism in an
adaptive way.

Thus, an insect may send a communication signal by doing something (e.g. make a noise, release a
chemical, or flash a light) or the signal may simply be an inherent part of the insect's physical makeup (e.g.
wing pattern, body color, or surface chemistry). In either case, the signal must elicit some behavioral change
in order for a human observer to recognize its existence.

Why do insects communicate?

Some form of intraspecific communication is a prerequisite for any behavior that involves the participation or
cooperation of two or more individuals. Intentionally or not, insects may also communicate with members of
other species (interspecific communication). The adaptive value of these communication signals may
include:

1. Recognition of kin or nestmates

2. Locating or identifying a member of the opposite sex
3. Facilitation of courtship and mating
4. Giving directions for location of food or other resources
5. Regulating spatial distribution of individuals -- aggregation or dispersal;
establishing and maintaining a territory
6. Warning of danger; setting off an alarm
7. Advertising one's presence or location
8. Expressing threat or submission (agonistic behaviors)
9. Deception / mimicry

How do insects communicate?

Like all other animals, insects use their five senses to acquire information about their environment; any of
these sensory modalities may serve as a pathway for the exchange of information. Taste and touch are both
contact senses, therefore, exchange of information can occur only when two individuals are touching one
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another. Vision, olfaction (smell), and hearing are remote senses -- information signals may propogate
through the air (or water) over considerable distances.

Each signal modality has unique advantages and disadvantages. Follow the links below to find examples of
each signal system and learn more about insect communication.

Tactile Communication

"Keep in touch!" For you, it's probably just a metaphor, but for some insects it's really a channel of
communication. Since many insects have poor vision and sound perception, physical contact provides an
important avenue of communication. In blister beetles (family Meloidae),
courtship begins with a series of antennal taps by the male on each side of
the female's body. She signals her receptivity by lifting her wing covers
(elytra) and allowing him to climb on her back. But to complete his quest,
the male must continue tapping, alternating from side to side at just the
right frequency until the female is stimulated to extend her genitalia and
begin mating.

Antennal tapping is also an essential component of communication in both ants and

termites. It's not clear exactly what information may be exchanged, but it certainly
involves nestmate recognition and leads to exchange of food through trophallaxis. In
some cases, instantaneous feedback allows the sender to monitor the receiver's
response and alter the signal if necessary. Antennal tapping on the hind legs is used
during tandem running in both ants and termites. This is a "follow-the-leader"
behavior in which the tapping informs the leader that she has not lost her disciple. If
tapping stops, the leader instinctively turns around and searches in ever-widening
circles until she re-establishes contact with the follower.

The "dance" language of honeybees is largely a tactile communication system, performed in total darkness
on the vertical surface of the honeycomb. A "round dance" signals to nestmates the presence of a nectar
source in close proximity to the hive (usually less than 80 feet). It consists of a series of circular runs with
more or less frequent changes in direction. The greater the frequency of direction changes, the better the
quality of the nectar source. The "waggle dance" is used for longer distances. It involves a figure eight
pattern with a series of abdominal waggles on a straight run after each half-circle turn (see figure at
left). Distance is indicated by the duration of the straight run and the frequency of the waggles. Direction is
indicated by the angle of the straight run (relative to vertical) and corresponds to the horizontal angle
between the sun and the direction of the food source.

Tactile cues generated by ripples on the water surface allow whirligig beetles (family Gyrinidae) to constantly
monitor the location of dozens of other nearby whirligigs. Thanks to this tactile communication system, the
whirligigs can swim rapidly in circles, avoid bumping into other members of their own species, and still detect
the presence of nearby predators or prey.

Certain treehoppers (order Hemiptera: family Membracidae) produce vibrations in the tissue of their host
plant that can be felt by all other treehoppers on the same plant. The signal travels throughout the plant in
much the same way that banging on water pipes in your apartment creates noise throughout the whole
building. The signals apparently work as an alarm system, and in some species, they may be used by
nymphs to elicit protective maternal behavior. Substrate vibrations can be a particularly effective
communication system for small insects who cannot generate an acoustic signal loud enough to be heard
more than few inches away.
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Acoustic Communication Pros and Cons of Tactile Communication

Advantages:

Instantaneous feedback
Sound is localized area
Individual recipient
Effective in the dark (e. g. caves, wood galleries)

Disadvantages:

produced when a physical object Not effective over distance

vibrates rapidly, disturbs nearby air Organisms must stay in direct contact
molecules (or other surrounding Message must be repeated to each recipient
medium), and generates Vibration signals can be intercepted by predators
compression waves that travel in all
directions away from the
source. We perceive these waves as sound when they collide with our ear drum and cause a mechanical
disturbance that is detected by sensory neurons in our inner ear. Both living and non-living objects produce
sound, but only animals use it as a means of communication.

Sound serves as a very effective communication modality. It can be made to vary in frequency (high pitch
vs. low pitch), amplitude (loudness), and periodicity (the temporal pattern of freqency and
amplitude). Together, these three variables can create an extremely wide and complex range of signals --
from an insect's mating call to human speech and vocal music. Since sound waves move rapidly through air
(about 331 m/sec), acoustic signals can be quickly started, stopped, or modified to send a time-sensitive
message.

At best, the human ear is able to Pros and Cons of Acoustic Communication
detect sound frequencies only Advantages:
within the range of about 20-20,000
hertz (vibrations per second). But Not limited by environmental barriers
some insects (as well as other
animals like bats and dolphins) Effective over distances and around corners
produce and detect sounds that are
well above this frequency Highly variable, fast change -- high information
range. Some grasshoppers and content
moths, for example, produce
ultrasonic sounds as high as 80,000 Disadvantages:
hertz. Entomologists study these
high-pitch sounds by using an audio May reveal location of sender to a potential predator
transducer, an electronic device
that converts inaudible high Less effective in "noisy" environments (e.g. seashore)
frequencies to lower audible
frequencies. May be metabolically "expensive" to produce

Most insects detect sound with a Attenuation -- intensity falls rapidly with distance from
tympanic mambrane in the source (cube-root function)
abdomen (e.g. grasshoppers and
moths) or in the tibiae of the front
legs (e.g. crickets and katydids). Mosquitoes have antennal hairs that resonate to certain frequencies of
sound. But sound vibrations can also travel through solid objects, and some insects (e.g. some species of
ants, bees, termites, and treehoppers) can sense substrate vibrations with mechanoreceptors (chordotonal
organs) in their legs. Since these signals are "felt" rather than "heard", they are usually regarded as a form
of tactile communication.

See and hear each insect in the table below by clicking on its common name.
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Chemical Communication

It is probably safe to say that insects rely more heavily on chemical signals than on any other form of
communication. These signals, often called semiochemicals or infochemicals, serve as a form of
"language" that helps to mediate interactions between organisms. Insects may be highly sensitive to low
concentrations of these chemicals -- in some cases, a few molecules may be enough to elicit a response.

Semiochemicals can be divided into Pros and Cons of Chemical Communication

two groups based on who "sends" a
message and who "receives" it: Advantages:

1. Pheromones are chemical Not limited by environmental barriers

signals that carry information Effective over distances and around corners
from one individual to Effective either day or night
another member of the same
Longer lasting than visual or auditory signals
species. These include sex
Metabolically "inexpensive" because only small
attractants, trail marking
quantities are needed
compounds, alarm
substances, and many other
intraspecific messages. Disadvantages:
2. Allelochemicals are signals
that travel from one animal to Low information content (presence/absence)
some member of a different Not effective in an upwind direction
species. These include
defensive signals such as
repellents, compounds used to locate suitable host plants, and a vast array of other substances that
regulate interspecific behaviors.

Allelochemicals can be further subdivided into three groups based on who "benefits" from the meassage:

1. Allomones benefit the sender -- such as a repellent, or defensive compound (e. g. cyanide) that
deters predation.
2. Kairomones benefit the receiver -- such as an odor that a parasite uses to find its host.
3. Synomones benefit both sender and receiver -- such as plant volatiles that attract insect pollinators.

Grape root borer moths (Vitacea polistiformis).

Male attracted by female's sex pheromone.

(click the "Play" button for video)

Insects use their sense of taste or smell to detect the presence of semiochemicals. Specialized receptors
may be located anywhere on the body, but are especially common on the feet, antennae, palps, and
ovipositor (see Chemoreception). The sense of smell (olfaction) is used for remote chemoreception --
detecting semiochemicals with low molecular weight that are volatile enough to become airborne. The sense
of taste (gustation) is used for contact chemoreception -- detecting molecules that adhere to a substrate or to
the outside of an insect's body.

Which type of signal is involved (pheromone, allomone, kairomone, or synomone)?

 11

1. A parasitic fly, Euclytia flavahe, is attracted to the odor of its host.

2. Ants are able to recognize their nestmates by the taste and smell of hydrocarbons (lipids) found on
the surface of the cuticle.
3. Caterpillars that feed on plants sometimes unwittingly induce those plants to release a defensive
compound that attracts parasitoids of the caterpillars.
4. Caterpillars of Lycaena arion produce a chemical that attracts ants and elicits care-giving behavior.
When carried into the ant's nest and placed among the brood, these caterpillars will eat the ant
larvae.

Visual Communication

Many insects communicate with visual signals. The color patterns and other markings on the
wings of butterflies and moths facilitate species recognition in much the same way colored
uniforms reveal the players' affiliations on a football field. Some insects use bright colors,
eyespots, or other distinctive patterns to scare away predators, to advertise their ability to sting, or
to mimic the appearance of another unpalatable species. Other insects use dance-like body
movements to attract a mate or to communicate with nestmates. Most of these signals are effective only as
long as they are visible in daylight. But a few insects (fireflies, for example) can generate their
own light and use visual signals that can be seen at night.

Passive signals, Pros and Cons of Visual Communication

such as
eyespots and Advantages:
color patterns
can serve as a
Effective over long distances
form of "free
advertising". The colorful wings Can be used while moving
of a butterfly, for example, are a Fast -- speed of light
"billboard" publicizing its Effective in all directions (independent of wind)
species identity. Individual Passive signals require no expenditure of energy
insects incur little or no
metabolic cost for displaying Disadvantages:
these messages because they
are an integral part of the Requires a clear line of sight
integument. It may be prudent
Visual signals may be intercepted by predators
to hide these signals from a
Only effective in daylight (in fireflies, only at night)
potential predator, so some
insects have a way to conceal Active signals may be metabolically "expensive" to produce
their message when
necessary. The red admiral butterfly, for example, has bright, distinctive markings on the upper wing surface
and drab, protective coloration on the underside.

Active signals, like body movements and light flashes, are more costly to
produce, but they can be withheld from use at inappropriate times. They
may also have a higher information content because signal frequency,
duration, or periodicity may convey additional meaning. In fireflies, for
example, pulses of light are used in a courtship dialogue between a male
(usually flying) and a female (usually perched in the vegetation). Each
species has a unique flash pattern and response time. Males of Photinus
pyralis emit a single "J"-shape flash during a rising flight movement. A
female responds with a single flash after a two second interval. In Photinus consumilis, males emit a series
of 3-5 short flashes and the females respond with a double flash. Roles are reversed in some tropical
species where the females fly and the males signal from perches in the vegetation.
 12

Unlike humans, many insects have the ability to see ultraviolet light. Not surprisingly, some species
communicate using wavelengths in this part of the spectrum. Female cabbage butterflies, for example, have
ultraviolet-reflecting scales on the dorsal wing surface. When they fly, each downstroke of the wing creates
a brief "flash" of UV that males apparently recognize as the flight signature of a potential mate. A "flashing
female" may attract several males who engage in aerial courtship displays.

Ultraviolet reflection from a male

alfalfa butterfly in simulated flight

In alfalfa butterflies, only males have the UV-reflective scales. They flutter in
front of the females to create a flickering courtship display. Missing scales
reduce the wings' reflectivity -- a sign of aging that impairs a male's ability to
seduce a mate.

Social Insects

Social insects are among the most dominant and prolific of all organisms on earth. In South America,
leafcutter ants (Atta spp.) consume more foliage than mammalian herbivores; in the southwestern
United States, more seeds are collected and eaten by harvester ants than by all other forms of life; in
some tropical habitats, the biomass of ants and termites exceeds that of all other animal species
combined; in the African savanah, a single colony of driver ants may contain as many as 20 million
workers; in Japan, a supercolony of Formica yessensis with 45,000 interconnecting nests contained
more than a million queens and 306 million workers within an area of 2.7 square kilometers.

Many insects exhibit "social" behaviors (e.g. feeding aggregations, parental care of the young, and
communal nest sites). In a broad sense, any insect that interacts with another member of its own species
could be called a social insect. But as a rule, entomologists do not regard these behaviors as sufficient
justification for classifying a species as truly social (i.e. eusocial). In order to qualify as eusocial, a species
must exhibit all four of the following characteristics:

1. Share a common nest site

2. Individuals of the same species cooperate in caring for the young
3. Reproductive division of labor -- sterile (or less fecund) individuals work for the benefit of a few
reproductive individuals
4. Overlap of generations -- offspring contribute to colony labor while their parents are still alive

Species that lack one or more of these characteristics are classified as presocial. Within this category are
subsocial species (in which the parents care for their offspring) and parasocial species (which have a
common nest site but lack one or more of the other eusocial characteristics). The table below helps clarify
the various terms used for social and presocial insects. Click on each term for a formal definition.

A Classification System for Presocial and Eusocial Insects

Common Cooperative Reproductive Generational

nest site brood care castes overlap

Solitary No No No No

Communal or Yes No No No
Subsocial

Quasisocial Yes Yes No No

 13

Semisocial Yes Yes Yes No

Eusocial Yes Yes Yes Yes

Relatively few insects are classified as eusocial -- the distinction is limited to the following groups:

Order Isoptera Order Hymenoptera

Termites -- all species Ants -- all species (family Formicidae)

Bees -- about 600 species in the family Apidae

Wasps -- about 700 species in the family Vespidae

Termites

All members of the order Isoptera are eusocial insects. Termites feed primarily on the cellulose and lignin
found in plant cell walls; these compounds are the main ingredients of wood and all paper
products. Termites cannot digest the cellulose directly so they rely upon symbiotic bacteria and protozoa
living within their intestines to supply most of the enzymes needed for cellulose digestion. Termites are
sometimes called white ants. They may resemble ants in size, but ants have a narrow waist and elbowed
antennae while termites have a thick waist and antennae that resemble a string of beads.

Ecologically, termites play an important role in the environment by helping to break down and recycle dead
wood and other plant tissues. They become pests when their appetite for wood and wood products extends
to human homes, fence posts, building materials, cardboard, and other valuable products. In tropical and
subtropical forests where termites are abundant, railroads must use expensive metal ties because wooden
ones are quickly destroyed.

Ecologically, termites play an important role in the environment by helping to break down and recycle dead
wood and other plant tissues. They become pests when their appetite for wood and wood products extends
to human homes, fence posts, building materials, cardboard, and other valuable products. In tropical and
subtropical forests where termites are abundant, railroads must use expensive metal ties because wooden
ones are quickly destroyed.

The Termite Colony

 14

Each termite lives in a nest or colony with

hundreds, thousands, or even millions of its << Termite Social Register >>
brothers and sisters. In fact, the termite colony
is really a large, extended family. Within this
Queen - Fertile adult female.
family, various groups of individuals have Reproductive
different functional roles according to a "caste Caste
system". King - Fertile adult male.

The worker caste Non fertile males and females with

Soldier
is the largest special morphological adaptations
Caste
group. It consists for defense of the colony.
entirely of
immatures, both
males and Immature males and females
females. These (nymphs) - offspring of the king and
Worker
soft-bodied, wingless individuals perform all of queen, with potential to molt into
Caste
the hard labor in the colony: they clean, replacement soldiers or
maintain, and repair the nest, gather food and reproductives if needed.
water, care for the young, and construct new
tunnels and galleries as the colony grows. These juveniles all have the genetic capacity to undergo
additional molts and become soldiers or reproductives, but most will spend their entire lives as workers.

Members of the soldier caste are larger in size but fewer in number than the workers.

They are also wingless, but they have large heads with powerful jaws. Their job is
to guard the nest site and protect it from attacks by
ants or other invaders. In some species the soldiers
lack jaws but have a large gland in the head that
shoots defensive chemicals through a
nozzle at the front of the head. The soldiers are
unable to care for themselves so they must
be fed and groomed by the workers.

The reproductive caste always includes a king (male) and a queen (female) who are the parents of the
termite family and founders of the colony. Some species also have a few supplemental reproductives who
share the egg laying duties. These are the only adult insects in the colony. The queen lays large numbers
of eggs which develop into more workers and soldiers as the family grows.

In every mature colony, there also develops an annual population of young winged
reproductives that swarm from the parent nest for a short mating flight. After flight, the
delicate wings break off, and the new king and queen set out to find another nest site
and start a new colony. Large colonies with multiple reproductives may also split into
two or more daughter colonies, a process known as "budding".

The termite's caste system is regulated by

pheromones. The king and queen each produce special pheromones that
circulate throughout the colony and inhibit workers of the same sex from
molting into reproductive adults. A death in the royal family (or an increase in
the size of the colony) results in a lower concentration of the corresponding
pheromone and, subsequently, one or more workers will molt into
replacement reproductives. Likewise, the concentration of sex-specific soldier
pheromones regulate the numbers of male and female soldiers to fall within
an optimal range based on colony size. Excess numbers of soldiers or
reproductives may be killed and eaten by the workers.
 15

Kinds of Termites.

About 2750 different species of termites are known. These can be divided into two groups: those
that live entirely within wood, and more advanced species that tunnel and nest in the soil. In
terms of their ecology and behavior, the most primitive species are similar to certain wood-
dwelling cockroaches with whom they may share a common ancestor. These primitive species
often have specialized habitat requirements, nesting only in rotten wood, damp wood, or dry wood. Their
colonies are rather small and persist only as long as the food resource lasts. All wood-dwelling termites
produce distinctive waste pellets which are often the first sign of an active infestation.

Subterranean termites construct underground nests and have the ability to tunnel through the soil to find new
food resources. These colonies are often long-lived and may grow to include several million
individuals. The subterranean termites that live in North America and Europe often invade wooden
structures above the ground by building earthen tubes that serve as protective tunnels between the nest and
their food source. These tubes are good evidence of a termite infestation.

In Africa and Australia, other subterranean species mix bits of soil with saliva to build nest mounds that are
up to 20 feet (6 meters) tall. The inside of the mound is divided into numerous chambers and galleries. The
king and queen live in a special cell deep inside the mound. The females abdomen grows in size until it is
large enough to hold many thousands of eggs. The queen
lays these eggs at the rate of several thousand a
day. Worker termites carry the eggs away to specially
constructed cells in the nest. There the workers care for the
young as they hatch from the eggs.

Some of the mound-building termites cultivate underground

fungus gardens. They collect dead plant material, mix it with
saliva and their own waste products to create a paste, and
inoculate this substance with the spores of a symbiotic
fungus. The termites feed on special structures produced by
the growing fungus.

Elements of Behavior

Behavior is what animals do. It can be defined more precisely as an internally directed system of adaptive
activities that facilitate survival and reproduction. Ethology is the scientific study of animal behavior --
particularly when that behavior occurs in the context of an animal's natural environment. Ethologists strive to
observe, record, and analyze each species' behavioral repertoire in order to understand the roles of
development, ecology, physiology, and evolution in shaping that behavior. Insects have always been
popular as subjects for behavioral research because, in comparison to vertebrates, they have relatively
simple nervous systems, they exhibit discrete responses to external stimuli, and they are more conducive to
ethical experimentation. Although they may lack the capacity for intelligence or forethought, they still display
a fascinating array of adaptive behavior that is a source of wonder and amazement to anyone who takes the
time to study it.

Any behavior we can observe by watching an animal is overt behavior. In insects, this usually includes
responses to external stimuli as well as spontaneous activities that are related to the animal's internal
(physiological) needs. Ethologists use the term "drive" ("hunger drive", "sex drive", etc.) to describe
motivational urges that compel animals to behave as they do. Insects also appear to have internal "drives"
for dispersal or migration as well as "drives" to complete stages in development such as constructing a nest
or spinning a cocoon.
 16

In general, overt behavior may be classified as innate, learned,

or complex. Many people use the term "instinctive behavior" as
a synonym for innate behavior. Although both terms refer to
natural, inborn patterns of behavior, some ethologists avoid the
word "instinct" because in common English usage it often
includes the connotation of acquired aptitudes or talents, as
in: "She has good business instincts." Purists avoid any
confusion in meaning by using the adjective "innate" for behaviors that are acquired through inheritance, and
"learned" for behaviors that are acquired through experience. Complex behavior is a blend of innate and
learned components.

Innate Behavior

Innate behavior is genetically programmed. Individuals inherit a suite of behaviors (often called an
ethogram) just as they inherit physical traits such as body color and wing venation. In general, innate
behaviors will always be:
6. Heritable -- encoded in DNA and passed from generation to generation
7. Intrinsic -- present in animals raised in isolation from others
8. Stereotypic -- performed in the same way each time by each individual
9. Inflexible -- not modified by development or experience
10. Consummate -- fully developed or expressed at first performance
Since innate behavior is encoded in DNA, it is subject to genetic change through mutation, recombination,
and natural selection. Just like physical traits, innate behaviors are phylogenetic adaptations that have an
evolutionary history.

Comparative study of similar species often sheds light on the selective

pressures that drive evolutionary changes in behavior. It may also help
explain the origin of some very unusual behavior. One species of dance fly,
for example, has a courtship ritual in which a male gives a ball of silk to a
female. She unravels the ball while he mates with her. By itself, this curious
behavior seems truely bizarre. But a study of courtship in other dance flies
reveals that males use a nuptial gift as a way to divert a female's aggressive
behavior long enough for insemination to occur. In "primitive" species, the
nuptial gift is an item of prey that the female consumes during copulation. In
more "advanced" species, males wrap the prey insilk, thus buying a little
extra time for copulation. In the species where males offer just a ball of silk,
they are exploiting the female's innate response to the stimulus of a nuptial
gift. Just another example of "selfish genes" at work!

Just because an insect's behavior is innate does not necessarily mean it is simple. Over time, natural
selection can lead to surprisingly intricate and sophisticated behavior such as the dance language of honey
bees or the courtship rituals of dance flies. These behaviors may appear purposeful and intelligent, but they
are merely the product of millions of years of genetic refinement through natural selection.
In general, innate behaviors are viewed as "programmed" responses to external stimuli. They usually fit into
one of the following categories:

Reflex. The most basic unit of innate behavior is a simple reflex arc. This is a neural pathway that may
involve as few as two neurons: a sensory neuron detects a stimulus and is linked with a motor neuron that
sets off a response in an effector cell (such as a muscle or a gland cell). More commonly, reflex arcs also
include an association neuron spliced between the sensory and motor neurons. The association neuron also
synapses with other neurons to relay information to the brain and other parts of the body. Most insects have
simple "startle" reflexes triggered by small disturbances as well as more comprehensive "escape" reflexes
triggered by larger disturbances.

Orientation Behaviors are coordinated movements (walking, flying, swimming, etc.) that occur in response
to an external stimulus. These behaviors have adaptive value for survival by helping the insect locate (or
avoid) the source of a stimulus. Orientation behaviors can be viewed as elements in a neural hierarchy. The
simplest behaviors involve input from only a single sensory receptor whereas more advanced behaviors
require bilateral input from a pair of receptors.
 17

Kinesis is a change in the speed of movement (orthokinesis) or a change in the

rate of turning (klinokinesis) which is directly proportional to the intensity of a
stimulus. Input from only a single sensory receptor is necessary. A kinesis is
non-directed orientation, that is, the animal exhibits a "random walk". The
change in speed or rate of turning increases the probability of locating the
stimulus but does not guarantee it.

Taxis is a movement directly toward (positive) or away from (negative) a stimulus. A klinotaxis involves side-
to-side motions of the head or body with successive comparison of stimulus intensity as the animal moves
forward. A tropotaxis requires bilateral input from paired sensory receptors such that the signal is equalized
in both receptors. Stimulus intensity increases with movement toward the source and decreases with
movement away from the source. A prefix may also be used to designate the type of stimulus involved (i.e.
phototaxis=light; geotaxis=gravity; thigmotaxis=contact with other objects
The dorsal light reaction is a special case (telotaxis) in which movement occurs at a constant 90 angle to
a light source. By keeping the sun directly overhead, a flying or swimming insect can insure that it travels
parallel to the ground (or water surface). A diving beetle, for example, can be fooled into swimming upside
down in an aquarium that is lit from below. The dorsal light reaction also explains why moths tend to circle a
street lamp at night.
The light compass reaction is another special case (menotaxis) in which insects (honey bees, for example)
fly away from the nest site at a fixed angle (x) to the sun and return at the supplementary angle (180-x) --
all without ever taking a class in geometry!

Fixed Action Pattern (FAP) is a sequence of coordinated movements that are

performed together as a "unit" without interruption. Each FAP is triggered by a unique
stimulus variously known as a sign stimulus, a key stimulus, or a releaser. A praying
mantis striking at prey is a typical example. The releaser for this FAP is any movement
by a small (prey-sized) object within striking distance. Once initiated, the mantis cannot
change direction in mid-strike or abort the mission if the prey escapes. Other common
examples of FAPs include courtship displays, hunting or food gathering, nest-building
activities, and attack or escape movements. Unlike simple reflexes, FAPs may involve a
whole-body response and often require a threshold level of internal readiness (drive).

A fixed action pattern is rarely triggered by the "big picture" (Gestalt) in an environmental
context. Instead, the sign stimulus is usually a highly specific signal that is consistently encountered at an
appropriate time. Thus a male fruit fly will perform a courtship display for a pheromone-impregnated cork
even though the cork doesn't look, taste, feel, or act like a female fruit fly. Sometimes, itis possible to find or
create a supernormal stimulus, essentially an exaggerated signal that produces a more vigorous or more
sustained response than a normal releaser. Apple growers, for example, hang large red spheres coated with
stick-um in their orchards to catch adults of the apple maggot (Rhagoletis pomonella). The red spheres are
a key stimulus for oviposition, and to female apple maggots, size matters!
Performing one FAP may lead an insect to encounter the releaser for a second FAP and that in turn may
lead to the releaser for a third FAP, etc. This type of behavioral cascade is common in insects. Niko
Tinbergen, one of the "fathers" of modern ethology, demonstrated that hunting behavior in a predatory wasp
proceeds through a stepwise series of three FAPs. The first releaser is visual: movement of a prey-sized
object triggers down-wind pursuit of the prey. At that point, prey odor is a releaser that triggers catching
behavior. Finally, tactile cues from the prey release stinging and egg laying behavior.

When a fixed action pattern fulfills or satisfies a physiological drive, it may be known as a consummatory
act. Taking a blood meal, for example, satisfies a mosquito's hunger drive. Any behavior that increases an
individual's probability of encountering the releaser for a consummatory act is often called an appetative
 18

behavior. Thus, chirping by a cricket may be regarded as an appetative behavior because it increases the
chances of finding a mate and satisfying the sex drive.

Learned Behavior

Learning can be defined as a persistent change in behavior that occurs as a result of experience. Since a
newborn nymph or larva has no prior experience, its first behaviors will be entirely innate. Each individual
starts life with a "clean slate": it acquires new skills and knowledge through trial and error, observation of
other individuals, or memory of past events. In general, learned behaviors will always be:
6. Nonheritable -- acquired only through observation or experience
7. Extrinsic -- absent in animals raised in isolation from others
8. Permutable -- pattern or sequence may change over time
9. Adaptable -- capable of modification to suit changing conditions
10. Progressive -- subject to improvement or refinement through practice

Although insects have relatively simple nervous systems and are not able to master college-level physics,
they have demonstrated the ability to "learn" in each of the following ways:

Habituation is learning to "ignore" stimuli that are unimportant, irrelevant, or repetitive. For
example, a puff of air on the cerci of a cockroach will cause the animal to scamper away. But
repeating the same stimulus over and over will lead to a decrease in the response and eventually to
no response at all. In some insect populations, widespread use of sex pheromone will disrupt mating
behavior. By making everything in the world smell like a virgin female, males become habituated to
the odor and stop responding to the signal. If a female cannot attract a mate, she will not produce
any offspring.

Classical Conditioning is learning to associate one stimulus with another, unrelated

stimulus. Honey bees, for example, learn to associate floral colors and fragrances with the presence
of nectar. They can be "trained" to collect sugar water from colored dishes on a feeding table. If a
blue dish with pure water sits next to a yellow dish with sugar water, worker bees will quickly learn to
associate "yellow" with "food" (even if the dishes are moved around). When solutions in the two
dishes are suddenly swapped (sugar to blue and water to yellow), the bees will ignore blue and
continue to forage at yellow until they eventually "learn" (by trial and error) to look for the blue dish.

Instrumental Learning depends on the animal's ability to

remember the outcome of past events and modify future
behavior accordingly. Good consequences (positive
feedback) reinforce the behavior

and increase its likelihood of occurrence in the future. Bad

consequences (negative feedback) have the opposite
effect. Cockroaches learning to run through a simple maze to find food is a simple example of
instrumental learning (also known as operant conditioning).

Latent Learning involves memory of patterns or events when there is no apparent reward or
punishment associated with the behavior. A sand wasp, for example, learns the location of her nest
site by taking a short reconnaissance flight each time she leaves the nest. She remembers the
pattern of surrounding landmarks to help her find the nest when she returns. Likewise, worker ants
can remember a series of landmarks along a trail and follow them (in reverse order) back home to
the nest site. Honey bees also show latent learning when they follow the waggle dance of a forager
and then use that information to find the reported nectar source.

Imprinting is a special case of programmed learning that occurs early in life and only within a short
time-window known as the "critical period". During this brief interval, the animal acquires an indelible
memory of certain salient stimuli in its "home" environment (taste of the host plant, smell of the nest
site, etc.). This memory is retained throughout life and recalled later when needed. Fruit fly larvae,
for example, will imprint on the taste and smell of their food. If reared on a diet that contains apple
 19

extract, adult females will show a strong preference for apples when they eventually search for a
place to lay their own eggs. Not just any stimulus will do. Imprinting is apparently regulated by an
innate "neural template" that restricts what can be remembered.

Complex Behavior

Ethologists are often careful to distinguish between learned and innate behaviors, but in reality the two are at
extreme opposite ends of a single continuum. Most overt behavior is neither 100% innate nor 100%
learned.

Sometimes innate behaviors may be modified (or modulated) through practice and experience.

In locusts, for example, the ability to fly is innate, but an older, experienced individual consumes less
energy (per unit time) than a novice flier. This suggests that the older insect has "learned" to fly more
efficiently. Similarly, learned behaviors may incorporate or depend upon elements of innate
behavior. Indeed, the ability to learn, to associate, or to remember is almost certainly an innate feature of the
insect's nervous system. Schematically, it may be useful to think of a box that represents the boundaries of
an animal's ethogram. All behavior must occur inside the physiological limits of this box (e.g. a beetle larva
does not have wings, therefore it cannot fly). Within the box, a set of innate behaviors can be simplistically
represented by straight lines. By following a zigzag route, an insect can use only innate behavior to get from
point "A" to point "B". But a learned behavior, superimposed on this innate grid, might provide a
"shortcut" that is more useful or more efficient. As in the locust example above, the innate ability to fly may
be refined and improved through experience.

Periodic Behavior
 20

Insects, like most other living

organisms, exhibit cycles of activity that
are synchronized with periodic events in
their environment. Hatching, molting,
foraging for food, courtship, and nest
building are all examples of common
behaviors that may recur at regular
intervals. When these periods of activity
are repeated at about the same time
each day, an animal is said to exhibit
diel periodicity (a daily activity cycle).
An actograph is a device that entomologists use to record
movement (or other activity) of a caged insect. Data from an
actograph is usually displayed by a chart recorder - much like
a seismograph registers earthquakes. Mouse over each term
highlighted at the left to see a corresponding pattern of daily
actograph data.

Diurnal insects are active during

daylight hours
Nocturnal insects are active at
night
Crepuscular insects are active
at dawn and dusk

Two additional sub-categories exist to

differentiate dawn and dusk periods:

Matinal behavior occurs only

near dawn
Vespertine behavior occurs only near dusk

In addition to daily rhythms, many insects also exhibit periodic behaviors with much longer cycles:

Circalunar rhythms are synchronized with the phases of the moon and have a cycle of about 28
days. Many insect species tend to molt or emerge as adults during the dark phase of the moon as
an adaptation to evade predators.
Circannual rhythms are synchronized with seasons of the year and have a cycle of about 12
months. Examples include molting in periodical cicadas and migratory flights in monarch butterflies.

Entrainment

Environmental cues such as temperature, light intensity, or location of the sun provide the information
animals need to synchronize their behavior with the earth's daily, monthly, or yearly cycles. The process of
establishing and maintaining this synchrony is commonly known as entrainment. In insects, entrainment of
periodic behavior usually occurs in one of two ways:

3. Exogenous entrainment involves direct response to environmental cues that "trigger" the onset and
termination of a particular behavior. In some species of crickets, for example, males begin chirping
at nightfall in response to low light intensity and they continue until the light returns at dawn. These
crickets will also chirp during a solar eclipse or any time they are covered by a box that keeps out the
light. They will not chirp at all if they are exposed to constant illumination. As the name implies,
exogenous behavior is regulated by forces "outside" the organism. An experimenter can control
when crickets chirp simply by changing the light intensity in their environment.
4. Endogenous entrainment involves an internal biological clock that "measures" the passage of time
and sends "start" and "stop" signals to the nervous system. These periodic behaviors are often
called circadian rhythms. In a cricket species where chirping behavior is controlled endogenously,
the males will always start to chirp around dusk, regardless of light intensity. They will not be fooled
into chirping during a solar eclipse, or when covered by a box, and they will start chirping around
dusk even if the lights are always on. Start and stop signals are generated from "inside" the
organism and cannot be arbitrarily changed by an experimentor.
 21

Even with this internal chronometer, an animal's circadian rhythms must still be synchronized with the
environment. As day lengths change throughout the year, insects have to adjust their activity cycles to
compensate. In fact, a slight adjustment is made each day in response to a cue (usually light or
temperature) from the environment. This signal, which serves as a daily "reset" mechanism for the biological
clock, is called a zeitgeber - derived from the German word for "time" (zeit) and the verb "to give" (geben).

An actograph of an entrained insect shows a constant pattern of diurnal activity under a regimen of 8
hours light and 16 hours dark. When held in constant light (or constant dark), the same insect shows
a phase shift in behavior as its biological clock becomes "free-running."

In the complete absence of any environmental cues, a biological clock becomes "free-running". Its
endogenous period is usually between 24.5 and 25.0 hours, so the animal's behavior tends to occur a little
later each day as the internal clock gradually drifts out of phase with the environment. Humans experience
this same phenomenon in sleep/wake cycles when housed under conditions of constant light or constant
dark. We also experience "jet lag" when we travel to a distant time zone. In both humans and insects, it
takes several days for the body to adjust to a new photoperiod regime.

Recent discoveries in insect genetics and physiology have revealed that there are different biological clocks
for different rhythms. The clock that controls when a fly lays an egg may not be the same as the clock that
controls when the fly forages for food. Biological clocks are emergent properties of individual cells and may
be widespread throughout the body. Although they regulate different functions, they all appear to have a
common genetic origin. In fruit flies (Drosophila spp.), the clock mechanism consists of four regulatory
proteins that interact through a negative feedback system. The clock cycle begins when two of these
proteins (named JRK and CYC) bind together in the nucleus where they "activate" genes that produce two
other proteins (PER and TIM). As the levels of PER and TIM accumulate over timethey inactivate the genes
that produce JRK and CYC. This, in turn, leads to reduced production of PER and TIM and, eventually,
renewed production of JRK and CYC to complete the cycle. Normally, this system completes one cycle
every 24 hours. Mutations in the genes that encode these four clock proteins can change the duration of the
cycle or completely disable the clock mechanism.

Periodic behavior is certainly not unique to insects -- it occurs, to some extent, in nearly every living
organism on earth. To a biologist, this fact suggests that there are strong selective pressures acting to
preserve the cells' genetic mechanisms for periodicity. For insects, daily periods of inactivity serve as an
opportunity to conserve energy and resources. Many mosquito species, for instance, seek a humid, shady
habitat during the day to avoid dehydration. Likewise, honeybees (Apis mellifera) return to the hive at dusk
and pass the night in a quiescent state that has been described as "sleep". For other species (e.g.
cockroaches and stoneflies) circadian rhythms in emergence and/or molting may reduce the threat of
predation. And finally, synchronized periods of activity within a species can expedite the timing of courtship
behavior and reduce the metabolic costs of reproduction. It's all a matter of timing!
 22

Insect Communication

Communication may be defined as any exchange of information between

individuals. For members of the Human species, it is an essential part of all social
interactions. We communicate through speech, written language, sign language, body
language, Braille, Morse code, and many other cultural and technological
inventions. There is even some evidence that we use body odor in
communication. Our brains are uniquely adapted for symbolic communication, but
most of our "language" skills are acquired through learning.

Insects also have many ways to communicate but, unlike humans, their "language" is almost entirely
innate. Each individual is born with a distinctive "vocabulary" that is shared only with other members of its
own species. Learning plays little or no role in the ability to produce these signals or to understand them.

An act of communication is not always overt or obvious. No physical entity passes from one individual to
another, so it is not always possible to know when exchange of information occurs. The situation is
analogous to an alien from another planet who comes to Earth and observes human behavior. Without
knowing our language and customs, the alien would be unlikely to recognize a black arm band, a coy wink,
or a "yellow ribbon 'round the old oak tree" as forms of human communication. We are in much the same
predicament when we study insect communication. The only way to distinguish communicative behavior
from non-communicative behavior is by looking for evidence of a change in the behavior (or sometimes,
physiology) of another individual.

For experimental purposes, ethologists (scientists who study animal behavior) often define communication
as:

An action or condition on the part of one organism that alters the behavior of another organism in an
adaptive way.

Thus, an insect may send a communication signal by doing something (e.g. make a noise, release a
chemical, or flash a light) or the signal may simply be an inherent part of the insect's physical makeup (e.g.
wing pattern, body color, or surface chemistry). In either case, the signal must elicit some behavioral change
in order for a human observer to recognize its existence.

Why do insects communicate?

Some form of intraspecific communication is a prerequisite for any behavior that involves the participation or
cooperation of two or more individuals. Intentionally or not, insects may also communicate with members of
other species (interspecific communication). The adaptive value of these communication signals may
include:

10. Recognition of kin or nestmates

11. Locating or identifying a member of the opposite sex
12. Facilitation of courtship and mating
13. Giving directions for location of food or other resources
14. Regulating spatial distribution of individuals -- aggregation or dispersal;
establishing and maintaining a territory
15. Warning of danger; setting off an alarm
16. Advertising one's presence or location
17. Expressing threat or submission (agonistic behaviors)
18. Deception / mimicry

How do insects communicate?

Like all other animals, insects use their five senses to acquire information about their environment; any of
these sensory modalities may serve as a pathway for the exchange of information. Taste and touch are both
contact senses, therefore, exchange of information can occur only when two individuals are touching one
 23

another. Vision, olfaction (smell), and hearing are remote senses -- information signals may propogate
through the air (or water) over considerable distances.

Each signal modality has unique advantages and disadvantages. Follow the links below to find examples of
each signal system and learn more about insect communication.

Tactile Communication

"Keep in touch!" For you, it's probably just a metaphor, but for some insects it's really a channel of
communication. Since many insects have poor vision and sound perception, physical contact provides an
important avenue of communication. In blister beetles (family Meloidae),
courtship begins with a series of antennal taps by the male on each side of
the female's body. She signals her receptivity by lifting her wing covers
(elytra) and allowing him to climb on her back. But to complete his quest,
the male must continue tapping, alternating from side to side at just the
right frequency until the female is stimulated to extend her genitalia and
begin mating.

Antennal tapping is also an essential component of communication in both ants and

termites. It's not clear exactly what information may be exchanged, but it certainly
involves nestmate recognition and leads to exchange of food through trophallaxis. In
some cases, instantaneous feedback allows the sender to monitor the receiver's
response and alter the signal if necessary. Antennal tapping on the hind legs is used
during tandem running in both ants and termites. This is a "follow-the-leader"
behavior in which the tapping informs the leader that she has not lost her disciple. If
tapping stops, the leader instinctively turns around and searches in ever-widening
circles until she re-establishes contact with the follower.

The "dance" language of honeybees is largely a tactile communication system, performed in total darkness
on the vertical surface of the honeycomb. A "round dance" signals to nestmates the presence of a nectar
source in close proximity to the hive (usually less than 80 feet). It consists of a series of circular runs with
more or less frequent changes in direction. The greater the frequency of direction changes, the better the
quality of the nectar source. The "waggle dance" is used for longer distances. It involves a figure eight
pattern with a series of abdominal waggles on a straight run after each half-circle turn (see figure at
left). Distance is indicated by the duration of the straight run and the frequency of the waggles. Direction is
indicated by the angle of the straight run (relative to vertical) and corresponds to the horizontal angle
between the sun and the direction of the food source.

Tactile cues generated by ripples on the water surface allow whirligig beetles (family Gyrinidae) to constantly
monitor the location of dozens of other nearby whirligigs. Thanks to this tactile communication system, the
whirligigs can swim rapidly in circles, avoid bumping into other members of their own species, and still detect
the presence of nearby predators or prey.

Certain treehoppers (order Hemiptera: family Membracidae) produce vibrations in the tissue of their host
plant that can be felt by all other treehoppers on the same plant. The signal travels throughout the plant in
much the same way that banging on water pipes in your apartment creates noise throughout the whole
building. The signals apparently work as an alarm system, and in some species, they may be used by
nymphs to elicit protective maternal behavior. Substrate vibrations can be a particularly effective
communication system for small insects who cannot generate an acoustic signal loud enough to be heard
more than few inches away.
 24

Acoustic Communication Pros and Cons of Tactile Communication

Advantages:

Instantaneous feedback
Sound is localized area
Individual recipient
Effective in the dark (e. g. caves, wood galleries)

Disadvantages:

produced when a physical object Not effective over distance

vibrates rapidly, disturbs nearby air Organisms must stay in direct contact
molecules (or other surrounding Message must be repeated to each recipient
medium), and generates Vibration signals can be intercepted by predators
compression waves that travel in all
directions away from the
source. We perceive these waves as sound when they collide with our ear drum and cause a mechanical
disturbance that is detected by sensory neurons in our inner ear. Both living and non-living objects produce
sound, but only animals use it as a means of communication.

Sound serves as a very effective communication modality. It can be made to vary in frequency (high pitch
vs. low pitch), amplitude (loudness), and periodicity (the temporal pattern of freqency and
amplitude). Together, these three variables can create an extremely wide and complex range of signals --
from an insect's mating call to human speech and vocal music. Since sound waves move rapidly through air
(about 331 m/sec), acoustic signals can be quickly started, stopped, or modified to send a time-sensitive
message.

At best, the human ear is able to Pros and Cons of Acoustic Communication
detect sound frequencies only Advantages:
within the range of about 20-20,000
hertz (vibrations per second). But Not limited by environmental barriers
some insects (as well as other
animals like bats and dolphins) Effective over distances and around corners
produce and detect sounds that are
well above this frequency Highly variable, fast change -- high information
range. Some grasshoppers and content
moths, for example, produce
ultrasonic sounds as high as 80,000 Disadvantages:
hertz. Entomologists study these
high-pitch sounds by using an audio May reveal location of sender to a potential predator
transducer, an electronic device
that converts inaudible high Less effective in "noisy" environments (e.g. seashore)
frequencies to lower audible
frequencies. May be metabolically "expensive" to produce

Most insects detect sound with a Attenuation -- intensity falls rapidly with distance from
tympanic mambrane in the source (cube-root function)
abdomen (e.g. grasshoppers and
moths) or in the tibiae of the front
legs (e.g. crickets and katydids). Mosquitoes have antennal hairs that resonate to certain frequencies of
sound. But sound vibrations can also travel through solid objects, and some insects (e.g. some species of
ants, bees, termites, and treehoppers) can sense substrate vibrations with mechanoreceptors (chordotonal
organs) in their legs. Since these signals are "felt" rather than "heard", they are usually regarded as a form
of tactile communication.

See and hear each insect in the table below by clicking on its common name.
 25

Chemical Communication

It is probably safe to say that insects rely more heavily on chemical signals than on any other form of
communication. These signals, often called semiochemicals or infochemicals, serve as a form of
"language" that helps to mediate interactions between organisms. Insects may be highly sensitive to low
concentrations of these chemicals -- in some cases, a few molecules may be enough to elicit a response.

Semiochemicals can be divided into Pros and Cons of Chemical Communication

two groups based on who "sends" a
message and who "receives" it: Advantages:

3. Pheromones are chemical Not limited by environmental barriers

signals that carry information Effective over distances and around corners
from one individual to Effective either day or night
another member of the same
Longer lasting than visual or auditory signals
species. These include sex
Metabolically "inexpensive" because only small
attractants, trail marking
quantities are needed
compounds, alarm
substances, and many other
intraspecific messages. Disadvantages:
4. Allelochemicals are signals
that travel from one animal to Low information content (presence/absence)
some member of a different Not effective in an upwind direction
species. These include
defensive signals such as
repellents, compounds used to locate suitable host plants, and a vast array of other substances that
regulate interspecific behaviors.

Allelochemicals can be further subdivided into three groups based on who "benefits" from the meassage:

4. Allomones benefit the sender -- such as a repellent, or defensive compound (e. g. cyanide) that
deters predation.
5. Kairomones benefit the receiver -- such as an odor that a parasite uses to find its host.
6. Synomones benefit both sender and receiver -- such as plant volatiles that attract insect pollinators.

Grape root borer moths (Vitacea polistiformis).

Male attracted by female's sex pheromone.

(click the "Play" button for video)

Insects use their sense of taste or smell to detect the presence of semiochemicals. Specialized receptors
may be located anywhere on the body, but are especially common on the feet, antennae, palps, and
ovipositor (see Chemoreception). The sense of smell (olfaction) is used for remote chemoreception --
detecting semiochemicals with low molecular weight that are volatile enough to become airborne. The sense
of taste (gustation) is used for contact chemoreception -- detecting molecules that adhere to a substrate or to
the outside of an insect's body.

Which type of signal is involved (pheromone, allomone, kairomone, or synomone)?

 26

5. A parasitic fly, Euclytia flavahe, is attracted to the odor of its host.

6. Ants are able to recognize their nestmates by the taste and smell of hydrocarbons (lipids) found on
the surface of the cuticle.
7. Caterpillars that feed on plants sometimes unwittingly induce those plants to release a defensive
compound that attracts parasitoids of the caterpillars.
8. Caterpillars of Lycaena arion produce a chemical that attracts ants and elicits care-giving behavior.
When carried into the ant's nest and placed among the brood, these caterpillars will eat the ant
larvae.

Visual Communication

Many insects communicate with visual signals. The color patterns and other markings on the
wings of butterflies and moths facilitate species recognition in much the same way colored
uniforms reveal the players' affiliations on a football field. Some insects use bright colors,
eyespots, or other distinctive patterns to scare away predators, to advertise their ability to sting, or
to mimic the appearance of another unpalatable species. Other insects use dance-like body
movements to attract a mate or to communicate with nestmates. Most of these signals are effective only as
long as they are visible in daylight. But a few insects (fireflies, for example) can generate their
own light and use visual signals that can be seen at night.

Passive signals, Pros and Cons of Visual Communication

such as
eyespots and Advantages:
color patterns
can serve as a
Effective over long distances
form of "free
advertising". The colorful wings Can be used while moving
of a butterfly, for example, are a Fast -- speed of light
"billboard" publicizing its Effective in all directions (independent of wind)
species identity. Individual Passive signals require no expenditure of energy
insects incur little or no
metabolic cost for displaying Disadvantages:
these messages because they
are an integral part of the Requires a clear line of sight
integument. It may be prudent
Visual signals may be intercepted by predators
to hide these signals from a
Only effective in daylight (in fireflies, only at night)
potential predator, so some
insects have a way to conceal Active signals may be metabolically "expensive" to produce
their message when
necessary. The red admiral butterfly, for example, has bright, distinctive markings on the upper wing surface
and drab, protective coloration on the underside.

Active signals, like body movements and light flashes, are more costly to
produce, but they can be withheld from use at inappropriate times. They
may also have a higher information content because signal frequency,
duration, or periodicity may convey additional meaning. In fireflies, for
example, pulses of light are used in a courtship dialogue between a male
(usually flying) and a female (usually perched in the vegetation). Each
species has a unique flash pattern and response time. Males of Photinus
pyralis emit a single "J"-shape flash during a rising flight movement. A
female responds with a single flash after a two second interval. In Photinus consumilis, males emit a series
of 3-5 short flashes and the females respond with a double flash. Roles are reversed in some tropical
species where the females fly and the males signal from perches in the vegetation.
 27

Unlike humans, many insects have the ability to see ultraviolet light. Not surprisingly, some species
communicate using wavelengths in this part of the spectrum. Female cabbage butterflies, for example, have
ultraviolet-reflecting scales on the dorsal wing surface. When they fly, each downstroke of the wing creates
a brief "flash" of UV that males apparently recognize as the flight signature of a potential mate. A "flashing
female" may attract several males who engage in aerial courtship displays.

Ultraviolet reflection from a male

alfalfa butterfly in simulated flight

In alfalfa butterflies, only males have the UV-reflective scales. They flutter in
front of the females to create a flickering courtship display. Missing scales
reduce the wings' reflectivity -- a sign of aging that impairs a male's ability to
seduce a mate.

Social Insects

Social insects are among the most dominant and prolific of all organisms on earth. In South America,
leafcutter ants (Atta spp.) consume more foliage than mammalian herbivores; in the southwestern
United States, more seeds are collected and eaten by harvester ants than by all other forms of life; in
some tropical habitats, the biomass of ants and termites exceeds that of all other animal species
combined; in the African savanah, a single colony of driver ants may contain as many as 20 million
workers; in Japan, a supercolony of Formica yessensis with 45,000 interconnecting nests contained
more than a million queens and 306 million workers within an area of 2.7 square kilometers.

Many insects exhibit "social" behaviors (e.g. feeding aggregations, parental care of the young, and
communal nest sites). In a broad sense, any insect that interacts with another member of its own species
could be called a social insect. But as a rule, entomologists do not regard these behaviors as sufficient
justification for classifying a species as truly social (i.e. eusocial). In order to qualify as eusocial, a species
must exhibit all four of the following characteristics:

5. Share a common nest site

6. Individuals of the same species cooperate in caring for the young
7. Reproductive division of labor -- sterile (or less fecund) individuals work for the benefit of a few
reproductive individuals
8. Overlap of generations -- offspring contribute to colony labor while their parents are still alive

Species that lack one or more of these characteristics are classified as presocial. Within this category are
subsocial species (in which the parents care for their offspring) and parasocial species (which have a
common nest site but lack one or more of the other eusocial characteristics). The table below helps clarify
the various terms used for social and presocial insects. Click on each term for a formal definition.

A Classification System for Presocial and Eusocial Insects

Common Cooperative Reproductive Generational

nest site brood care castes overlap

Solitary No No No No

Communal or Yes No No No
Subsocial

Quasisocial Yes Yes No No

 28

Semisocial Yes Yes Yes No

Eusocial Yes Yes Yes Yes

Relatively few insects are classified as eusocial -- the distinction is limited to the following groups:

Order Isoptera Order Hymenoptera

Termites -- all species Ants -- all species (family Formicidae)

Bees -- about 600 species in the family Apidae

Wasps -- about 700 species in the family Vespidae

Termites

All members of the order Isoptera are eusocial insects. Termites feed primarily on the cellulose and lignin
found in plant cell walls; these compounds are the main ingredients of wood and all paper
products. Termites cannot digest the cellulose directly so they rely upon symbiotic bacteria and protozoa
living within their intestines to supply most of the enzymes needed for cellulose digestion. Termites are
sometimes called white ants. They may resemble ants in size, but ants have a narrow waist and elbowed
antennae while termites have a thick waist and antennae that resemble a string of beads.

Ecologically, termites play an important role in the environment by helping to break down and recycle dead
wood and other plant tissues. They become pests when their appetite for wood and wood products extends
to human homes, fence posts, building materials, cardboard, and other valuable products. In tropical and
subtropical forests where termites are abundant, railroads must use expensive metal ties because wooden
ones are quickly destroyed.

Ecologically, termites play an important role in the environment by helping to break down and recycle dead
wood and other plant tissues. They become pests when their appetite for wood and wood products extends
to human homes, fence posts, building materials, cardboard, and other valuable products. In tropical and
subtropical forests where termites are abundant, railroads must use expensive metal ties because wooden
ones are quickly destroyed.

The Termite Colony

 29

Each termite lives in a nest or colony with

hundreds, thousands, or even millions of its << Termite Social Register >>
brothers and sisters. In fact, the termite colony
is really a large, extended family. Within this
Queen - Fertile adult female.
family, various groups of individuals have Reproductive
different functional roles according to a "caste Caste
system". King - Fertile adult male.

The worker caste Non fertile males and females with

Soldier
is the largest special morphological adaptations
Caste
group. It consists for defense of the colony.
entirely of
immatures, both
males and Immature males and females
females. These (nymphs) - offspring of the king and
Worker
soft-bodied, wingless individuals perform all of queen, with potential to molt into
Caste
the hard labor in the colony: they clean, replacement soldiers or
maintain, and repair the nest, gather food and reproductives if needed.
water, care for the young, and construct new
tunnels and galleries as the colony grows. These juveniles all have the genetic capacity to undergo
additional molts and become soldiers or reproductives, but most will spend their entire lives as workers.

Members of the soldier caste are larger in size but fewer in number than the workers.

They are also wingless, but they have large heads with powerful jaws. Their job is
to guard the nest site and protect it from attacks by
ants or other invaders. In some species the soldiers
lack jaws but have a large gland in the head that
shoots defensive chemicals through a
nozzle at the front of the head. The soldiers are
unable to care for themselves so they must
be fed and groomed by the workers.

The reproductive caste always includes a king (male) and a queen (female) who are the parents of the
termite family and founders of the colony. Some species also have a few supplemental reproductives who
share the egg laying duties. These are the only adult insects in the colony. The queen lays large numbers
of eggs which develop into more workers and soldiers as the family grows.

In every mature colony, there also develops an annual population of young winged
reproductives that swarm from the parent nest for a short mating flight. After flight, the
delicate wings break off, and the new king and queen set out to find another nest site
and start a new colony. Large colonies with multiple reproductives may also split into
two or more daughter colonies, a process known as "budding".

The termite's caste system is regulated by

pheromones. The king and queen each produce special pheromones that
circulate throughout the colony and inhibit workers of the same sex from
molting into reproductive adults. A death in the royal family (or an increase in
the size of the colony) results in a lower concentration of the corresponding
pheromone and, subsequently, one or more workers will molt into
replacement reproductives. Likewise, the concentration of sex-specific soldier
pheromones regulate the numbers of male and female soldiers to fall within
an optimal range based on colony size. Excess numbers of soldiers or
reproductives may be killed and eaten by the workers.
 30

Kinds of Termites.

About 2750 different species of termites are known. These can be divided into two groups: those that live
entirely within wood, and more advanced species that tunnel and nest in the soil. In terms of their ecology
and behavior, the most primitive species are similar to certain wood-dwelling cockroaches with whom they
may share a common ancestor. These primitive species often have specialized habitat requirements,
nesting only in rotten wood, damp wood, or dry wood. Their colonies are rather small and persist only as
long as the food resource lasts. All wood-dwelling termites produce distinctive waste pellets which are often
the first sign of an active infestation.

Subterranean termites construct underground nests and have the ability to tunnel through the soil to find new
food resources. These colonies are often long-lived and may grow to include several million
individuals. The subterranean termites that live in North America and Europe often invade wooden
structures above the ground by building earthen tubes that serve as protective tunnels between the nest and
their food source. These tubes are good evidence of a termite infestation.

In Africa and Australia, other subterranean species mix bits of soil with saliva to build nest mounds that are
up to 20 feet (6 meters) tall. The inside of the mound is divided into numerous chambers and galleries. The
king and queen live in a special cell deep inside the mound. The females abdomen grows in size until it is
large enough to hold many thousands of eggs. The queen
lays these eggs at the rate of several thousand a
day. Worker termites carry the eggs away to specially
constructed cells in the nest. There the workers care for the
young as they hatch from the eggs.

Some of the mound-building termites cultivate underground

fungus gardens. They collect dead plant material, mix it with
saliva and their own waste products to create a paste, and
inoculate this substance with the spores of a symbiotic
fungus. The termites feed on special structures produced by
the growing fungus.