Introduction to

The Manduca Project

Introduction 1
 The Manduca Project
In 1991, a science outreach program for grades K-3 called
The Manduca Project was initiated through the joint efforts
of the University of Arizona Department of Biochemistry and
Tucson Unified School District (TUSD). This project provides
resources for student explorations with Manduca sexta. It has
demonstrated the insect’s usefulness both as a research subject
and as a bridge between the University and the public schools.
The Manduca Project began simply—teachers received
caterpillars, artificial diet, instructions, and help with planning a
week of classroom activities around the insects. Soon, teachers
and their students expanded this activity into interdisciplinary
lessons covering weeks and even months.

2 The Manduca Project

Insects in the classroom
Insects lend themselves particularly well to exploratory classroom
activities. Insects are easy to find, and there are several that are
simple to raise in the classroom. Insects give students hands-on
experience observing and designing experiments with live animals,
experience that cannot be duplicated using books and models. Most
importantly, students of all ages are fascinated by insects. They
capture students’ attention and open their minds—crucial steps in
the learning process. When the mind is open, anything can happen!

Why Manduca sexta?

The tobacco hornworm (Manduca sexta) is useful for examining a
diverse array of biological problems. Scientists around the world
study this insect to answer questions ranging from ecology and
growth to agriculture, neurobiology, muscle physiology, and the
biochemistry of camouflage.
Why use Manduca in research? First, they are good model
physiological systems—like us in many ways, but much simpler.
Second, they are safe, easy, and inexpensive to raise (compared to,
say, African elephants). Finally, there is little controversy regarding
using non-endangered insects in scientific research. For these
same reasons, Manducas make excellent subjects for raising and
investigating animals in the classroom.

Introduction 3

Chill out!
Wiggling larvae are difficult
to handle. Before working
with larvae, they’re chilled
in a bucket of crushed ice
to slow them down.
Processing Manducas
Carefully cutting them open.
Extracting hemolymph (blood).
Removing the midgut (stomach).
4 The Manduca Project
Manducas in research
Why use Manduca?
Manduca sexta is an invaluable tool for scientific research. It
has been the subject of years of investigations ranging from
behavioral studies to biochemical and genetic processes to
helping model how muscles and hormones are affected by space
travel. Manduca is easy and inexpensive to raise, is relatively large,
has simple body systems, has a complete (and relatively fast) life
cycle, and is free of the ethical issues surrounding more complex
or endangered organisms. For these same reasons, they are also
ideal experimental subjects for the classroom.
Biochemistry
Lipid metabolism
Lipids are a group of chemical
compounds that include fats,
oils, and similar substances
that are not soluble in water.
A good example is corn oil.
Lipids are a necessary and
important part of most animals’
diets. Along with proteins and
carbohydrates, they are a basic
building material for cells.
Manducas are easy to raise
Lilian Canavoso collects hemolymph from about
in large numbers, are relatively 200 Manducas per day to study the process of
large as insects go, and are lipid metabolism.
basically little green food
processors. That makes them ideal living laboratories for learning
how animals metabolize (digest) lipids.
When Manducas eat, the food enters the large stomach or mid-
gut, where enzymes break down the lipids and transport them
through the walls of the mid-gut out into the hemolymph, or
blood. The hemolymph carries the metabolized lipids throughout
the Manduca’s body. Some is used right away as the larva grows,
and some is stored in the fat bodies—structures surrounding
the midgut—to be used during molting and pupation, when the
Manduca can’t eat.
Researchers feed Manduca larvae special lipids containing
radioactive tracers that allow them to measure how fast the
lipids are digested, and to identify exactly where they go. The
mid-guts can even be removed from the larvae and kept alive (and
metabolizing) for up to two hours!
This research is helping scientists better understand how
lipids are processed by animals (including humans). Practical
applications are in food and nutrition, disease prevention and
treatment, and in insect pest control. (Many insecticides are
toxins dissolved in a lipid base.)

Pheromones
Pheromones are specialized chemical compounds produced by
animals that affect the behavior of other members of their
species. Female Manducas produce sex pheromones that attract Males vs. Females
males for mating. Researchers are studying both the production When it comes to sex
of these pheromones in females and the sensory systems of pheromones, male and
female Manducas are very
males. Scientists are now able to create synthetic Manduca different. Females produce
pheromones. These are being used to study ways to control pheromones, but don’t
Manduca reproduction as a means of pest management, as well have the specialized
as controlling Manduca’s flight behavior. sensory organs to detect
them. The males have a
highly-developed sense of
Parasitism smell for detecting
pheromones, but don’t
Parasites are organisms that grow or feed in or on a “host” produce them.
organism without providing any benefit to the host. In nature, Scientists studying the
tiny wasps of the Polistes family puncture the skin of Manduca chemistry of pheromones
larvae and lay their eggs. When the eggs hatch, the wasp larvae work primarily with
burrow back out through the skin and attach themselves to the females, while those
interested in olfaction
caterpillar. There, they “hitch a ride”, feeding off the Manduca larva (smell) work with males.
until they reach adulthood.
The important feature of this parasitic relationship is that
the Manduca larvae’s normally
Courtesy of Clemson University Dept. of Entomology.

strong immune system fails to

notice that it has been invaded.
Researchers have found that,
along with the eggs, the wasp
injects a virus that interferes
with the Manducas immune
response. These wasps are
already used in agriculture as Hitchhikers? Larvae of the wasp Cotesia
a part of integrated pest congregata parasitize a fifth instar Manduca.

Introduction 5
 Photos courtesy of Chip Hedgecock, ARLDN
Skip Vaught prepares an
experiment to study odor-
guided flight in Manducas.
Photo courtesy of Chip Hedgecock, ARLDN
Researcher Pete Test uses a
dissecting microscope to
implant electrodes in a
Manduca’s brain.
6 The Manduca Project
management, for controlling the plant-munching Manduca.
Scientists are trying to better understand the biochemical
process by which the virus works, in hopes of using this knowledge
to develop better and safer pest control techniques, and more
effective treatments of immune system disorders in humans.
Neurobiology
Neurobiologists at the University of Arizona are studying
Manduca to learn about how brains are “wired.” Manducas make
great study animals because their brains are very similar to the
brains of other animals, only much simpler. Scientists learn what
makes moths tick by observing their behavior and studying how
their brains work in response to pheromones and plant odors.
Many of the scientists at the Arizona Research Labs Division
of Neurobiology (ARLDN) at the University study olfaction in
the adult moth. Olfaction is the scientific term for “smelling.”
Nearly all animals depend
on their sense of smell for Photo courtesy of Chip Hedgecock, ARLDN
their survival. In Manduca,
like other insects, smell
is especially important for
locating food sources,
mates, and places to lay
eggs. Moths have an
extraordinary sense of
smell—some scientists
think it’s a million times
more sensitive than that Dr. Tom Christensen of ARLDN prepares a wind tunnel
of humans. experiment to study the production and release of sex
pheromones in female Manducas.
Olfaction
Some ARLDN scientists are looking at which smells attract
moths. Some odor sources, like the chemicals found in
pheromones or in the tomato plant, are important to the moth’s
survival. Moths innately recognize these important odors and will
fly toward their source, while ignoring strange odors that they
don’t typically encounter in the wild. Moths are tested either in
laboratory wind tunnels or outdoors, in large netted enclosures.
Once researchers identify a group of odors that attracts the
moths, neurobiologists study the moths’ brains and nervous
systems. They use extremely small electrodes to measure the
brains’ reaction to a variety of plant and pheromone odors. After
the brain activity is recorded, fluorescent dyes are injected into
the part of the brain where the recordings were taken. Later,
Photo courtesy of Chip Hedgecock, ARLDN
At left, graduate student Katja
Selchow monitors Manduca
data during a wind tunnel
experiment.

Wired for flight

Tiny instrument packages
have been developed that
can be attached to the
moth without affecting
its ability to fly. The
package includes a radio
that transmits data from
scientists cut the brain into very thin sections to see, using a the moth’s nervous
microscope, exactly which group of cells reacted to a particular system to a receiver. (A
odor. After many studies of this kind, they can begin assembling a major challenge has been
“map” of the brain and how it processes odor information. finding a lighter battery
for the instrument!)
Odor-guided flight
Understanding how the adult Manduca’s brain responds to familiar
odors opens up many intriguing possibilities. Dr. John G. Hildebrand,
director of the ARLDN, is leading research into practical uses for Other research
these “talented” moths. Some possibilities include using them to Elsewhere, scientists are
using their understanding
“sniff ” for drugs, explosives, biological and chemical weapons, toxic of the Manduca’s bio-
chemicals, or other environmental hazards. chemical, neurological, and
mechanical functions to
In the not so distant future, scientists may be able to guide the develop an amazing array
insect’s flight by “remote control.” This could be done by applying of new devices, many with
either electrical stimuli directly to the brain or “puffs” of odor to potential biomedical uses:
• Biobots—organisms
the antennae. These “RC Moths,” equipped with a radio “backpack,” equipped with devices to
could be used for environmental monitoring and other situations control their behavior and
where the use of humans or mechanical robots is either unsafe or interact with the environ-
ment.
too expensive. • Biosensors—devices
that combine biological
tissues with synthetic
devices to perform
sensory tasks, such as
artificial “super noses”.
• Biomimetics—artificial
devices that mimic the
neurology and mechanics
of organisms, such as
self-guided, odor-
following robots.

Introduction 7
 Learning with Manduca
Using Manduca sexta, students learn observation skills by keeping
detailed logs of the insects’ growth and behavior. They learn the
elements of graphing by plotting the growth of the insect using
simple student-made balances to measure mass, and string to
measure changes in length. Students use Manduca as the basis
for writing, art, poetry, and music projects. Typically, individual
students have their own insect to observe and care for, instilling a
sense of responsibility and heightening their observation skills.

Still have questions?

For answers to Manduca
science questions, e-mail
Dr. Michael Wells at:
mawells@email.arizona.edu

For answers to questions

about rearing Manducas
and ordering Manduca
eggs and diet, contact Looking ahead
Mary Hernandez at:
520/626-4373 The Manduca Project continues to evolve, both in terms of its
520/621-1772 breadth and number of participants. The University’s Department
of Biochemistry sponsors workshops for elementary educators
throughout the Tucson area. To date, more than 15,000
students have studied Manduca sexta in the classroom. The
program’s success is a direct result of the teachers’ dedication,
creativity, and energy in developing lessons and in recruiting
and training other teachers. We hope that these participating
teachers will continue to share their ideas with us and with each
other as we introduce The Manduca Project to a wider audience
across the country.
For more information, updates, and additional classroom
activities, visit The Manduca Project web site at:
http://insected.arizona.edu/manduca/

8 The Manduca Project

The World of
Insects

The World of Insects 9


Living in a world
without insects
Advantages
• No more “icky bugs”
• Agriculture without
pesticides *
• Reduce spread of insect-
borne diseases
Disadvantages
• *Without pollinators, crop
yields would plummet!
• Starvation of species
that eat insects
• Collapse of the food chain!
• We’d be buried in waste!
• Loss of insect products
like silk, honey, etc.
• A world with less wonder
and beauty!
Where would we be without the
latest Manduca larva fashions?
10 The Manduca Project
Insects all around us
Your life without insects?
Imagine for a moment, a world without
insects... “Wonderful!”, you say, “Finally I
can explore the great outdoors unbothered
by all those nasty creepy-crawlies!
Picnics without ants! Apples without
worms! Camping without mosquito
bites! Paradise! Where do I sign up?”
Before you get too excited, consider this.
Without insects, our world would be a very different place.
If insects didn’t play their roles as pollinators and as food
sources for other animals, our food supply and selection would be
drastically reduced. As a result, you might have trouble putting
together that “ant-free picnic” you were so looking forward to.
Likewise, a pleasant camping spot might be difficult to come by
in a world full of animal waste and dead plants and animals that
would exist without insects to help break them down.
While it’s true that some insects — like those bloodsucking
mosquitoes or plant-eating pests — directly harm humans, the
vast majority don’t. Rather, they’re all a vital part of this planet’s
ecology. They provide food for other animals (including humans
in some cultures), recycle natural materials, and help plants to
reproduce. They even create products we use every day, such
as silk and honey. On a scientific level, insects provide systems
for exploring and modeling human body processes. So, you see,
insects help us to better understand ourselves and the world
around us.
At the purely aesthetic level, insects provide our world with
beauty, drama, and wonder. They have, it seems, a limitless variety
of colors and shapes, behaviors and habits. As you look through
a hand lens at an insect, you enter a world more bizarre and
outrageous than any science fiction author could dream up.
They can be fascinating, entertaining, beautiful, or ugly. They are
insects, and they are everywhere.
So many insects
In terms of both number of species and number of individuals,
insects are a dominant form of life on Earth. There are somewhere
between 800,000 and 1,000,000 insect species known—
that’s more than all other animals combined! What’s more,
scientists estimate that with those insect species yet to be
discovered, there are between 80 and 100 million species of
insects sharing the planet with us.
The sheer number of individual insects is even more staggering.
It is estimated that at any given moment, Earth is home to a
billion billion insects. Spread out evenly over the land surface, this A billion billion?
would be nearly 8,000 insects per square meter! (About 750 How do you write out a
per square foot, five per square inch, or almost one per square billion billion as a number?
centimeter.) With six billion humans in the world, this works out to 1,000,000,000,000,000,000
170 million insects per person. Another way to say this
number is “one quintillion”.
To break it down another way, consider a single group of insects, No matter how you say or
the termites. If you weighed all of the termites in Africa, their write it, that’s a lot of
combined weight would be more than that of all of the elephants insects!
in Africa! Certainly, problems of poaching and
habitat destruction have taken their toll on
the numbers of elephants, but even the
idea of enough termites to outweigh one
elephant is mind-boggling!

They’re everywhere!
Not only are insects incredibly numerous, they successfully
occupy nearly every conceivable environment, and are constantly
adapting to new ones. Habitats range from the Arctic tundra and
glaciers, where you can find types of insects that are so cold-
adapted they will actually die of the heat if you hold them in your
hand, to the most scorching deserts.
Insects have adapted to life in pools of
crude oil in California and the Great Salt
Ahh…lovely weather
we’re having, hmm?
Lake in Utah, where the water is six times
saltier than the ocean. Some can be totally
dried out for months at a time, only to
“come back to life” when it rains. Insects
thrive in cities and rural areas, on land, in
water, and in the air. If you can imagine a
habitat, chances are that there will be at
least one species of insect making use of it.

The World of Insects 11

 The secrets to insects’ success
Why are insects so successful? There are many theories, but no
one knows for sure. One involves insects’ ability to fly. Anyone who
has watched a housefly land on the ceiling or been mesmerized by
the acrobatics of a dragonfly skimming over a pond can appreciate
the amazing feats accomplished by insects and their wings. From
a practical standpoint, flight allows insects to escape predators
and move easily from habitat to habitat in search of food and
potential mates.
“But wait!” you say, “If all it takes to be successful are wings,
how come we’re not overrun with birds and bats?” True, flight by
itself probably cannot explain all of insects’ success and diversity.
Another theory credits the fact that insects are, in general, fairly
small. Their small size allows them to occupy a wide range of
habitats other animals are unable to take advantage of. Habitats
can range from frozen tundra to hot, dry deserts to small,
temporary pools of water that collect in plants growing in tropical
rainforests — insects live everywhere! Insects’ hard, waterproof
outer shell or exoskeleton is a major contributor to their ability
to live practically anywhere. To some extent, the exoskeleton
protects insects from changes in their environment.
In addition to their small size, insects’ body structures are
incredibly diverse and highly adapted to their environments.
For instance, aquatic insects often have gills or other special
structures that allow them to breathe underwater.
Insects feed on almost anything. There are insects that feed
on leaves, nectar, pollen, roots, stems, and fruit of plants. This
includes some that eat leaves of plants that are poisonous to all
other animals. Some insects feed on the blood of mammals, while
others feed on the droppings left behind by those mammals. Like
termites, some ants even eat wood. Somewhere out there in the
world, there is probably an insect that can eat just about anything
you could imagine!
Another important factor that probably contributes to insects’
success is their ability to reproduce quickly and in great numbers.
Mammals usually have only a few babies at a time whereas many
insects can lay from hundreds to thousands of eggs at a time!
So, you see, there are many reasons why insects are a huge
evolutionary success story!

12 The Manduca Project

The insect “family tree”
With so many insects to account for, it’s not surprising that the
insects’ “family tree” or phylogeny can get pretty complicated. To
understand the insect family tree, you need to have an idea how
living things are classified and named. A taxonomy—a scientific
classification of organisms—is an important and useful tool.
There are several ways to classify living things. One of the
oldest and simplest taxonomies was developed in the 1730’s by
Swedish scientist Carl Linné, later renamed Carolus Linnaeus.

The Linnaean System

Linnaeus developed his taxonomy of living things “from the bottom
up”. He started by grouping individual species of plants with
similar characteristics into a group called a genus. Then, he
Carolus Linnaeus (1707–1778)
grouped similar genera (plural of genus) into families, similar
families into orders and so on, until all plants were joined into
one huge group, the Plant Kingdom. Later, Linnaeus repeated this Linnaeus’ Scheme
process for the animal kingdom. From the most general
level to the most specific:
Linnaeus used binomial (two-part) Latin names in his taxonomy, Kingdom
combining the genus (capitalized) and the species (not Phylum
capitalized) names, usually written in italics: Manduca sexta. Class
We still use this binomial system today. Here is the Linnaean Order
classification of the tobacco hornworm: Family
Genus
Species
Classification Name Description
Kingdom Animalia all animals

Phylum Arthropoda exoskeletons and jointed legs Why Latin?

Latin was the universal
Class Insecta insects - 6 legs, insect body plan language of science in
Linnaeus’ day. Scientists
Order Lepidoptera butterflies, moths, and skippers still use Latin because it
follows a strict set of
Family Sphingidae hawk moths grammatical rules and is
not used today by any
Genus Manduca culture, eliminating any
appearance of playing
Species sexta the tobacco hornworm “cultural favorites”.

To begin with, Manducas are arthropods — a large group that arthro = joint
includes such animals as lobsters, spiders, scorpions, centipedes, pods = feet
and millipedes, in addition to insects. All arthropods have hard
exoskeletons (skeletons on the outside) and jointed legs, traits
that distinguish them from all other animals.

The World of Insects 13


Allergy Alert!
Butterfly and moth scales
are antigenic. That means
that they can cause an
allergic reaction in some
people. For some, this
reaction can be very
severe, and they avoid
handling or working closely
with these insects.
Moth scales X 175
The scales easily detach
from the wings. This may
be an adaptation for
escaping spiders and
other predators.
14 The Manduca Project
Insect orders - cousins on the family tree
No one is sure how all the arthropods are related to each
other, but the relationships between the major orders of insects
are fairly well established (although like many other subjects of
scientific research, there is still lively debate about the placement
of some groups). Manducas belong to the order of “scaly-winged”
insects, the Lepidoptera. Lepidoptera include butterflies, moths,
and skippers.
Lepidoptera, the scaly-winged insects
The name Lepidoptera comes from two Greek words: lepis, which
means “scale” and ptera, which means “wing”. Of the 165,000
species of Lepidoptera identified to date, only 5% are butterflies;
the other 95% are moths.
If you look closely at the wings of butterflies and moths, you will
see that they are covered with tiny scales. These scales make the
colors and patterns on the wings of moths and butterflies, and
are commonly recognized as the powder that comes off the wings
when you touch them. In the pictures below, the wing of a Cecropia
moth is magnified to show the scales covering the wings and the
structure of the scales themselves.
Jim Kalisch, University of Nebraska at
Lincoln Entomology. Used with permission.
© 1997 Museum of Science, Boston. Used with permission.
Scale structure X 7,500
To fly, moths must be light. A
close-up of a scale shows that
it is mostly open space, adding
little weight.
© 1997 Museum of Science, Boston. Used with permission.
Phylogeny of major insect orders
A phylogeny, like the one shown below, is a lot like a family tree.
Groups of insects that are on branches near each other are
more closely related than those on branches farther apart. On
the insect phylogeny shown here, the groups nearer the top (e.g.
Springtails, Silverfish, etc.) are considered more “primitive”. This
means that they’ve been around for a very long time and are
relatively “simple”. The groups nearer the bottom (e.g. Beetles, Manny’s Full Name
Flies, Moths, etc.) are considered to be more “advanced”, and have
With so many insects to
evolved more complexity than the so-called primitive insects. classify, scientists must
use a more detailed
scheme of super-groups,
Springtails sub-groups, and
inter-groups of related
Silverfish organisms. Here’s a more
complete classification for
Mayflies Manny:
Animalia (Kingdom)
Dragonflies Metazoa
Arthropoda (Phylum)
Walking Sticks Hexapoda
Insecta (Class)
Grasshoppers Pterygota
Neoptera
Endopterygota
Praying Mantids Lepidoptera (Order)
Glossata
Cockroaches Neolepidoptera
Heteroneura
Termites * Yes, there
really is a large group of
Ditrysia
insects called “true bugs”. Sphingoidea
Earwigs These insects truly can be called
“bugs”! Some examples are the Sphingidae (Family)
Milkweed Bug, the Kissing Bug, and
the Giant Water Bug.
Sphinginae
Lice Manduca (Genus)
sexta (Species)
True bugs* Whew! Aren’t you glad you
don’t have to remember all
Lacewings that?

Beetles
Fleas
“It’s me - Manny!”
Flies
Moths & Butterflies
Bees, Wasps, & Ants

The World of Insects 15

 Insect anatomy
What makes an insect an insect?
So far you’ve learned a little bit about Arthropods and seen how all
of the major groups of insects are related to each other. By now
you may be wondering, “Just what is an insect, anyway?”.
In spite of the enormous amount of variation within insects, they
all have two things in common:
• Insects have three, and only three pairs of legs.
• Insects have three major body sections; the head, thorax,
and abdomen.
These features distinguish insects from the other arthropod
groups. All other arthropods have more than three pairs of legs,
and have either more or less than three body sections.
Insects seem to be put together inside-out and upside-down.
Their skeletons are on the outside of their bodies, their heart runs
in a thin line down their backs, and their nerve cords are found
along their bellies. Some even have more than one “brain”!
This figure shows the external anatomy of a “generalized” insect,
or one with very few modifications to the basic insect body plan:
Head Thorax Abdomen
Wings
Compound eye

Antenna
Spiracles

Legs
The insect’s external anatomy is only slightly less bizarre than its
internal anatomy. Moving from front to back:
• head — has eyes and antennae for sensing the environment,
and an assortment of mouthparts for consuming food.
• thorax — a three-segmented section that is the center for
locomotion. Legs and wings (for those insects which have
wings) are attached to the thorax.
• abdomen — this many-segmented section is the center for
digestion and reproduction. Breathing holes called spiracles
are found in a line along the sides of the thorax and abdomen.
These lead to a branching network of tiny tubes throughout
the insect’s body that supply oxygen to every cell.

16 The Manduca Project

Mouth Parts
One of the most interesting parts of insect anatomy is the
mouth. The most basic type of mouth is the simple “chewing type”.
Simple eyes

Compound eye Antenna (cut)

Maxilla Maxilla
Labrum

Palp Palp

Mandible Mandible

Palp

Labium

Even this “simple” mouth has what first appears to be a mind-

boggling assortment of parts. To begin with, there is the labrum, a
flap-like structure that acts more or less like an upper lip. Directly
beneath the labrum are the mandibles, or functional “jaws”. Unlike
our jaws, insect mandibles work side-to-side, rather than up-and-
down. Behind the mandibles are the maxillae; multi-segmented
structures that are partly for sensing and manipulating food, and
partly for chewing it. Finally, behind the other mouthparts is the
labium; which serves as a combination lower lip and, by way of the
attached palps, as a sensory structure.
Not all insects have all of these mouth parts. In many
groups, especially those more “advanced” ones, various parts
my be reduced or even absent, giving those insects’ very
different mouths from the ones pictured here. Mouth type is
often indicative of food and lifestyle choice; insects will have
mouthparts that best suit the environments in which they live.

The World of Insects 17

 The Life of
Manduca sexta

18 The Manduca Project

Manduca life cycle
Manduca sexta is commonly called a Sphinx moth or Hawk moth.
Like the other advanced insects, this moth has a life cycle
showing complete metamorphosis. Complete metamorphosis is
a pattern of growth and change characterized by four predictable,
very different life stages, the egg, larva, pupa, and adult. The
complete cycle usually takes from 30 to 50 days, depending on
environmental conditions.
W A larger, “poster” version of this
illustration is included in the back
of this guide, or can be downloaded
from The Manduca Project web site.
2 days http://insected.arizona.edu/manduca
Eggs 3t
1.4 mg o5
da
ys
s
Hiding out
y
da
3

In nature, Manduca pupae

2
to

Adult moth 1st instar larva bury themselves about

19

1 mg
9–15 inches deep

1st
2 to
underground, in loose soil

Stadiu
3 days
or leaf litter. This protects
them from predators and

m
Pupa
the weather.
5th Stadium
5 days

2nd instar larva

8 mg
What’s a Stadium?
2 to

2nd S

A stadium is the period of

3

tad

time between larval molts.

day
iu m

During a typical stadium,

s

5th instar larva Manducas spend about

1100 mg 48 hours eating and 30
s

ay hours actually molting.

ium

d 4d 3rd instar larva

a
St h 2t 30 mg
4t days
o3
3rd
Sta
dium Manduca’s favorite
4th instar larva “natural” foods
150 mg
• Peppers
• Tomarillo
The egg stage • Jimson Weed, Loco Weed
• Devil’s Claw
Manduca sexta lays its eggs on the underside
• Tomato
of the leaves of its favorite food plant. • Tobacco
The tiny, pale eggs are a little over • Petunia
1 millimeter in diameter and weigh • Tomatillo & Ground
about 1 milligram. A Manduca adult Cherry
lays eggs just once—but she typically • various ornamental
lays up to 200 of them! plants and vines
University of Arizona / ARLDN

The Life of Manduca sexta 19


X A larger, “poster” version of this
illustration is included in the back
of this guide, or can be downloaded
from The Manduca Project web site.
http://insected.arizona.edu/manduca
Manduca Mouth
Manduca larvae spend
most of their lives eating,
so their mouths are well-
adapted for chewing food.
Below, a manduca larva
holds a piece of Manduca
diet with its clawed “feet”
as it eats.
Courtesy of Chip Hedgecock, ARLDN
20 The Manduca Project
The larval stage
Larvae, also known as caterpillars, hatch from their tiny, pale
green eggs three to five days after the eggs are laid. These
“newborn” or first instar larvae are only a few millimeters long and
very pale green, but that changes quickly. As the larvae get older
and larger they become bright green with seven pairs of lateral
white stripes bordered with black. At the hind end of the body is
a horn or spinelike process that is usually red or reddish-brown.
This horn is characteristic of most species in this family of
moths, and serves as the basis of another common name: the
tobacco hornworm. It is not clear what purpose the horn serves,
although there is speculation that it may be useful in protective
camouflage or mimicry. One thing it does not do however, is house
a stinger, although it is a common myth that it does.
Thorax Abdomen
Head
Chewing Leg Prolegs Spiracle
mouthparts
The name Manduca means “glutton” or “chewer”. Looking at the
larvae, it’s easy to see where this name comes from. The larvae
have mouthparts adapted for chewing, and they put these to use
eating everything in sight. In the wild, they feed voraciously on
such plants as tobacco, tomato, potato, pepper, and nightshade
(Family: Solanaceae). Gardeners familiar with this insect know
that a mature (5th instar) larva can consume an entire tomato or
chili plant in a single night!
It’s important to note that the larvae don’t eat all the time. Right
before each larval molt, the larvae stop eating and void their gut
contents so that they’re empty for the actual molt. During this
phase, which can last 48 hours or more, the old hard head capsule
slips forward to make room for the developing new one. The larvae
appear bubble-headed at this point and are informally referred to
as head-caps. Eventually, the old skin (exoskeleton) is shed and is
usually eaten by the larva in a perfect example of recycling.
The molting process repeats itself four times, with each How many legs?
successive molt yielding a larger larva. At the end of the fifth According to the picture,
Manduca larvae appear to
and final larval instar, something different happens. The huge larva have eight pairs of legs.
climbs down from its host plant and begins a period of vigorous Aren’t insects supposed
walking or wandering that typically lasts for about five days. to have just three pairs?
During this stage, gut contents are again voided, body moisture The three pairs of legs on
is lost, and a dark bluish line or dorsal heart becomes visible along the thorax are the true
legs. These will become the
the larva’s back. At this point, the larva seeks out loose soil or leaf six legs of the adult moth.
litter and buries itself. After a few days, the buried larva molts a The larvae’s stubby
final time to form a light green pupa. prolegs, complete with
claw-like “crockets”, shown
below, are part of the
The pupal stage abdomen. They are used
for crawling, clinging, and
Insects’ pupal stage is frequently referred to as a “resting stage”. climbing, and are changed
into other structures
While it’s true that the insect is no longer walking and feeding, during metamorphosis.
amazing changes are taking place. Over the first few hours,
chemical reactions cause the pale green skin or cuticle of the
new pupa to harden and turn dark brown. During the three-week
pupation period, tissues are reorganized to form the structures
of the adult Manduca. Looking closely at the cuticle, you can
see the location of the developing wings, eyes, antennae, and
Courtesy of Chip Hedgecock, ARLDN
proboscis (tongue) of the adult moth. In a sense, the cuticle is
a mold into which the developing adult is formed. When the adult
emerges, the cuticle is split open and left behind.
Thorax Abdomen
Head W A larger, “poster” version of this
illustration is included in the back
of this guide, or can be downloaded
from The Manduca Project web site.
http://insected.arizona.edu/manduca
Spiracle
Compound eye Antenna Wing
Proboscis

The pupa loses the ability to walk, but it can still move. If the
pupa is disturbed, the end of the abdomen thrashes violently. This
response is defensive, and may startle a potential predator into
dropping the pupa and seeking a less lively meal. The thrashing is
also effective against small predators, and may decapitate ants
attacking by way of the spiracles.

The Life of Manduca sexta 21

 The adult stage
The pupa completes its development in 19 to 23 days and the
adult Manduca sexta emerges. The adult is a large, gray moth with
a wingspan of up to 10 centimeters (4 inches). It has dense
scaling, banded hindwings, and six orange-yellow spots along each
side of the abdomen (hence the scientific name “sexta”). The
proboscis, which first appeared as a conspicuous “jug-like handle”
in the pupa, becomes a tightly-coiled siphon in the adult. Unlike
the chewing mouthparts of the larvae, this siphon can be uncoiled
and used like a soda straw to sip nectar, the adults’ only food.
Adults are strong flyers, active at dawn and dusk, and may often
be seen hovering hummingbird-like from flower to flower. Within
two days after emerging from the pupa, adults begin mating. The
females lay eggs once—up to 200 of them—on the kinds of host
plants their larvae prefer to eat. After an average life
span of about a week, the adults die, leaving the Wing
garden to the next generation of hungry
caterpillars. Antenna
Head
X A larger, “poster” version of this Proboscis
illustration is included in the back
of this guide, or can be downloaded
from The Manduca Project web site. Compound Eye Abdomen
http://insected.arizona.edu/manduca
Thorax
Leg

22 The Manduca Project