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1 Introduction to the Arthropods

What Is an Arthropod?
The subjects of this book are the arthropods that live among us, primarily the
insects but also some of their relatives, such as arachnids, millipedes, centipedes,
and a few crustaceans. When formally classified, these animals are placed in the
phylum Arthropoda, which comprises a huge number of species with a tremendous
diversity of forms and habits. Nonetheless, all arthropods share certain features
that together define them as a distinct form of life:

(a) (b)

(c)
F I G U R E  1-1
Three representative arthropods. (a) Dragonfly (insect), (b) julid millipede, and (c) windscorpion
(arachnid). All show the basic external features of arthropods including an exoskeleton, segmenta-
tion of the body, jointed appendages, and a body design that is bilaterally symmetrical. Photograph
of the dragonfly courtesy of Brian Valentine; photograph of the millipede courtesy of Jim Kalisch/
University of Nebraska; photograph of the windscorpion by Jack Kelly Clark and provided courtesy
of the University of California IPM Program.

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2    CHAPT ER 1

• All arthropods have a body supported by a

hardened external skeleton (­ exoskeleton), a reverse The Diversity and
type of engineering compared to our internal
skeleton. To allow growth, this exoskeleton must be Abundance
periodically shed, and a new one rebuilt.
• The body of an arthropod is divided into segments,
of Arthropods
a feature shared by some other animal groups, The arthropods are, by far, the most diverse life form
such as earthworms (phylum Annelida) and on the planet. Insects alone, with approximately
velvet worms (phylum Onychophora). 970,000 known species, comprise over one-half of all
• The appendages of arthropods—their legs, kinds of life known to occur on the planet. Yet despite
antennae, and mouthparts—are jointed. This is the impressive numbers, these reflect only “known
the feature that defines the phylum. (In Greek, species,” ones that have been suitably described in the
arthropod means “jointed foot.”) scientific literature and accepted as distinct species.
• Internally, the nerve cord runs along the lower This number represents only a small fraction of the
(ventral) part of the body and is not enclosed in number of species estimated to be present on the
a protective spinal column. These features planet today. This number is also a tiny fraction of all
contrast with those found in phylum Chordata the insects that ever were on the planet. It has been
to which we belong. suggested that perhaps 95% of all insects that have
• Blood is moved by the aid of a tube-like heart, ever existed, since their first appearance some 400
located along the back (dorsal) part of the million years ago (mya), are now extinct.
body. Today, the number of insect species thought to
• The overall body arrangement is bilaterally occur is often estimated at about four to five million
symmetrical, so that, if the body were cut species. The great majority of these, at least 80%,
through the center from head to tail, the two remain unknown to science so far. Progress is being
halves would be a mirror image of one another. made to close this gap, with over 7,000 new insect

Other life forms

(Prokaryota, Cyanophyta,
Chromista, Viruses,
Fungi and Protoclista)
Lichens

Insects

Plants
(excluding Algae)

Vertebrates
F I G U R E  1-2
The relative number of different kinds of life forms
known on Earth, based on the number of known
species. Of the approximately 1.9 million presently
recognized species, just over half are insects. Figures Other
based on Numbers of Living Species in Australia and invertebrates
the World, 2nd ed. (2009). Photographs courtesy of
Tom Murray. Non-insect arthropods

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I N T R O D U C T I O N T O T H E A RT H RO P O D S    3

species being described annually, over 20 per day on pasture during November 1943. About 2.5 billion
average. At this rate of new discovery, impressive as arthropods were estimated per hectare, with mites
it is, perhaps we can expect a full catalog of the five comprising some 62% and springtails 23% of the
million insects to be ready in about 550 years or so. total number. On the basis of surveys such as this it
A much more difficult question to answer is has been estimated that the insects, springtails, mites,
“How abundant are insects and other arthropods in and other land-dwelling ­arthropods outnumber
terms of total population numbers?” One of the humans by as much as 250 million to 1. Furthermore,
problems is that the overwhelming number of these arthropods collectively comprise over 80% of
arthropods are minute and live in soil. For example, the total biomass of the terrestrial animals, far
one of the first attempts at counting all of the outweighing all the other land dwellers such as
arthropods in a sample of soil was done in an English earthworms, reptiles, birds, and mammals.

(a) (b)
F I G U R E 1- 3
(a and b) Springtails and soil-dwelling mites are the most abundant kinds of animal life on the planet. A billion or more may typically
be found in a hectare of fields, pasture, or lawn. Photographs courtesy of Brian Valentine.

The Many Roles combined. Furthermore, the roles of arthropods in

ecosystems are myriad, but central to the functioning
of Arthropods of planet Earth:

If all mankind were to disappear, the world

would regenerate back to the rich state of Pollination of flowering plants. Insects are essential
­equilibrium that existed ten thousand years to the pollination of most flowering plants, and
ago. If insects were to vanish, the environment many of the flowering plants are the result of
would collapse into chaos. (E. O. Wilson, coevolution with their insect pollinators. The
The Diversity of Life) tremendous variety of flower types reflect
different ways that plants have evolved to more
Although small in size, arthropods, in their efficiently attract pollinators. In response, new
tremendous numbers, collectively account for the species of insects have arisen to better exploit
most biomass of all land animals. In the Amazon rain these sources of nectar and pollen. In addition to
forest, the weight of just one family of insects, the native plants, essentially all fruits, vegetables, and
ants, is estimated to be four times more than all the many of the forage crops (e.g., clover, alfalfa) are
mammals, birds, fish, reptiles, and amphibians dependent on insects to produce seed.

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4    CHAPT ER 1

F I G U R E 1- 4
A leafcutter bee pollinating sweet pea. Many plants are F I G U R E   1- 5
dependent on insects for their pollination. Photograph by Blow flies colonizing fresh carrion. Insects help in the
Whitney Cranshaw/Colorado State University. ­decomposition of dead plant and animal matter and have central
roles in the cycling of nutrients in natural systems. Photograph
by Whitney Cranshaw/Colorado State University.
Recycling plant and animal matter. Many insects
develop by feeding on dead plant matter, dead
animal matter, or animal dung. In this role, they
function as macrodecomposers that are in the
first-line “clean-up crew” essential to the recovery
and recycling of nutrients. Through insect feeding,
these substances are broken down into much
smaller particles and partially digested, which
greatly accelerates the process of decay that frees
the nutrients to nourish later generations of plants.
In the absence of insects, nutrient-recycling systems
break down and organic matter accumulates.
Soil formation and mixing. The great majority of
terrestrial arthropods live within the soil. These
animals help to turn the soil and incorporate organic
matter and nutrients. The impacts of these activities
can be very dramatic, with some of the social insects
(e.g., the ants and termites) moving and mixing
tremendous amounts of soil as they tunnel. These
processes are critical to soil formation and the
maintenance of soil fertility. Without these insects,
plant growth would be reduced and restricted.
Centrality to animal food chains. Through their
feeding activities, plant-feeding insects (about
25% of the species on the planet) convert plant
biomass to animal biomass. In turn, these
F I G U R E   1- 6
creatures serve as the primary source of food for Mound-building termites are central to soil formation and
other insects (another 25% of the planet’s mixing in large areas of Africa. Photograph courtesy of USDA
species) and for many birds, fish, and mammals APHIS PPQ Archives/Bugwood.org.

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I N T R O D U C T I O N T O T H E A RT H RO P O D S    5

that are, in turn, food for yet still more animals. Unfortunately, most people recognize only those
Thus, plant-eating insects are the critical link arthropods that are directly and immediately affecting
between plants and much of the rest of animal human activities. These species are often considered
life on Earth (including humans). negatively, as competitors, because of their ability to
cause several types of harm—destruction of crops,
damage to stored products or structures, transmission
of plant and animal pathogens, and stings or bites—
or merely some degree of annoyance. Those that do
affect us in these ways are judged to be “pests,” a
subjective and very flexible term that is defined by
how much impact they are personally perceived to
have. It is important to keep in mind that only a tiny
fraction of all arthropods are ever elevated to this
infamous status. A listing of all insects worldwide that
are considered pests for one reason or another would
include fewer than 10,000 species, approximately 1%
of the total number of known insect species. A list of
species that are directly beneficial to humans may be
larger by an order of magnitude.
All too often people try to separate the insect
world into “good bugs” and “bad bugs.”
Alternatively, one often hears the question “What
good is this insect/scorpion/spider?” These types of
categorizations fail to recognize the tremendous
F I G U R E 1-7
Many types of wildlife utilize insects as an important part of
importance of the arthropods to the functioning of
their diet. Photograph courtesy of David Leatherman. this planet, usually in ways we little understand. It is
perhaps important to keep in mind the words of
Maintenance of plant communities. Although the effects pioneer conservationist/naturalist John Muir: “When
of large plant-feeding mammals are conspicuous, it is we try to pick out anything by itself, we find it
the activities of insects that most often determine hitched to everything else in the Universe.”
what plant life is present. Insects do this in many Insects are neither good nor bad. They are, along
ways, including feeding on plants (phytophagy), with all other extant life forms, a representation of
feeding on seeds, pollination, and dispersing seeds. the latest expression of what has evolved on Earth.

Classification of the
Arthropods
In the classification of biological organisms, all life
forms are grouped according to how related they are,
usually based on physical features. Within this
organization, all life forms are arranged in a series of
subgroupings that become increasingly specialized.
This science of classification is known as taxonomy
and is conducted by specialists known as taxonomists.
Closely associated with taxonomy, and often guiding
the classification arrangements, is the science of
F I G U R E 1- 8
Through their foraging activities, leafcutting ants can have
systematics that seeks to determine the relatedness of
dramatic effects on the kinds of plant life that occur. Photograph different life forms. Systematists make extensive use
courtesy of Ronald F. Billings/Texas Forest Service/Bugwood.org. of the fossil records of extinct species along with all

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6    CHAPT ER 1

PROTURA

COLLEMBOLA
springtails

DIPLURA

MICROCORYPHIA
bristletails

THYSANURA
silverfish
EPHEMEROPTERA
mayflies
ODONATA
dragon-/damselflies
ORTHOPTERA
crickets, grasshoppers
PHASMATODEA
stick insects
DERMAPTERA
earwigs
GRYLLOBLATTODEA
? ice crawlers
MANTOPHASMATODEA
rock crawlers

PLECOPTERA
stoneflies
EMBIIDINA
web spinners
ZORAPTERA

BLATTODEA
cockroaches
ISOPTERA
termites
MANTODEA
mantises
PHTHIRAPTERA
true lice
PSOCOPTERA
bark lice
HEMIPTERA
plant lice, true bugs
THYSANOPTERA
thrips
COLEOPTERA
beetles

NEUROPTERA
lacewings, dobsonfiles, snakeflies

HYMENOPTERA
wasps
LEPIDOPTERA
moths, butterflies
TRICHOPTREA
caddisflies
SIPHONAPTERA
fleas
MECOPTERA
scorpionflies

DIPTERA
true flies
? STREPSIPTERA
twisted wings

F I G U R E 1- 9
A diagram of a proposed phylogenetic relationship of the insect orders. Phylogenetics is the study of the evolutionary relations between
organisms, and in a diagram such as this the orders that are most closely clumped are most closely related. Phylogenetics is a very active
field that has been greatly aided by genetics. With new information, evolutionary relationships and taxonomic groupings are constantly
being reevaluated, and changes in taxonomic arrangement are frequent. (Modified from Grimaldi and Engel, 2005.)

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I N T R O D U C T I O N T O T H E A RT H RO P O D S    7

manner of biological features of present (extant) book. Also discussed, in part, are a group of
species. Increasingly, genetics also guides systematics. arthropod classes collectively known as crustaceans
The powerful new tools that allow sequencing of (subphylum Crustacea).
genes are producing a revolution in the systematics The classes are subdivided into orders.
of insects (and many other organisms) during which For example, once you have identified something as
we are seeing many taxonomic arrangements being an insect (from the class Hexapoda), the next
modified and many new species being recognized. grouping is the order of insects where it has been
Using this system, all animals are classed placed. Butterflies and moths, insects that have
together within the broadest type of grouping, a characteristic scale-covered wings, are placed by
kingdom—specifically the kingdom Animalia. taxonomists in the order Lepidoptera. Beetles that
The kingdom containing all animals is next have a hardened front pair of wings are in the order
subdivided into various phyla (singular, phylum), Coleoptera, while the flies, with their unique single
one of which is Arthropoda—the arthropods that pair of wings, are classified in the order Diptera.
are the focus of this book. (Examples of some Because of differences in how scientists classify the
other animal phyla include Annelida, the insects, you may see some differences in the number
segmented worms; Nematoda, the round worms; of and names for the orders among the many books
Mollusca, the mollusks; and Chordata, the animals describing insect life and in their names. The
with a hollow, ventral nerve cord, which includes classification system used for this book follows that
humans.) In turn, a phylum is subdivided into of the 7th edition of Borror and DeLong’s
sections, each known as a class. Four classes of Introduction to the Study of Insects (2005), which
arthropods (millipedes, centipedes, arachnids, lists in the class Hexapoda 28 orders of insects and
hexapods/insects) are the primary focus of this 3 orders of entognathous hexapods.

(a) (b)

F I G U R E   1-10
(a–c) Representatives of three insect orders: sulfur butterfly (Lepidoptera), blatellid
cockroach (Blattodea), and ground beetle (Coleoptera). Photographs courtesy of
David Cappaert/Michigan State University/Bugwood.org, Ken Gray/Oregon State
(c) University, and Jim Kalisch/University of Nebraska, respectively.

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8    CHAPT ER 1

Orders are subdivided into families, scientific weevils and bark beetles (Curculionidae), and leaf
names usually ending in “idae.” For example, the beetles (Chrysomelidae). Each family is divided
beetles (order Coleoptera) are divided into scores into genera (singular genus), and each genus into
of families, including lady beetles (Coccinellidae), various species.

(a) (b) (c)

F I G U R E 1-11
(a–c) Representatives of three families within the order Coleoptera (beetles): lady beetle (Coccinellidae), weevil (Curculionidae), and
darkling beetle (Tenebrionidae). Photographs courtesy of Whitney Cranshaw/Colorado State University, Brian Valentine, and Jim
Kalisch/University of Nebraska, respectively.

Each species of insect, as well as all other life giving each species a scientific name that is universally
forms, has its own scientific name. This name is recognized was formalized by Carolus Linnaeus
constructed by combining the genus name and what is (sometimes Anglicized as Carl Linnaeus), a Swedish
known as the specific epithet. The genus name is physician and biologist (1707–1778). The outline he
capitalized, the specific epithet is not, and both are developed, published in the book Systema Naturae
written in italics. For example, the scientific name of (1st edition 1735), was revolutionary and remains the
the house fly is Musca domestica and that of the tomato fundamental framework whereby all living organisms
hornworm is Manduca quinquemaculata. The idea of are classified, based on shared features.

(a) (b) (c)

F I G U R E 1-12
(a–c) Representatives of three different species within the beetle family Coccinellidae (lady beetles): Hippodamia parenthesis,
Harmonia axyridis, Coleomegilla maculata. Photographs by Whitney Cranshaw/Colorado State University.

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I N T R O D U C T I O N T O T H E A RT H RO P O D S    9

Since each scientific name has two parts, it is

described as binomial nomenclature. This has become
the universally recognized standard for discussing the
identity of different organisms in a world that shares few
other common languages. In the formal naming of an
organism, the person who originally described it is also
placed at the end of the name. Therefore in the scientific
literature the house fly would be Musca domestica
Linnaeus and the tomato hornworm Manduca
quinquemaculata (Haworth), recognizing that these two
insects were originally described by Linnaeus and
Haworth, respectively. In this book, the descriptor names
are left out for simplification, not to diminish in any way
the contributions of those who took it upon themselves
to first identify the insect as being a unique species.
Several mnemonic phrases have been developed
to help reinforce learning of the basic taxonomic
groups—kingdom, phylum, class, order, family,
genus, species—including the following:

King Philip cuts open five green snakes. F I G U R E   1-13

The principles that guide the classification of living
Kings play cards on fat green stools. ­o rganisms was first formalized in the book Systema Naturae,
Kings play chess on Fridays, generally speaking. written by Carolus Linnaeus (1707–1778). Painting by
King Philip cried out—“for goodness sake!” Alexander Roslin.

TABLE 1-1   Taxonomic position of the honey bee, Apis mellifera Linnaeus, and the southern black widow, Latrodectus mactans (Fabricius).‡

HONEY BEE (COMMON NAME)* SOUTHERN BLACK WIDOW (COMMON NAME)†

F I G U R E 1-14 F I G U R E   1-15
The honey bee, Apis mellifera Linnaeus. Photograph courtesy of The southern black widow, Latrodectus mactans (Fabricius).
Joseph Berger/Bugwood.org. Photograph courtesy of Clemson University/Bugwood.org.

Phylum—Arthropoda Phylum—Arthropoda

Class—Hexapoda Class—Arachnida
(continued)

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10    CHAPT ER 1

TAB LE 1 - 1

HONEY BEE (COMMON NAME)* SOUTHERN BLACK WIDOW (COMMON NAME)†

Order—Hymenoptera Order—Araneae

Family—Apidae Family—Theridiidae

Genus—Apis Genus—Latrodectus

Species—mellifera Species—mactans

Original descriptor—Linnaeus Original descriptor—Fabricius ‡

*Common name accepted by the Entomological Society of America.

†
Common name accepted by the American Arachnological Society.
‡
The original 1775 description by Fabricius used the genus name Aranea. In later revisions, the southern black widow was placed in a different genus (Latrodectus).
This change from the original is indicated by the parentheses surrounding the name of the original descriptor.

It must be recognized that whatever type of As our understanding of how different organisms
classification is used, it is a human construct and thus are related has increased, additional groupings have been
subject to change. Orders, families, and even classes required. These are most often created by the prefix
of organisms may be rearranged following revisions “sub” or “super.” For example, a subclass is a division
made by taxonomists as new information becomes of a class but will still contain within it one or more
available through discoveries of new species, better orders of the class. A superfamily will contain one or more
fossil records, and the use of modern molecular families within the same order. The taxonomic
genetic techniques. arrangement used for this book is presented in table 1-2.

TAB LE  1 - 2   Primary taxonomic divisions of the phylum Arthropoda. Orders have been included for the terrestrial or freshwater
­arthropods that are the focus of this book (classes Arachnida and Hexapoda; subphylum Crustacea in brief).

Subphylum Trilobita—trilobites (fossils only)

Subphylum Chelicerata
Class Merostomata—eurypterids (fossils only) and horseshoe crabs
Class Arachnida—arachnids
Order Scorpiones—scorpions
Order Palpigradi—micro whipscorpions
Order Thelyphonida (Uropygi)—whipscorpions
Order Schizomida—shorttailed whipscorpions
Order Amblypygi—tailless whipscorpions, whipspiders
Order Araneae—spiders
Order Ricinulei—hooded tickspiders
Order Opiliones—harvestmen, daddy longlegs
Order Acari—mites and ticks
Order Pseudoscorpiones—pseudoscorpions
Order Solifugae—windscorpions, sunspiders
Class Pycnogonida—sea spiders
Subphylum Crustacea—crustaceans
Class Cephalocarida
(continued)

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I N T R O D U C T I O N T O T H E A RT H R O P O D S    11

TAB LE 1 -2

Class Branchiopoda
Order Anostraca—fairy shrimp
Order Notostraca—tadpole shrimp
Order Conchostraca—clam shrimp
Order Cladocera—water fleas
Class Ostracoda
Class Copepoda
Class Mystacocarida
Class Remipedia
Class Tantulocarida
Class Branchiura
Class Cirripedia
Class Malacostraca
Order Amphipoda—amphipods
Order Isopoda—isopods
Order Stomatopoda—mantis shrimp
Order Decapoda—lobsters, crayfish, crabs, shrimp
Subphylum Atelocerata
Class Diplopoda*—millipedes
Class Chilopoda*—centipedes
Class Pauropoda*—pauropods
Class Symphyla*—symphylans
Class Hexapoda—hexapods (includes insects)
Subclass Entognatha †
Order Protura—proturans
Order Diplura—diplurans
Order Collembola—springtails
Subclass Insecta—insects
Order Microcoryphia ‡—jumping bristletails
Order Thysanura ‡—silverfish
Order Ephemeroptera—mayflies
Order Odonata—dragonflies and damselflies
Order Orthoptera—grasshoppers, crickets, katydids
Order Phasmatodea—walkingsticks and leaf insects
Order Grylloblattodea—rock crawlers
Order Mantophasmatodea—heelwalkers or gladiators
Order Dermaptera—earwigs
Order Plecoptera—stoneflies
Order Embidiina—webspinners
Order Zoraptera—zorapterans, angel insects
Order Isoptera—termites
Order Mantodea—mantids
Order Blattodea—cockroaches
Order Hemiptera—true bugs, cicadas, hoppers, psyllids, whiteflies, aphids, and scale insects
Order Thysanoptera—thrips
Order Psocoptera—psocids
Order Phthiraptera—lice
(continued)

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12    CHAPT ER 1

TAB LE 1 - 2

Order Coleoptera—beetles
Order Neuroptera—alderflies, dobsonflies, fishflies, snakeflies, lacewings, antlions, and owlflies
Order Hymenoptera—sawflies, parasitic wasps, ants, wasps, and bees
Order Trichoptera—caddisflies
Order Lepidoptera—butterflies and moths
Order Siphonaptera—fleas
Order Mecoptera—scorpionflies
Order Strepsiptera—twisted-wing parasites
Order Diptera—flies

*
Arthropods that are often referred to as Myriapoda, the myriapods.
†
The classification of the various entognathous hexapods is subject to debate, although each of the three groups is considered distinct. Some classification schemes
consider them as separate subclasses or even classes.
‡
The orders Microcoryphia and Thysanura (alternately named as Archaeognatha and Zygentoma) consist of insects with primitive features that originated before
the development of wings. As such they are sometimes considered together as the Apterygota, in contrast with the other insect orders (Pterygota) that have
physical features associated with wings. This arrangement is subject to debate, as many other features of the Thysanura indicate that they are more closely related
to the insects that developed wings than to the Microcoryphia.

Common Name or United States. However, when outbreaks of the forest

tent caterpillar (Malacosoma disstria) occur in forests
Scientific Name? of northern Minnesota and Wisconsin and the
caterpillars are seen marching across roads, this very
Some insect orders and families, and many individual different insect is called an “armyworm” and elsewhere
species of insects, have a common name. This is the other caterpillars seen in bands are often referred as
familiar insect name in English, in contrast to the “armyworms.” Similarly, an odd group of insects known
more formal scientific name. For example “beetles” as Jerusalem crickets are known locally by a wide variety
is the common name for the order Coleoptera, of names such as “children of the earth,” “old baldheaded
“swallowtails” is the common name for the butterflies man,” and “potato bugs.” (In turn, a great number of
within the family Papilionidae, and “house fly” is the other generally round-bodied arthropods are known as
common name for the insect Musca domestica. “potato bugs,” including pillbugs and the Colorado
Scientific names are universal; they are the same in potato beetle.) Therefore the use of formally accepted
every country. That is their utility and appeal—although common names provides a means to discuss and write
names of genera and even families are sometimes about insects in a manner that allows the identification
rearranged when new information (now usually of the species to be consistently recognized.
genetics) leads to taxonomic revisions. However, most The Entomological Society of America attempts to
people find it easier to learn and use the common standardize the common names of insects used in the
names when discussing local insects. Unfortunately, United States in the publication Common Names of
such common names may be used for very different Insects and Related Organisms. Common names of
insects in different locations, thus leading to some arachnids are similarly codified by the American
confusion. For example, an insect formally known as Arachnological Society. Around the world, similar
the armyworm (Mythimna unipuncta) is a common publications have been developed by various professional
pest caterpillar of grain crops in much of the central organizations committed to the study of arthropods.

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I N T R O D U C T I O N T O T H E A RT H R O P O D S    13

(a)

F I G U R E   1-16
(a) The armyworm, Mythima unipuncta, and (b) the forest tent caterpillar,
Malacosoma disstria, each have formalized common names through the
Entomological Society of America. The armyworm is a pest of grain crops,
and the forest tent caterpillar feeds on various deciduous trees. During
outbreaks, forest tent caterpillars are sometimes referred to as “armyworms,”
which can cause confusion as to the species in question. Photographs by
(b) Frank Peairs and Whitney Cranshaw/Colorado State University.

(a) (b) (c)

F I G U R E 1-17
Jerusalem crickets (a), Stenopelmatus spp., may locally be called by many different names including “children of the earth,” “old
baldheaded man,” and “potato bug.” Among the other arthropods that are sometimes called “potato bugs” are (b) pillbugs,
Armadillidium vulgare, and (c) the Colorado potato beetle, Leptinotarsa decemlineata. Photographs by Ken Gray/Oregon State
University, Whitney Cranshaw/Colorado State University, and David Cappaert/Michigan State University/Bugwood.org, respectively.

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