Hickman−Roberts−Larson: 12.

Arthropods Text © The McGraw−Hill

Animal Diversity, Third Companies, 2002
Edition

FIRST PAGES
12

chapter
• • • • • •

t w e l v e
Arthropods

A Winning Combination
Tunis, Algeria—Treating it as an invading army, Tunisia, Alge-
ria, and Morocco have mobilized to fight the most serious
infestation of locusts in over 30 years. Billions of the insects
have already caused extensive damage to crops and are threat-
ening to inflict great harm to the delicate economies of North
Africa.
Source: New York Times, April 20, 1988

Humans suffer staggering economic losses due to insects, of

which outbreaks of billions of locusts in Africa are only one
example. In the western United States and Canada, an outbreak of
mountain pine beetles in the 1980s and 1990s killed pines on huge
acreages, and the 1973 to 1985 outbreak of spruce budworm in
fir/spruce forests killed millions of conifer trees. These examples
serve to remind us of our ceaseless struggle with the dominant
group of animals on earth today: insects. Insects far outnumber all
other species of animals in the world combined, and numbers of
individuals are equally enormous. Some scientists have estimated
that there are 200 million insects for every human alive today!
Insects have an unmatched ability to adapt to all land environments
and to virtually all climates. Having originally evolved as land
animals, insects developed wings and invaded the air 150 million
years before flying reptiles, birds, or mammals. Many have exploited
freshwater and saltwater (shoreline) habitats, where they are now
widely prevalent; only in the seas are insects almost nonexistent, but
there are vast numbers of crustaceans in marine habitats.
How can we account for the enormous success of these crea-
tures? Arthropods have a combination of valuable structural and
physiological adaptations, including a versatile exoskeleton,
metamerism, an efficient respiratory system, and highly developed
sensory organs. In addition, many have a waterproofed cuticle and
have extraordinary abilities to survive adverse environmental condi-
tions. We describe these adaptations and others in this chapter.

Spotted spiny lobster,

Panulirus guttatus.
212
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

Arthropods 213

hylum Arthropoda (ar-throp´o-da) (Gr. arthron, joint, +

P pous, podos, foot) embraces the largest assemblage of

living animals on earth. It includes spiders, scorpions,
ticks, mites, crustaceans, millipedes, centipedes, insects, and
some smaller groups. In addition there is a rich fossil record
extending back to the mid-Cambrian period (figure 12.1).
Arthropods are eucoelomate protostomes with well-
developed organ systems, and their cuticular exoskeleton con-
taining chitin is a prominent characteristic. Like annelids, they
are conspicuously segmented; their primitive body pattern is a
linear series of similar somites, each with a pair of jointed
appendages. However, unlike annelids, arthropods have
embellished the segmentation theme: variation occurs in the
pattern of somites and appendages in the phylum. Often
somites are combined or fused into functional groups, called
tagmata, for specialized purposes. Appendages, too, are fre-
quently differentiated and specialized for walking, swimming,
A
flying, or eating.
Few arthropods exceed 60 cm in length, and most are far
below this size. The largest is a Japanese crab (Macrocheira
kaempferi), which has approximately a 3.7 m span; the small-
est is a parasitic mite, which is less than 0.1 mm long.
Arthropods are usually active, energetic animals. How-
ever we judge them, whether by their great diversity or their
wide ecological distribution or their vast numbers of species,
the answer is the same: they are the most abundant and
diverse of all animals.
Although arthropods compete with us for food supplies
and spread serious diseases, they are essential in pollination of
many food plants, and they also serve as food, yield drugs and
dyes, and create such products as silk, honey, and beeswax.

Ecological Relationships
Arthropods are found in all types of environment from low
ocean depths to very high altitudes and from the tropics far into
both north and south polar regions. Some species are adapted
for life on land or in fresh, brackish, and marine waters; others B
live in or on plants and other animals. Most species use flight to
varying degrees to move among their favored habitats.Some live f i g u r e 12.1
in places where no other animal could survive. Fossils of early arthropods. A, Trilobite fossils, dorsal view. These
Although all types—carnivorous, omnivorous, and her- animals were abundant in the mid-Cambrian period. B, Eurypterid fossil;
bivorous—occur in this vast group, the majority are herbivo- eurypterids flourished in Europe and North America from Ordovician
rous. Most aquatic arthropods depend on algae for their to Permian periods.
nourishment, and most land forms live chiefly on plants. There
are many parasites.In diversity of ecological distribution arthro-
pods have no rivals. 1. A versatile exoskeleton. Arthropods possess an
exoskeleton that is highly protective without sacrificing
Why Have Arthropods Achieved Such mobility. The skeleton is the cuticle, an outer covering
secreted by underlying epidermis.
Great Diversity and Abundance? The cuticle consists of an inner and thicker
Arthropods have achieved a great diversity, number of species, procuticle and an outer, relatively thin epicuticle.
wide distribution, variety of habitats and feeding habits, and The procuticle is divided into exocuticle, which is
power of adaptation to changing conditions. These are some secreted before a molt, and endocuticle, which is
of the structural and physiological patterns that have been secreted after molting. Both layers of the procuticle
helpful to them: contain chitin bound with protein. Chitin is a tough,
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

214 chapter twelve

and insects the cuticle forms ingrowths for muscle

characteristics attachment. It may also line the foregut and hindgut,
line and support the trachea, and be adapted for a
of phylum arthropoda variety of purposes.
1. Bilateral symmetry; metameric body, tagmata of The nonexpansible cuticular exoskeleton does,
head and trunk; head, thorax, and abdomen; or however, impose important conditions on growth. To
cephalothorax and abdomen grow, an arthropod must shed its outer covering at inter-
2. Appendages jointed; primitively, one pair to each vals and grow a larger one—a process called ecdysis,
somite (metamere), but number often reduced; or molting. Arthropods molt from four to seven times
appendages often modified for specialized functions before reaching adulthood, and some continue to molt
3. Exoskeleton of cuticle containing protein, lipid, after that. Much of an arthropod’s physiology centers on
chitin, and often calcium carbonate secreted by molting, particularly in young animals—preparation,
underlying epidermis and shed (molted) at intervals molting itself, and then all processes that must be com-
4. Muscular system complex, with exoskeleton for pleted in the postmolt period.
attachment; striated muscles for rapid action; An exoskeleton is also relatively heavy and becomes
smooth muscles for visceral organs; no cilia proportionately heavier with increasing size. Weight of
5. Coelom reduced; most of body cavity consisting of the exoskeleton tends to limit ultimate body size.
hemocoel (sinuses, or spaces, in the tissues) filled 2. Segmentation and appendages for more efficient
with blood locomotion. Typically each somite has a pair of jointed
6. Complete digestive system; mouthparts modified appendages, but this arrangement is often modified,
from appendages and adapted for different methods with both segments and appendages specialized for
of feeding adaptive functions. Limb segments are essentially
7. Circulatory system open, with dorsal contractile hollow levers that are moved by muscles, most of which
heart, arteries, and hemocoel are striated for rapid action. The jointed appendages are
8. Respiration by body surface, gills, tracheae (air equipped with sensory hairs and are variously modified
tubes), or book lungs for sensory functions, food handling, and swift and effi-
9. Paired excretory glands called coxal, antennal, or cient walking or swimming.
maxillary glands present in some; some with other 3. Air piped directly to cells. Most land arthropods have
excretory organs, called Malpighian tubules a highly efficient tracheal system of air tubes, which
10. Nervous system similar to annelid plan, with dorsal delivers oxygen directly to tissues and cells and makes a
brain connected by a ring around the gullet to a high metabolic rate possible. Aquatic arthropods
double nerve chain of ventral ganglia; fusion of gan- breathe mainly by some form of gill.
glia in some species; well-developed sensory organs 4. Highly developed sensory organs. Sensory organs
11. Sexes usually separate, with paired reproductive are found in great variety, from compound (mosaic) eyes
organs and ducts; usually internal fertilization; to senses of touch, smell, hearing, balancing, and chemi-
oviparous or ovoviviparous; often with meta- cal reception. Arthropods are keenly alert to what goes
morphosis; parthenogenesis in a few forms; on in their environment.
growth with ecdysis 5. Complex behavior patterns. Arthropods exceed most
other invertebrates in complexity and organization of
their activities. Innate (unlearned) behavior unquestion-
ably controls much of what they do, but learning also
resistant, nitrogenous polysaccharide that is insoluble in
plays an important part in the lives of many arthropods.
water, alkalis, and weak acids. Thus the procuticle not
6. Reduced competition through metamorphosis.
only is flexible and lightweight but also affords protec-
Many arthropods pass through metamorphic changes,
tion, particularly against dehydration. In most
including a larval form quite different from adults in struc-
crustaceans, the procuticle in some areas is also impreg-
ture. Larval forms are often adapted for eating a different
nated with calcium salts, which reduce its flexibility.
kind of food from that of adults and occupy a different
In the hard shells of lobsters and crabs, for instance,
space, resulting in less competition within a species.
this calcification is extreme. The outer epicuticle is
composed of protein and often lipids. The protein is
stabilized and hardened by a chemical process called
tanning, adding further protection. Both the procuticle Subphylum Trilobita
and the epicuticle are composed of several layers each.
The cuticle may be soft and permeable or may Trilobites (figure 12.1A) probably had their beginnings a mil-
form a veritable coat of armor. Between body segments lion or more years before the Cambrian period in which they
and between segments of appendages it is thin and flex- flourished. They have been extinct some 200 million years, but
ible, permitting free movement of joints. In crustaceans were abundant during the Cambrian and Ordovician periods.
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

Arthropods 215

Their name refers to the trilobed shape of the body, caused by abdominal appendages book gills (flat leaflike gills) are
a pair of longitudinal grooves. They were bottom dwellers, exposed. Horseshoe crabs can swim awkwardly by means of
probably scavengers. Most of them could roll up like pill bugs. their abdominal plates and can walk on their walking legs.
They feed at night on worms and small molluscs and are harm-
less to humans.
Subphylum Chelicerata
Chelicerate arthropods are a very ancient group that includes Class Pycnogonida: Sea Spiders
eurypterids (extinct), horseshoe crabs, spiders, ticks and mites,
scorpions, sea spiders, and others. They are characterized by Pycnogonids are curious little marine animals that are much
having six pairs of appendages that include a pair of che- more common than most of us realize. They stalk about on
licerae, a pair of pedipalps, and four pairs of walking legs their four pairs of long, thin walking legs, sucking juices from
(a pair of chelicerae and five pairs of walking legs in horseshoe hydroids and soft-bodied animals with their large suctorial
crabs). They have no mandibles and no antennae. Most proboscis (figure 12.3). They often have a pair of ovigerous
chelicerates suck liquid food from their prey. legs (ovigers) with which males carry the egg masses. Their
odd appearance is enhanced by the much reduced abdomen
attached to an elongated cephalothorax. Most are only a few
Class Merostomata millimeters long, although some are much larger. They are
common in all oceans.
Subclass Eurypterida
Eurypterids, or giant water scorpions (figure 12.1B), lived 200
to 500 million years ago and some were perhaps the largest of
Class Arachnida
all arthropods, reaching a length of 3 m. They had some resem- Arachnids (Gr. arachnē, spider) are a numerous and diverse
blances to marine horseshoe crabs (figure 12.2) and to scorpi- group, with over 50,000 species described so far. They include
ons, their terrestrial counterparts. spiders, scorpions, pseudoscorpions, whip scorpions, ticks,
mites, harvestmen (daddy longlegs), and others. The arachnid
tagmata are a cephalothorax and an abdomen.
Subclass Xiphosurida: Horseshoe Crabs
Xiphosurids are an ancient marine group that dates from the
Cambrian period. There are only three genera (five species) liv-
Order Araneae: Spiders
ing today. Limulus (L. limus, sidelong, askew) (figure 12.2), Spiders are a large group of 35,000 recognized species, distrib-
which lives in shallow water along the North American uted all over the world. The cephalothorax and abdomen
Atlantic coast, goes back practically unchanged to the Triassic show no external segmentation, and the tagmata are joined by
period. Horseshoe crabs have an unsegmented, horseshoe- a narrow, waistlike pedicel (figure 12.4).
shaped carapace (hard dorsal shield) and a broad abdomen, All spiders are predaceous and feed largely on insects
which has a long spinelike telson, or tailpiece. On some (figure 12.5). Their chelicerae function as fangs and bear ducts

Pedipalp

First walking leg

Chelicera
Mouth
Simple eye Gnathobase
Hinge

Opisthosoma Chilarium
(abdomen)
Eye Gills
Genital
operculum
Telson Abdomen
Carapace
Anus
Gill opercula
A B Telson

f i g u r e 12.2
A, Dorsal view of horseshoe crab Limulus (class Merostomata). They grow to 0.5 m in length. B, Ventral view.
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

216 chapter twelve

Chelicera crush or chew up prey, aiding digestion by enzymes from their

Proboscis mouth. Many spiders provision their young with previously
captured prey.
Palp
Spiders breathe by book lungs or tracheae or both.
Book lungs, which are unique to spiders, consist of many par-
Oviger
allel air pockets extending into a blood-filled chamber (see fig-
ure 12.4). Air enters the chamber by a slit in the body wall.
Abdomen Tracheae are a system of air tubes that carry air directly to tis-
sues from openings called spiracles. Tracheae are similar to
those in insects (p. 236), but are much less extensive.
Spiders and insects have a unique excretory system of
Malpighian tubules (see figure 12.4), which work in conjunc-
f i g u r e 12.3 tion with specialized rectal glands. Potassium and other solutes
Pycnogonid, Nymphon sp. In this genus all anterior appendages and waste materials are secreted into the tubules, which drain
(chelicerae, palps, and ovigers) are present in both sexes, although the fluid, or “urine,” into the intestine. Rectal glands reabsorb
ovigers are often not present in females of other genera. most of the potassium and water, leaving behind such wastes as
uric acid. By this cycling of water and potassium, species living
in dry environments conserve body fluids, producing a nearly
from their poison glands, with which they effectively dispatch dry mixture of urine and feces. Many spiders also have coxal
their prey. Some spiders chase their prey, others ambush them, glands, which are modified nephridia that open at the coxa, or
and many trap them in a net of silk. After a spider seizes its base, of the first and third walking legs
prey with its chelicerae and injects venom, it liquefies tissues Spiders usually have eight simple eyes, each provided
with a digestive fluid and sucks up the resulting broth into the with a lens, optic rods, and a retina (see figure 12.4B). Chiefly
stomach. Spiders with teeth at the bases of their chelicerae they perceive moving objects, but some, such as those of the

Tagmata

Abdomen
Pedicel Eyes
Cephalothorax

Pedipalp
Walking
A legs B Chelicerae

Pumping Pericardial
Anterior sinus Ostium
stomach
Brain aorta
Eyes Digestive gland
Lateral blood
vessel

Poison
gland Malpighian tubule

Stercoral
pocket
Anus
Chelicera
Pharynx Coxal Ventral Book lung Spinnerets
Vagina Silk glands
Digestive gland sinus Ovary
ceca Subesophageal Seminal
receptacle
C ganglion

f i g u r e 12.4
A, External anatomy of a jumping spider. B, Anterior view of head. C, Internal anatomy of a spider.
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

Arthropods 217

f i g u r e 12.5
A, A camouflaged crab spider,
Misumenoides sp., awaits its insect
prey. Its coloration matches the
petals among which it lies, thus
deceiving insects that visit the flow-
ers in search of pollen or nectar.
B, A jumping spider, Eris aurantius.
This species has excellent vision
and stalks an insect until it is close
enough to leap with unerring preci-
sion, fixing its chelicerae into its prey.

A B

hunting and jumping spiders, may form images. Because vision tilized eggs in a silken cocoon, which she may carry about or
is usually poor,a spider’s awareness of its environment depends may attach to a web or plant. A cocoon may contain hundreds
especially on its hairlike sensory setae. Every seta on its sur- of eggs, which hatch in approximately two weeks.Young usu-
face is useful in communicating some information about its sur- ally remain in their egg sac for a few weeks and molt once
roundings, air currents, or changing tensions in the spider’s before leaving it. Several molts occur before adulthood.
web. By sensing vibrations of its web, a spider can judge the
size and activity of its entangled prey or can receive a message
Are Spiders Really Dangerous? It is truly amazing that
tapped out on a silk thread by a prospective mate.
such small and helpless creatures as spiders have generated so
much unreasoned fear in humans. Spiders are timid creatures,
Web-Spinning Habits The ability to spin silk is an impor- which, rather than being dangerous enemies to humans, are
tant factor in the lives of spiders, as it is in some other arach- actually allies in our continuing conflict with insects. The
nids. Two or three pairs of spinnerets containing hundreds of venom produced to kill prey is usually harmless to humans.
microscopic tubes connect to special abdominal silk glands Even the most poisonous spiders bite only when threatened or
(see figure 12.4C). A protein secretion emitted as a liquid hard- when defending their eggs or young. American tarantulas (fig-
ens on contact with air to form a silk thread. Spiders’ silk ure 12.8), despite their fearsome appearance, are not danger-
threads are stronger than steel threads of the same diameter ous. They rarely bite, and their bite is not considered serious.
and are said to be second in tensional strength only to fused
quartz fibers. The threads will stretch one-fifth of their length
before breaking.
The spider web used for trapping insects is familiar to
most people. Webs of some species consist merely of a few
strands of silk radiating out from a spider’s burrow or place of
retreat. Other species spin beautiful, geometric orb webs.
However, spiders use silk threads for many purposes besides
web making. They use silk threads to line their nests; form
sperm webs or egg sacs; build draglines; make bridge lines,
warning threads, molting threads, attachment discs, or nursery
webs; or to wrap up prey securely (figure 12.6). Not all spiders
spin webs for traps. Some, such as the wolf spiders, jumping
spiders (see figure 12.5B), and fisher spiders (figure 12.7), sim-
ply chase and catch their prey.

Reproduction Before mating, a male spins a small web,

deposits a drop of sperm on it, and then picks the package up
and stores it in special cavities of his pedipalps. When he f i g u r e 12.6
mates, he inserts his pedipalps into a female’s genital opening Grasshopper, snared and helpless in the web of a golden garden spider
to store the sperm in his mate’s seminal receptacles. A (Argiope aurantia), is wrapped in silk while still alive. If the spider is not
courtship ritual usually precedes mating. A female lays her fer- hungry, its prize will be saved for a later meal.
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
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Edition

218 chapter twelve

f i g u r e 12.7
A fisher spider, Dolomedes triton, feeds on a minnow. This handsome
spider feeds mostly on aquatic and terrestrial insects but occasionally
captures small fishes and tadpoles. It pulls its paralyzed victim from
the water, pumps in digestive enzymes, then sucks out the predigested
contents.

f i g u r e 12.9
A, A black widow spider, Latrodectus mactans, suspended on her web.
Note the orange “hourglass” on the ventral side of her abdomen.
B, The brown recluse spider, Loxosceles reclusa, is a small venomous
spider. Note the small violin-shaped marking on its cephalothorax.
The venom is hemolytic and dangerous.
f i g u r e 12.8
A tarantula, Brachypelma vagans.
Some spiders in other parts of the world are dangerous,
for example, funnel-web spiders Atrax robustus in Australia.
Two genera in the United States can give severe or even Most dangerous of all are certain ctenid spiders in South Amer-
fatal bites: Latrodectus (L.latro, robber,+ dektes, biter),and Lox- ica, for example, Phoneutria fera. In contrast to most spiders,
osceles (Gr. loxos, crooked, + skelos, leg). The most important these are quite aggressive.
species are Latrodectus mactans, or black widows, and Lox-
osceles reclusa, or brown recluse. Black widows are moderate
Order Scorpionida: Scorpions
to small in size and shiny black, with a bright orange or red
“hourglass” on the underside of their abdomen (figure 12.9). Although scorpions are more common in tropical and sub-
Their venom is neurotoxic;that is,it acts on the nervous system. tropical regions, some occur in temperate zones. Scorpions are
About four or five out of each 1000 bites reported are fatal. generally secretive, hiding in burrows or under objects by day
Brown recluse spiders, which are smaller than black wid- and feeding at night. They feed largely on insects and spiders,
ows, are brown, and bear a violin-shaped dorsal stripe on their which they seize with clawlike pedipalps and tear up with
cephalothorax (figure 12.9). Their venom is hemolytic rather jawlike chelicerae.
than neurotoxic, destroying tissues and skin surrounding a A scorpion’s body consists of a rather short cephalotho-
bite. Their bite can be mild to serious and occasionally fatal. rax, which bears appendages and from one to six pairs of eyes,
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
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Edition

Arthropods 219

f i g u r e 12.10
A, An emperor scorpion (order Scorpionida),
Paninus imperator, with young, which stay with
their mother until their first molt. B, A harvest-
man (order Opiliones). Harvestmen run rapidly
on their stiltlike legs. They are especially notice-
able during the harvesting season, hence the
common name.

A B

and a clearly segmented abdomen without appendages. The

abdomen is divided into a broader preabdomen and tail-like
postabdomen, which ends in a stinging apparatus used to
inject venom (figure 12.10). Venom of most species is not
harmful to humans, although that of certain species of Androc-
tonus in Africa and Centruroides in Mexico, Arizona, and New
Mexico can be fatal unless antivenom is available.
Scorpions bear living young, which their mother carries
on her back until after the first molt.

Order Opiliones: Harvestmen

Harvestmen, often known as “daddy longlegs,” are common in
the United States and other parts of the world (figure 12.10). f i g u r e 12.11
These curious creatures are easily distinguished from spiders A wood tick, Dermacentor variabilis (order Acarina).
by a broad joining of their abdomen and cephalothorax with-
out constriction of a pedicel, and by presence of external seg-
mentation of their abdomen. They have four pairs of long, have to be important ecologically, but many acarines have
spindly legs, and without apparent ill effect, they can cast off more direct effects on our food supply and health. Spider mites
one or more legs if they are grasped by a predator (or human (family Tetranychidae) are serious agricultural pests on fruit
hand). The ends of their chelicerae are pincerlike, and they trees, cotton, clover, and many other plants. Larvae of genus
feed much more as scavengers than do most arachnids. Trombicula are called chiggers or redbugs. They feed on der-
mal tissues of terrestrial vertebrates, including humans, and
cause an irritating dermatitis; some species of chiggers trans-
Order Acari: Ticks and Mites mit a disease called Asiatic scrub typhus. Hair-follicle mites,
Acarines differ from all other arachnids in having their Demodex (figure 12.13), are apparently nonpathogenic in
cephalothorax and abdomen completely fused, with no sign of humans; they infect most of us although we are unaware of
external division or segmentation (figure 12.11). Their mouth- them. Other species of Demodex and other genera of mites
parts are carried on a little anterior projection, or capitulum. cause mange in domestic animals.
They are found almost everywhere—in both fresh and salt
water, on vegetation, on the ground, and parasitic on verte-
brates and invertebrates. Over 25,000 species have been The inflamed welt and intense itching that follows a chigger
described, many of which are important to humans, but this is bite is not the result of a chigger burrowing into the skin,
probably only a fraction of the species that exist. as is popularly believed. Rather the chigger bites through
Many species of mites are entirely free living. Der- skin with its chelicerae and injects a salivary secretion con-
matophagoides farinae (Gr. dermatos, skin, + phago, to eat, + taining powerful enzymes that liquefy skin cells. Human
eidos, likeness of form) (figure 12.12) and related species are skin responds defensively by forming a hardened tube that
denizens of house dust all over the world, sometimes causing the larva uses as a sort of drinking straw and through
allergies and dermatoses. Some mites are marine, but most which it gorges itself with host cells and fluid. Scratching
aquatic species are found in fresh water. They have long, hair- usually removes the chigger but leaves the tube, which is a
like setae on their legs for swimming, and their larvae may be source of irritation for several days.
parasitic on aquatic invertebrates. Such abundant organisms
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

220 chapter twelve

spotted fever, a poorly named disease because most cases

occur in the eastern United States. Dermacentor also transmits
tularemia and agents of several other diseases. Texas cattle
fever, also called red-water fever, is caused by a protozoan par-
asite transmitted by the cattle tick Boophilus annulatus. Many
more examples could be cited.

In the 1970s people in the town of Lyme, Connecticut,

experienced an epidemic of arthritis. Subsequently known
as Lyme disease, it is caused by a bacterium and carried by
ticks of the genus Ixodes. Now thousands of cases are
reported each year in Europe and North America, and
other cases have been reported from Japan, Australia, and
South Africa. Many people bitten by infected ticks recover
spontaneously or do not suffer any ill effects. Others, if not
f i g u r e 12.12 treated at an early stage, develop a chronic, disabling
Scanning electron microgaph of house dust mite, Dermatophagoides farinae. disease. Lyme disease is now the leading arthropod-borne
disease in the United States.

Subphylum Crustacea
Crustaceans traditionally have been included as a class in sub-
phylum Mandibulata, along with insects and myriapods. Mem-
bers of all of these groups have, at least, a pair of antennae,
mandibles, and maxillae on the head. Whether Mandibulata
constitutes a monophyletic grouping has been debated, and
we discuss this question further on page 248.
The 30,000 or more species of Crustacea (L. crusta, shell)
include lobsters,crayfishes,shrimp,crabs,water fleas,copepods,
and barnacles. It is the only arthropod class that is primarily
aquatic; they are mainly marine, but many freshwater and a few
terrestrial species are known. The majority are free living, but
many are sessile, commensal, or parasitic. Crustaceans are often
f i g u r e 12.13 very important components of aquatic ecosystems, and several
Demodex follicuorum, human follicle mite. This tiny mite (100 to 400 µm) have considerable economic importance.
lives in follicles, particularly around the nose and eyes. Its prevalence Crustaceans are the only arthropods with two pairs of
ranges from about 20% in persons 20 years of age or younger to nearly antennae (figure 12.14). In addition to antennae and
100% in the aged. mandibles, they have two pairs of maxillae on the head,
followed by a pair of appendages on each body segment
(although appendages on some somites are absent in some
Ticks are usually larger than mites. They pierce the skin groups). All appendages, except perhaps the first antennae
of vertebrates and suck blood until enormously distended; (antennules), are primitively biramous (two main branches),
then they drop off and digest their meal. After molting, they and at least some appendages of all present-day adults show
are ready for another meal. In addition to disease conditions that condition. Organs specialized for respiration, if present,
that they themselves cause, ticks are among the world’s pre- are in the form of gills. Crustaceans lack Malpighian tubules.
mier disease vectors, ranking second only to mosquitos. They Crustaceans primitively have 60 segments or more, but
carry a greater variety of infectious agents than any other most tend to have between 16 and 20 somites and increased
arthropods; such agents include protozoan, rickettsial, viral, tagmatization. The major tagmata are head, thorax, and
bacterial, and fungal organisms. Species of Ixodes carry the abdomen, but these are not homologous throughout the sub-
most common arthropod-borne infection in the United States, phylum (or even within some classes) because of varying
Lyme disease (see accompanying note). Species of Dermacen- degrees of fusion of somites, for example, as in the cephalo-
tor (see figure 12.11) and other ticks transmit Rocky Mountain thorax.
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Arthropods 221

Antenna

Gill

Cephalothorax Abdomen
13 segments 6 segments
Rostrum
Carapace Coxa
Eye
Basis
Antennule Thorax
Exopod
Mandible Endopod
Maxillae

Maxillipeds

Cheliped Swimmerets Telson

(first leg)
Uropod

f i g u r e 12.15
Parts of a biramous crustacean appendage (third maxilliped
Walking legs of a crayfish).

f i g u r e 12.14
Archetypical plan of Malacostraca. Note that maxillae and maxillipeds
have been separated diagrammatically to illustrate general plan. Typically
in living animals only the third maxilliped is visible externally. In order ber of appendages and toward a variety of modifications that fit
Decapoda the carapace covers the cephalothorax, as shown here. them for many functions. Some are foliaceous (flat and leaflike),
as are the maxillae; some are biramous, as are the swimmerets,
maxillipeds, uropods, and antennae; some have lost one branch
and are uniramous, as are the walking legs.
In many crustaceans, the dorsal cuticle of the head In crayfishes we find the first three pairs of thoracic
extends posteriorly and around the sides of the animal to appendages, called maxillipeds, serving along with the two
cover or fuse with some or all thoracic and abdominal somites. pairs of maxillae as food handlers; the other five pairs of
This covering is called a carapace. In some groups the cara- appendages are lengthened and strengthened for walking
pace forms clamshell-like valves that cover most or all of the and defense (figure 12.16). The first pair of walking legs,
body. In decapods (including lobsters, shrimp, crabs, and oth- called chelipeds, are enlarged with a strong claw, or chela,
ers) the carapace covers the entire cephalothorax but not the for defense. Abdominal swimmerets serve not only for loco-
abdomen. motion, but in males the first pair is modified for copulation,
and in females they all serve as a nursery for attached eggs
and young. The last pair of appendages, or uropods, are
Form and Function wide and serve as paddles for swift backward movements,
and, with the telson, they form a protective device for eggs or
Appendages
young on the swimmerets.
Some modifications of crustacean appendages may be illus-
trated by those of crayfishes and lobsters (class Malacostraca,
order Decapoda, p. 228). Swimmerets, or abdominal Terminology applied by various workers to crustacean
appendages, retain the primitive biramous condition. Such an appendages has not been blessed with uniformity. At least
appendage consists of inner and outer branches, called the two systems are in wide use. Alternative terms to those we
endopod and exopod, which are attached to one or more use, for example, are protopodite, exopodite, endopodite,
basal segments collectively called a protopod (figure 12.15). basipodite, coxopodite, and epipodite. The first and second
There are many modifications of this plan. In the primitive pairs of antennae may be called antennules and antennae,
character state for crustaceans, all trunk appendages are rather and the first and second maxillae are often called maxillules
similar in structure and adapted for swimming.The evolutionary and maxillae. A rose by any other name . . .
trend, shown in crayfishes, has been toward reduction in num-
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222 chapter twelve

Antenna
Antenna

Cheliped Antennules Cheliped

(first walking leg)
Antennules
Rostrum

Eye Exopod of first maxilliped

Second maxilliped
Second
walking leg Mouth

Third Third maxilliped

walking leg
Second walking leg
Carapace Third walking leg

Cervical groove Fourth walking leg

First (copulatory)
Fourth swimmeret of male
walking leg
Second swimmeret
Fifth
walking leg Swimmerets 3 to 5
Abdomen Sternum
Anus
Telson
Telson
Uropod Uropod
A B

f i g u r e 12.16
External structure of crayfishes. A, Dorsal view. B, Ventral view.

Ecdysis molt period when its defenseless condition makes it particu-

larly vulnerable to predation.
The problem of growth despite a restrictive exoskeleton is That ecdysis is under hormonal control has been demon-
solved in crustaceans, as in other arthropods, by ecdysis (Gr. strated in both crustaceans and insects, but the process is
ekdysis, to strip off), a periodic shedding of old cuticle and for- often initiated by a stimulus perceived by the central nervous
mation of a larger new one. Molting occurs most frequently system. The action of the stimulus in decapods is to decrease
during larval stages and less often as an animal reaches adult- production of a molt-inhibiting hormone from neurosecre-
hood. Although actual shedding of the cuticle is periodic, the tory cells in the X-organ of the eyestalk. The sinus gland, also
molting process and preparations for it, involving a storage of in the eyestalk, releases the hormone. When the level of molt-
reserves and changes in the integument, are a continuous inhibiting hormone drops, Y-organs near the mandibles pro-
process going on during most of an animal’s life. duce molting hormone. This hormone initiates processes
During each premolt period the old cuticle becomes leading to premolt. Y-organs are homologous to prothoracic
thinner as inorganic salts are withdrawn from it and stored in glands of insects, which produce ecdysone.
tissues. Other reserves, both organic and inorganic, also accu-
mulate and are stored. The underlying epidermis begins to
grow by cell division; it secretes first a new inner layer of epi- Neurosecretory cells are modified nerve cells that secrete
cuticle and then enzymes that digest away the inner layers of hormones. They are widespread in invertebrates and also
old endocuticle (figure 12.17). Gradually a new cuticle forms occur in vertebrates. Cells in the vertebrate hypothalamus
inside the degenerating old one. Finally actual ecdysis occurs and in the posterior pituitary are good examples.
as the old cuticle ruptures, usually along the middorsal line,
and the animal backs out (figure 12.18). By taking in air or
water the animal swells to stretch the new larger cuticle to its
full size. During the postmolt period the cuticle thickens,its
Other Endocrine Functions
outer layer hardens by tanning, and its inner layer is strength- Body color of crustaceans is largely a result of pigments in spe-
ened as salvaged inorganic salts and other constituents are cial branched cells (chromatophores) in the epidermis. Chro-
redeposited. Usually an animal is very secretive during its post- matophores change color by concentrating pigment granules in
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Arthropods 223

Intermolt condition In premolt, old As new exocuticle is

procuticle separates secreted, molting fluid
from epidermis, which dissolves old endocuticle,
secretes new epicuticle and solution products
are reabsorbed

the center of each cell, which causes a lightening effect, or by dispers-

ing pigment throughout each cell, which causes darkening. Neurose-
cretory cells in the eyestalk control pigment behavior. Neurosecretory
hormones also control pigment in the eyes for light and dark adapta-
tion,and other neurosecretory hormones control rate and amplitude of
heartbeat.
Androgenic glands, which are not neurosecretory, occur in
male malacostracans, and their secretion stimulates expression of
male sexual characteristics. If androgenic glands are artificially im-
planted in a female, her ovaries transform to testes and begin to pro-
duce sperm, and her appendages begin to acquire male characteristics
at the next molt.

Feeding Habits
Feeding habits and adaptations for feeding vary greatly among crus-
taceans. Many forms can shift from one type of feeding to another
At ecdysis, the
depending on environment and food availability, but fundamentally old epicuticle and
the same set of mouthparts is used by all. Mandibles and maxillae are exocuticle are discarded
involved in actual ingestion; maxillipeds hold and crush food. In preda-
tors the walking legs, particularly chelipeds, serve in food capture.
Many crustaceans, both large and small, are predatory, and some
have interesting adaptations for killing prey. One shrimplike form,
Lygiosquilla, has on one of its walking legs a specialized digit that can
be drawn into a groove and released suddenly to pierce passing prey.
Pistol shrimp, Alpheus, have one enormously enlarged chela that can
be cocked like the hammer of a gun and snapped shut at great speed,
forming a cavitation bubble that implodes with a snap sufficient to
stun its prey.
Food of crustaceans ranges from plankton, detritus, and bacteria,
used by suspension feeders, to larvae, worms, crustaceans, snails,
and fishes, used by predators, and dead animal and plant matter, used
by scavengers. Suspension feeders, such as fairy shrimps, water fleas,
and barnacles, use their legs, which bear a thick fringe of setae, to cre-
ate water currents that sweep food particles through the setae. Mud In postmolt, new
cuticle is stretched and
shrimps, Upogebia, use long setae on their first two pairs of thoracic unfolded, and endocuticle
appendages to strain food material from water circulated through is secreted
their burrow by movements of their swimmerets.
Crayfishes have a two-part stomach. The first contains a gastric f i g u r e 12.17
mill in which food, already torn up by the mandibles, can be ground Cuticle secretion and reabsorption
up further by three calcareous teeth into particles fine enough to pass in ecdysis.
through a filter of setae in the second part of the stomach; food parti-
cles then pass into the intestine for chemical digestion.
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224 chapter twelve

Respiration, Excretion, and Circulation

Gills of crustaceans vary in shape—treelike, leaflike, or
Abdomen
filamentous—all provided with blood vessels or sinuses. They
Left usually are attached to appendages and kept ventilated by
cheliped
Carapace movement of appendages through water. The overlapping cara-
pace usually protects the branchial chambers. Some smaller
Rupture of membrane
crustaceans breathe through their general body surface.
A and abdomen between carapace Excretory and osmoregulatory organs in crustaceans
are paired glands located in their head, with excretory pores
Old carapace
separates and rises opening at the base of either antennae or maxillae, thus called
antennal glands or maxillary glands, respectively (figure
12.19). Antennal glands of decapods are also called green
Old
glands. They resemble coxal glands of chelicerates. Waste
products are mostly ammonia with some urea and uric acid.
New
Some wastes diffuse through the gills as well as through the
excretory glands.
Circulation, as in other arthropods, is an open system
consisting of a heart, either compact or tubular, and arteries,
B which transport blood to different areas of the hemocoel.
Some smaller crustaceans lack a heart. An open circulatory sys-
tem depends less on heartbeats for circulation because move-
ment of organs and limbs circulates blood more effectively in
open sinuses than in capillaries. Blood may contain as respira-
tory pigments either hemocyanin or hemoglobin (hemocyanin
in decapods), and it has the property of clotting to prevent loss
Abdomen of blood in minor injuries.
emerging
C
Nervous and Sensory Systems
f i g u r e 12.18 A cerebral ganglion above the esophagus sends nerves to the
Molting sequence in a lobster, Homarus americanus. A, Membrane
anterior sense organs and connects to a subesophageal gan-
between carapace and abdomen ruptures, and carapace begins a slow
elevation. This step may take up to two hours. B and C, Head, thorax,
glion by a pair of connectives around the esophagus. A double
and finally abdomen are withdrawn. This process usually takes no more ventral nerve cord has a ganglion in each segment that sends
than 15 minutes. Immediately after ecdysis, chelipeds are desiccated and nerves to viscera, appendages, and muscles (figure 12.19).
the body is very soft. The lobster now begins rapid absorption of water Giant fiber systems are common among crustaceans.
so that within 12 hours its body increases about 20% in length and 50% Sensory organs are well developed. There are two types
in weight. Tissue water will be replaced by protein in succeeding weeks. of eyes—a median, or nauplius, eye and compound eyes. A

f i g u r e 12.19 Tagmata
Internal structure of a male crayfish.
Cephalothorax Abdomen

Eye Brain Stomach Heart

Pericardium
Antenna Rostrum Ostium
Antennule Testis
Intestine

Uropod

Antennal gland
Vas Swimmeret
Mouth
deferens Copulatory Anus
Ganglion swimmeret
Digestive Telson
gland
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Arthropods 225

Corneal
lens
f i g u r e 12.20
Compound eye of an insect. A single
ommatidium is shown enlarged to
Crystalline the right.
cone
Rhabdom

Pigment
Ommatidium cells
Nerve fiber
Retinular
cells
Optic nerve
Ommatidium

Cross section of
ommatidium

Compound eye

median eye consists usually of a group of three pigment cups A hatchling of a crayfish is a tiny juvenile similar in form
containing retinal cells, and it may or may not have a lens. to the adult and has a complete set of appendages and somites.
Median eyes are found in nauplius larvae and in some adult However, most crustaceans produce larvae that must go
forms, and they may be an adult’s only eye, as in copepods. through a series of changes, either gradual or abrupt over a
Most crustaceans have compound eyes similar to series of molts, to assume adult form (metamorphosis). The
insect eyes. In crabs and crayfishes they are on the ends of primitive larva of crustaceans is nauplius (figure 12.21). It has
movable eyestalks (figure 12.19). Compound eyes are precise an unsegmented body, a frontal eye, and three pairs of
instruments, different from vertebrate eyes, yet especially appendages, representing the two pairs of antennae and the
adept at detecting motion; they can analyze polarized light. mandibles. Developmental stages and postlarvae of different
The convex corneal surface gives a wide visual field, particu- groups of Crustacea are varied and have special names.
larly in stalked eyes where the surface may cover an arc of 200
degrees or more.
Compound eyes are composed of many tapering units Class Branchiopoda
called ommatidia set close together (figure 12.20). Facets, or
Members of class Branchiopoda (bran´kee-op´o-da) (Gr.
corneal surfaces, of ommatidia give the surface of the eye an
branchia, gills, + pous, podos, foot) have several primitive
appearance of a fine mosaic. Most crustacean eyes are adapted
characteristics. Four orders are recognized: Anostraca (fairy
either to bright or to dim light, depending on their diurnal or
shrimp and brine shrimp), which lack a carapace; Notostraca
nocturnal habits, but some are able, by means of screening pig-
(tadpole shrimp such as Triops), whose carapace forms a large
ments, to adapt, to some extent at least, to both bright and dim
dorsal shield covering most trunk somites; Conchostraca
light. The number of ommatidia varies from a dozen or two in
(clam shrimp such as Lynceus), whose carapace is bivalved
some small crustaceans to 15,000 or more in a large lobster.
and usually encloses the entire body; and Cladocera (water
Some insects have approximately 30,000.
fleas such as Daphnia, figure 12.22), with a carapace typically
Other sensory organs include statocysts, tactile setae on
covering the entire body but not the head. Branchiopods have
the cuticle of most of the body, and chemosensitive setae, espe-
reduced first antennae and second maxillae. Their legs are flat-
cially on antennae, antennules, and mouthparts.
tened and leaflike (phyllopodia) and are the chief respiratory
organs (hence, the name branchiopods). Legs also are used in
Reproduction and Life Cycles suspension feeding in most branchiopods, and in groups other
than cladocerans, they are used for locomotion as well. The
Most crustaceans have separate sexes, and numerous special- most important and diverse order is Cladocera, which often
izations for copulation occur among different groups. Barna- forms a large segment of freshwater zooplankton.
cles are monoecious but generally practice cross-fertilization.
In some ostracods males are scarce, and reproduction is usu-
ally parthenogenetic. Most crustaceans brood their eggs in
some manner—branchiopods and barnacles have special
Class Maxillopoda
brood chambers, copepods have egg sacs attached to the sides Class Maxillopoda includes a number of crustacean groups tra-
of their abdomen (see figure 12.24), and malacostracans usu- ditionally considered classes themselves. Specialists have rec-
ally carry eggs and young attached to their appendages. ognized evidence that these groups descended from a
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
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226 chapter twelve

Egg

Adult f i g u r e 12.23
An ostracod (subclass Ostracoda, class Maxillopoda).
Nauplius

Protozoea

Postlarval stage

Mysis

f i g u r e 12.21 f i g u r e 12.24
Life cycle of a Gulf shrimp Penaeus. Penaeids spawn at depths of 40 to 90 m. Young larval A copepod with attached ovisacs (subclass Copepoda,
forms are planktonic and move inshore to water of lower salinity to develop as juveniles. class Maxillopoda).
Older shrimp return to deeper water offshore.

Members of subclass Ostracoda (os-trak´o-da) (Gr.

ostrakodes, testaceous, that is, having a shell) are, like con-
chostracans, enclosed in a bivalved carapace and resemble tiny
clams, 0.25 to 8 mm long (figure 12.23). Ostracods show con-
siderable fusion of trunk somites, and numbers of thoracic
appendages are reduced to two or none.
Subclass Copepoda (ko-pep´o-da) (Gr. kōpē, oar, + pous,
podos, foot) is an important group of Crustacea, second only
to Malacostraca in number of species. Copepods are small
(usually a few millimeters or less in length), rather elongate,
tapering toward the posterior end, lacking a carapace, and
retaining a simple, median, nauplius eye in adults (figure
f i g u r e 12.22 12.24). They have four pairs of rather flattened, biramous, tho-
A water flea, Daphnia (order Cladocera), photographed with polarized racic swimming appendages, and a fifth, reduced pair. Their
light. These tiny forms occur in great numbers in northern lakes and are abdomen bears no legs. Many symbiotic as well as free-living
an important component of the food chain leading to fishes. species are known. Many parasites may be so highly modified
as adults (and may depart so far from the description just
given) that they can hardly be recognized as arthropods. Eco-
common ancestor and thus form a clade within Crustacea. logically, free-living copepods are of extreme importance,
They basically have five cephalic, six thoracic, and usually four often dominating the primary consumer level (herbivore) in
abdominal somites plus a telson, but reductions are common. aquatic communities.
No typical appendages occur on the abdomen. Eye of nauplii Subclass Branchiura (bran-kee-u´ra) (Gr. branchia, gills,
(when present) have a unique structure termed a maxillopo- + ura, tail) is a small group of primarily fish parasites, which,
dan eye. despite its name, has no gills (figure 12.25). Members of this
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
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Arthropods 227

barnacles) (figure 12.26A). Typically, a carapace (mantle) sur-

rounds their body and secretes a shell of calcareous plates.
Their head is reduced, their abdomen absent, and their tho-
racic legs are long, many-jointed cirri with hairlike setae. Cirri
are extended through an opening between the calcareous
plates to filter from water small particles on which the animal
feeds (figure 12.26B).
f i g u r e 12.25
Fish louse (subclass Branchiura, class Maxillopoda).
Barnacles frequently foul ship bottoms by settling and
growing there. So great may be their number that the
speed of a ship may be reduced 30% to 40%, requiring
expensive drydocking of the ship to clean them off.

Class Malacostraca
Class Malacostraca (mal´a-kos´tra-ka) (Gr. malakos, soft, +
ostrakon, shell) is the largest class of Crustacea and shows
great diversity. We will mention only 4 of its 12 to 13 orders.
The trunk of malacostracans usually has eight thoracic and six
abdominal somites, each with a pair of appendages. There are
A many marine and freshwater species.
Isopoda (i-sop´o-da) (Gr. isos, equal, + pous, podos, foot)
are commonly dorsoventrally flattened, lack a carapace, and
have sessile compound eyes. Their abdominal appendages bear
gills. Common land forms are sow bugs or pill bugs (Porcellio
and Armadillidium, figure 12.27A), which live under stones
and in damp places. Asellus is common in fresh water, and Ligia
is abundant on sea beaches and rocky shores. Some isopods are
parasites of other crustaceans or of fish (figure 12.28).
Amphipoda (am-fip´o-da) (Gr. amphis, on both sides, +
pous, podos, foot) resemble isopods in that members have no
carapace and have sessile compound eyes. However, they are
usually compressed laterally, and their gills are in the thoracic

f i g u r e 12.26
A, Acorn barnacles, Semibalanus cariosus (subclass Cirripedia) are found
on rocks along the Pacific Coast of North America. B, Common goose-
neck barnacles, Lepas anatifera. Note the feeding legs, or cirri, on Lepas.
Barnacles attach themselves to a variety of firm substrates, including
rocks, pilings, and boat bottoms.

group are usually between 5 and 10 mm long and may be found

on marine or freshwater fish. They typically have a broad,
shieldlike carapace, compound eyes, four biramous thoracic
appendages for swimming,and a short,unsegmented abdomen.
The second maxillae have become modified as suction cups. A
Subclass Cirripedia (sir-i-ped´i-a) (L. cirrus, curl of hair,
+ pes, pedis, foot) includes barnacles, individuals of which are f i g u r e 12.27
usually enclosed in a shell of calcareous plates, as well as three A, Four pill bugs, Armadillidium vulgare
smaller orders of burrowing or parasitic forms. Barnacles are (order Isopoda), common terrestrial
sessile as adults and may be attached to their substrate by a forms. B, Freshwater sow bug,
stalk (gooseneck barnacles) (figure 12.26B) or directly (acorn Caecidotea sp., an aquatic isopod. B
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228 chapter twelve

A B

f i g u r e 12.28
An isopod parasite (Anilocra sp.) on a coney (Cephalopholis fulvus) inhab-
iting a Caribbean coral reef (order Isopoda, class Malacostraca).

position, as in other malacostracans. There are many marine C

amphipods (figure 12.29), such as beach fleas, Orchestia, and
numerous freshwater species. f i g u r e 12.29
Marine amphipods. A, Free-swimming amphipod, Anisogammarus sp.
Euphausiacea (yu-faws´i-a´se-a) (Gr. eu, well, + phausi,
B, Skeleton shrimp, Caprella sp., shown on a bryozoan colony, resemble
shining bright, + acea, L. suffix, pertaining to) is a group of
praying mantids. C, Phronima, a marine pelagic amphipod, takes over
only about 90 species, important as oceanic plankton known the tunic of a salp (subphylum Urochordata, Chapter 15). Swimming by
as “krill.”They are about 3 to 6 cm long (figure 12.30) and com- means of its abdominal swimmerets, which protude from the opening
monly occur in great oceanic swarms, where they are eaten by of the barrel-shaped tunic, the amphipod maneuvers to catch its prey.
baleen whales and many fishes. The tunic is not seen (order Amphipoda, class Malacostraca).
Decapoda (de-cap´o-da) (Gr. deka, ten, + pous, podos,
foot) have five pairs of walking legs of which the first is often
modified to form pincers (chelae) (see figures 12.14 and Myriapods include Chilopoda (centipedes), Diplopoda (milli-
12.16). These are lobsters, crayfishes (see figure 12.14), pedes), Pauropoda (pauropods), and Symphyla (symphylans).
shrimps (see figure 12.21), and crabs, the largest of the crus- Insects have evolved a pattern of three tagmata—head,
taceans (figure 12.31). True crabs differ from others in having thorax, and abdomen—with appendages on the head and tho-
a broader carapace and a much reduced abdomen (figure rax but greatly reduced or absent from the abdomen. The com-
12.31A and C). Familiar examples are fiddler crabs, Uca, which mon ancestor of insects probably resembled myriapods in
burrow in sand just below high-tide level (figure 12.31C), dec- general body form.
orator crabs, which cover their carapaces with sponges and The head of myriapods and insects resembles the crus-
sea anemones for camouflage, and spider crabs, such as Li- tacean head but has only one pair of antennae, instead of
binia. Hermit crabs (figure 12.31B) have become adapted to two. It also has mandibles and two pairs of maxillae (one
live in snail shells; their abdomen, which lacks a hard exoskele- pair of maxillae in millipedes). The legs are all uniramous.
ton, is protected by the snail shell. Respiratory exchange is by body surface and tracheal
systems, although juveniles, if aquatic, may have gills.
Subphylum Uniramia
Appendages of members of Uniramia (yu´ni-ra´me-a) (L. unus,
Class Chilopoda: Centipedes
one, + ramus, a branch) are unbranched, as the name implies. Centipedes are active predators with a preference for moist
This subphylum includes insects and myriapods. The term places such as under logs or stones, where they feed on earth-
myriapod (Gr. myrias, a myriad, + podos, foot) refers to several worms, insects, etc. Their bodies are somewhat flattened
classes that have evolved a pattern of two tagmata—head and dorsoventrally, and they may contain from a few to 177
trunk—with paired appendages on most or all trunk somites. somites (figure 12.32). Each somite, except the one behind the
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
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Edition

Arthropods 229

f i g u r e 12.30
Meganyctiphanes, order Euphausiacea,“northern krill.”

A B C

D E

f i g u r e 12.31
Decapod crustaceans. A, A bright orange tropical rock crab, Grapsus grapsus, is a conspicuous exception to the rule that most crabs bear cryptic
coloration. B, A hermit crab, Elassochirus gilli, which has a soft abdominal exoskeleton, lives in a snail shell that it carries about and into which it can
withdraw for protection. C, A male fiddler crab, Uca sp., uses its enlarged cheliped to wave territorial displays and in threat and combat. D, A red
night shrimp, Rhynchocinetes rigens, prowls caves and overhangs of coral reefs, but only at night. E, Spiny lobster Panulirus argus (order Decapoda,
class Malacostraca).
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

230 chapter twelve

Antenna Eye First tergum Second tergum

Second maxilla

First maxilla
A

Maxilliped with First leg Second leg

B poison fang

f i g u r e 12.32
A, A centipede, Scolopendra (class Chilopoda) from the Amazon Basin, Peru. Most segments have one pair of appendages each. First segment bears a
pair of poison claws, which in some species can inflict serious wounds. Centipedes are carnivorous. B, Head of centipede.

head and the last two, bears one pair of appendages. Those of ments. The four thoracic segments bear only one pair of legs
the first body segment are modified to form poison claws, each, but abdominal segments each have two pairs, a condition
which they use to kill their prey. Most species are harmless to that may have evolved from fusion of somites. Two pairs of spi-
humans. racles occur on each abdominal somite, each opening into an
Their head bears a pair of eyes, each consisting of a group air chamber that gives rise to tracheal tubes.
of ocelli (simple eyes). Respiration is by tracheal tubes with a Millipedes are less active than centipedes and are gener-
pair of spiracles in each somite. Sexes are separate, and all ally herbivorous, living on decayed plant and animal matter and
species are oviparous. Young are similar to adultS. Common sometimes living plants. They prefer dark moist places under
house centipedes Scutigera, with 15 pairs of legs, and Scolopen- stones and logs. Females lay eggs in a nest and guard them care-
dra (figure 12.32), with 21 pairs of legs, are familiar genera. fully. Larval forms have only one pair of legs per somite.

Class Diplopoda: Millipedes Class Insecta: Insects

Diplopods, or “double-footed” arthropods, are commonly called Insects are the most numerous and diverse of all groups of
millipedes, which literally means “thousand feet”(figure 12.33). arthropods (figure 12.34). There are more species of insects
Although they do not have a thousand legs,they do have a great than species in all the other classes of animals combined. The
many. Their cylindrical bodies are made up of 25 to 100 seg- number of insect species named has been estimated at close to

Eyes First segment Second segment

Antenna

Mandible
base
A B Mandible First leg

f i g u r e 12.33
A, A tropical millipede with warning coloration. Note the typical doubling of appendages on most segments, hence diplosegments. B, Head of millipede.
 Hickman−Roberts−Larson: 12. Arthropods Text © The McGraw−Hill
Animal Diversity, Third Companies, 2002
Edition

Arthropods 231

1 million, with thousands, perhaps millions, of other species

yet to be discovered and classified.
It is difficult to appreciate fully the significance of this
extensive group and its role in the biological pattern of animal
life. The study of insects (entomology) occupies the time and
resources of thousands of skilled men and women all over the
world. The struggle between humans and insect pests seems
to be endless, yet paradoxically, insects are so interwoven into
the economy of nature in so many roles that we would have a
difficult time without them.
Insects differ from other arthropods in having three
pairs of legs and usually two pairs of wings on the thoracic
region of the body (figure 12.35), although some have one pair
of wings or none. In size insects range from less than 1 mm to
20 cm in length, the majority being less than 2.5 cm long.

Distribution and Adaptability

Insects have spread into practically all habitats that can sup-
port life, but only a relatively few are marine. They are com-
mon in brackish water, in salt marshes, and on sandy beaches.
They are abundant in fresh water, soils, forests, and plants, and
they are found even in deserts and wastelands, on mountain-
tops, and as parasites in and on the bodies of plants and ani-
mals, including other insects.
Their wide distribution is made possible by their powers
of flight and their highly adaptable nature. In many cases they
can easily surmount barriers that are impassable to many other
animals. Their small size and well-protected eggs allow them
to be carried great distances by wind, water, and other animals.
The amazing adaptability of insects is evidenced by their
f i g u r e 12.34 wide distribution and enormous diversity of species. Such
Pie diagram indicating relative numbers of species of insects to the rest diversity enables this vigorous group to take advantage of all
of the animal kingdom and protozoan groups. available resources of food and shelter.

Forewing
Hindwing
Metathorax

Compound
eye Tympanum MALE
Prothorax
Mesothorax
A Antenna
Ocelli Cercus
Frons
f i g u r e 12.35 Gena
Ovipositor
A, A pair of grasshoppers, Schistocerca obscura
(order Orthoptera), copulating. The African Tibia Spiracles
Clypeus Labial
desert locust mentioned in the chapter prologue Tergum
(p. 212) is Schistocerca gregaria.B, External features palp Tarsus
Labrum Coxa
of a female grasshopper. The terminal segment of Maxillary Femur Sternum
Mandible palp Trochanter
a male with external genitalia is shown in inset. B