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Miller the Zoology Book


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Miller

OF 269

Harley: Zoology,Fifth
EditionII. Animal

Like Protistsand
Animalia14. The
Arthropods:Blueprint for
Success
The McGraw

HillCompanies, 2001

CHAPTER14The Art
hropods

213
The circulatory system
of arachnids, like that of
mostarthropods, is an
open system in which a
dorsal contractile
vessel(usually called
the dorsal aorta or
heart) pumps blood
into tissuespaces. In
arthropods, the coelom

is reduced to cavities
surround-ing the gonads
and sometimes the
coxal glands. Large
tissuespaces, or sinuses,
are derived from the
blastocoel and are
calledthe
hemocoel.
Blood bathes the tissues
and then returns to

thedorsal aorta through


openings in the aorta
called ostia.
Arachnidblood contains
the dissolved
respiratory pigment
hemocyanin andhas
amoeboid cells that aid
in clotting and body
defenses.The nervous
system of all arthropods

is ventral and, in ancestral arthropods, must


have been laid out in a
pattern similar tothat of
the annelids
(see gure 13.8a)
. With the exception of
scor-pions, the nervous
system of arachnids is
centralized by fusion
of ganglia.The body of

an arachnid has
a variety of sensory
structures.Most
mechanoreceptors and
chemoreceptors are
modications of the
exoskeleton, such as
projections, pores, and
slits, togetherwith
sensory and accessory

cells. Collectively,
these receptors arecalled
sensilla.
For example, setae
are hairlike, cuticular
modica-tions that may
be set into membranous
sockets. Displacement
of aseta initiates a nerve
impulse in an associated
nerve cell (gure14.10

a
). Vibration receptors
are very important to
some arach-nids.
Spiders that use webs
to capture prey, for
example, determineboth
the size of the insect
and its position on the
web by the vi-brations
the insect makes while

struggling to free itself.


Thechemical sense of
arachnids is comparable
to taste and smell
invertebrates. Small
pores in the exoskeleton
are frequently associated with peglike, or
other, modications of
the exoskeleton, andthey
allow chemicals to

stimulate nerve cells.


Arachnids possessone
or more pairs of eyes,
which they use
primarily for
detectingmovement and
changes in light
intensity (gure 14.10
b
). Theeyes of
some hunting spiders

probably form
images.Arachnids are
dioecious. Paired
genital openings are
on theventral side of the
second abdominal
segment. Sperm transfer
isusually indirect. The
male often packages
sperm in a spermatophore, which is

then transferred to the


female. Courtship rituals conrm that
individuals are of the
same species, attract a
fe-male to the
spermatophore, and
position the female
to receive
thespermatophore. In
some taxa (e.g.,

spiders), copulation
occurs,and sperm is
transferred via a
modied pedipalp of
the male.Development
is direct, and the young
hatch from eggs as
minia-ture adults. Many
arachnids tend their
developing eggs and

youngduring and after


development.

Order
Scorpionida
Members of the order
Scorpionida (skor
pe-ah-ni

dah) are thescorpions


(gure 14.11
a
). They are common
from tropical towarm
temperate climates.
Scorpions are secretive
and nocturnal,hiding
during most daylight
hours under logs and
stones.Scorpions have

small chelicerae that


project anteriorly
fromthe front of the
carapace (gure 14.11
b
). A pair of
enlarged,chelate
pedipalps is posterior
to the chelicerae. The
opisthosomais divided.
An

anterior preabdomen
contains the slitlike
openingsto book lungs,
comblike tactile and
chemical receptors
calledpectines, and
genital openings. The
postabdomen
(commonlycalled the
tail) is narrower than
the preabdomen and is

curveddorsally and
anteriorly over the body
when aroused. At
the tip of the
postabdomen is a sting.
The sting has a bulbular
base that con-tains
venom-producing glands
and a hollow, sharp,
barbed point.Smooth
muscles eject venom

during stinging. Only a


few scorpi-ons have
venom that is highly
toxic to humans.
Species in thegenera
Androctonus
(northern Africa) and
Centuroides
(Mexico,

Lamellaeof lungBlood
flowbetween lamellaeAir flowing
inthrough spiracle

FIGURE 14.9
Arachnid Book Lung.
Air and blood moving on
opposite sides of alamella
of the lung exchange
respiratory gases by
diffusion. Figure
14.12shows the location of
book lungs in spiders.

LensHypodermisArticularmembr
aneLight-sensitivecellNerve
fibersto brainReceptor cell

(
)

FIGURE 14.10
Arthropod Seta and
Eye (Ocellus).
(a)
A seta is a hairlike
modica-tion of the cuticle
set in a membranous

socket. Displacement of
the setainitiates a nerve
impulse in a receptor cell
(sensillum) associated
withthe base of the seta.
(b)
The lens of this spider eye
is a thickened, trans-parent
modication of the cuticle.
Below the lens and
hypodermis arelightsensitive sensillae with

pigments that convert light


energy intonerve impulses.

Miller

Harley: Zoology,Fifth
EditionII. Animal


Like Protistsand
Animalia14. The
Arthropods:Blueprint for
Success
The McGraw

HillCompanies, 2001

214

PART TWOAnimallike Protists and


Animalia
Arizona, and New
Mexico) have been
responsible for
humandeaths. Other
scorpions from the
southern and
southwestern areasof
North America give

stings comparable to
wasp stings.Prior to
reproduction, male and
female scorpions have a
pe-riod of courtship that
lasts from ve minutes
to several hours.Male
and female scorpions
face each other and
extend their ab-domens
high into the air. The

male seizes the female


with hispedipalps, and
they repeatedly walk
backward and then
forward.The lower
portion of the male
reproductive tract forms
a sper-matophore that is
deposited on the
ground. During
courtship, themale

positions the female so


that the genital opening
on her ab-domen is
positioned over the
spermatophore.
Downward pressureof
the females abdomen
on a triggerlike
structure of the spermatophore releases
sperm into the females

genital chamber.Most
arthropods are
oviparous;
females lay eggs that
de-velop outside the
body. Many scorpions
and some arthropods are
ovoviviparous;
development is internal,
although large,

yolkyeggs provide all


the nourishment for
development. Some
scorpi-ons, however, are
viviparous,
meaning that the mother
providesnutrients to
nourish the embryos.
Eggs develop in
diverticula of the ovary
that are closely

associated with
diverticula of the digestive tract. Nutrients pass
from the digestive tract
diverticula to
thedeveloping embryos.
Development requires
up to 1.5 years,
and 20to 40 young are
brooded. After birth, the
young crawl onto

themothers back, where


they remain for up to a
month.

Order
Araneae
With about 34,000
species, the order
Araneae (ah-ran

a-e) is thelargest group


of arachnids (gure
14.12). The prosoma of
spidersbears chelicerae
with poison glands and
fangs. Pedipalps are
leg-like and, in males,
are modied for sperm
transfer. The
dorsal,anterior margin
of the carapace usually

has six to eight eyes.A


slender, waistlike
pedicel attaches the
prosoma to
theopisthosoma. The
abdomen is swollen or
elongate and
containsopenings to the
reproductive tract, book
lungs, and tracheae.
Italso has six to

eight conical
projections, called
spinnerets, that
areassociated with silk
glands. The protein
that forms silk is
emittedas a liquid, but
hardens as it is drawn
out. Spiders produce
severalkinds of silk,
each with its own use.

In addition to webs for


cap-turing prey (gure
14.13), silk is used to
line retreats, to lay
a safetyline that fastens
to the substrate to
interrupt a fall, and to
wrapeggs into a case for
development. Air
currents catch silk lines
thatyoung spiders

produce and disperse


them. Silk lines have
carriedspiders at great
altitudes for hundreds
of kilometers. This is
calledballooning.
FIGURE 14.11
Order Scorpionida.
(a)Centruroides sculpturat
us
is shown here.

(b)
Ex-ternal anatomy of a
scorpion.
PedipalpsC h e l i c e r a W
a l k i n g l e g s Median
eyeLateral eyeAbdominal
stingerP r o s o m a P r e a b d o m e
n P o s t a b d o m e n Opisthosoma

(b)(a)
ProsomaBook lung
slitSpiracle SpinneretsSternum
Chelicerawith
fangOpisthosomaEyesCheliceraP
edipalpPedicelAbdomen

(a)(b)

FIGURE 14.12
External Structure of a
Spider.
(a)
Dorsal view.
(b)
Ventral view.
Sources: (a) After Sherman and
Sherman. (b) After the Kastons.

Miller

Harley: Zoology,Fifth
EditionII. Animal

Like Protistsand
Animalia14. The
Arthropods:Blueprint for
Success
The McGraw

HillCompanies, 2001

CHAPTER14The Art
hropods

215
Most spiders feed on
insects and other
arthropods that
theyhunt or capture in
webs. A few (e.g.,
tarantulas or bird
spiders)feed on small
vertebrates. Spiders bite
their prey to paralyze
themand then sometimes

wrap prey in silk. They


puncture the preysbody
wall and
inject enzymes. The
spiders pumping
stomach thensucks
predigested prey
products into the
spiders digestive
tract.The venom of
most spiders is harmless

to humans. Black
widowspiders
(Lactrodectus)
and brown recluse
spiders
(Loxosceles)
areexceptions, since
their venom is toxic to
humans (gure
14.14).Mating of

spiders involves
complex behaviors that
includechemical, tactile,
and/or visual signals.
Females deposit
chemicalscalled
pheromones on their
webs or bodies to
attract
males.(Pheromones are
chemicals that one

individual releases into


theenvironment to
create a behavioral
change in another
member of the same
species.) A male may
attract a female by
plucking thestrands of a
females web. The
pattern of plucking is
species spe-cic and

helps identify and


locate a potential mate
and preventsthe male
spider from becoming
the females next meal.
The tips of a males
pedipalps possess a
bulblike reservoir with
an ejaculatoryduct and a
penislike structure
called an embolus. Prior

to mating,the male lls


the reservoir of his
pedipalps by depositing
sperm ona small web
and then collecting
sperm with his
pedipalps.
Duringmating, the
pedipalp is engorged
with blood, the embolus
is in-serted into the

females reproductive
opening, and sperm
aredischarged. The
female deposits up to
3,000 eggs in a silken
eggcase, which she then
seals and attaches to
webbing, places in a retreat, or carries with
her.

Order
Opiliones
Members of the order
Opiliones (o
pi-le
on-es) are the harvestmen or daddy longlegs.
The prosoma broadly

joins to the opisthosoma, and thus, the


body appears ovoid.
Legs are very long
andslender. Many
harvestmen are
omnivores (they feed on
a varietyof plant and
animal material), and
others are strictly
predators.They seize

prey with their


pedipalps and ingest
prey as describedfor
other arachnids.
Digestion is both
external and
internal.Sperm transfer
is direct, as males have
a penislike structure.
(a)(b)

FIGURE 14.13

Order Araneae.
Members of the family
Araneidae, the orb
weavers,produce some of
the most beautiful and
intricate spider webs.
Manyspecies are relatively
large, like this garden
spider

Argiope.

A web isnot a permanent


construction. When webs
become wet with rain
ordew, or when they age,
they lose their stickiness.
The entire web, or atleast
the spiraled portion, is
then eaten and replaced.
FIGURE 14.14
Two Venomous Spiders.
(a)
A black widow spider

(Lactrodectus mac-tans)
is recognized by its shiny
black body with a red
hourglass patternon the
ventral surface of its
opisthosoma.
(b)
A brown recluse spider
(Loxosceles reclusa)
is recognized by the dark
brown, violin-shaped

markon the dorsal aspect


of its prosoma.

Miller

Harley: Zoology,Fifth
EditionII. Animal

Like Protistsand
Animalia14. The
Arthropods:Blueprint for
Success
The McGraw

HillCompanies, 2001

216

PART TWOAnimallike Protists and


Animalia
Females have a tubular
ovipositor that projects
from a sheath atthe time
of egg laying. Females
deposit hundreds of
eggs in damplocations
on the ground.

Order Acarina
Members of the order
Acarina (ak
ar-i
nah) are the mites
andticks. Many are
ectoparasites (parasites
on the outside of

thebody) on humans
and domestic animals.
Others are free-living
inboth terrestrial and
aquatic habitats. Of all
arachnids, acarineshave
had the greatest impact
on human health and
welfare.Mites are 1 mm
or less in length. The
prosoma and opistho-

soma are fused and


covered by a single
carapace. An anterior
pro-jection called the
capitulum carries
mouthparts. Chelicerae
andpedipalps are
variously modied for
piercing, biting,
anchoring,and sucking,
and adults have four

pairs of walking
legs.Free-living mites
may be herbivores or
scavengers. Herbivorous mites, such as
spider mites, damage
ornamental and agricultural plants. Scavenging
mites are among the
most commonanimals
in soil and in leaf litter.

These mites include


some pestspecies that
feed on our, dried fruit,
hay, cheese, and animal
fur(gure
14.15).Parasitic mites
usually do not
permanently attach to
theirhosts, but feed for a
few hours or days and
then drop to the

ground.One mite, the


notorious chigger or red
bug
(Trombicula),
is a para-site during one
of its larval stages on all
groups of
terrestrial verte-brates.
A larva enzymatically
breaks down and sucks
host skin,causing local

inammation and
intense itching at the
site of thebite. The
chigger larva drops
from the host and then
molts to thenext
immature stage, called a
nymph.
Nymphs eventually
molt toadults, and both
nymphs and adults feed

on insect eggs.A few


mites are permanent
ectoparasites. The
follicle mite,
Demodex folliculorum,
is common (but
harmless) in hair
follicles of most of the
readers of this text. Itch
mites cause scabies in

humansand other
animals.
Sarcoptes scabei
is the human itch mite.
It tun-nels in the
epidermis of human
skin, where females lay
about 20eggs each day.
Secretions of the mites
irritate the skin,
and infec-tions are

acquired by contact with


an infected
individual.Ticks are
ectoparasites during their
entire life history.
Theymay be up to 3 cm
in length, but are
otherwise similar to
mites.Hooked
mouthparts are used to
attach to their hosts and

to feedon blood. The


female ticks, whose
bodies are less
sclerotized thanthose of
males, expand
when engorged with
blood. Copulation occurs on the host, and
after feeding, females
drop to the ground tolay
eggs. Eggs hatch into

six-legged immatures
called seed
ticks.Immatures feed on
host blood and drop to
the ground for
eachmolt. Some ticks
transmit diseases to
humans and domestic
ani-mals. For example,
Dennacentor andersoni

transmits the
bacteriathat cause
Rocky Mountain
spotted fever and
tularemia, and
Ixodes scapularis
transmits the bacteria
that cause Lyme
disease(gure
14.16).Other orders of
arachnids include whip

scorpions, whipspiders,
pseudoscorpions,
and others.

CLASS
PYCNOGONIDA
Members of the class
Pycnogonida (pik
no-gon

i-dah) are the


seaspiders. All are
marine and are most
common in cold
waters(gure 14.17).
Pycnogonids live on the
ocean oor
and frequentlyfeed on
cnidarian polyps and
ectoprocts. Some sea
spiders feed bysucking

prey tissues through a


proboscis. Others tear at
prey withtheir
chelicerae.Pycnogonids
are dioecious. Gonads
are
U
-shaped, andbranches of
the gonads extend into
each leg. Gonopores are
onone of the pairs

of legs. As the female


releases eggs, the male
fer-tilizes them, and the
fertilized eggs are
cemented into
sphericalmasses and
attached to a pair of
elongate appendages of
the male,called ovigers,
where they are brooded
until hatching.

SUBPHYLUM
CRUSTACEA
Some members of the
subphylum Crustacea
(krus-tas
e-ah) (L.
crustaceus,
hard shelled), such as
craysh, shrimp,

lobsters, andcrabs, are


familiar to nearly
everyone. Many others
are lesser-known but
very common taxa.
These include copepods,
cladocer-ans, fairy
shrimp, isopods,
amphipods, and
barnacles. Except
forsome isopods and

crabs, crustaceans are


all aquatic.Crustaceans
differ from other living
arthropods in two
ways.They have two
pairs of antennae,
whereas all other
arthropodshave one pair
or none. In addition,
crustaceans possess
biramousappendages,

each of which consists of


a basal segment, called
the
protopodite,
with two rami (distal
processes that give the
ap-pendage a
Y
shape) attached. The
medial ramus is the

endopodite,
and the lateral ramus is
the
exopodite
(gure 14.18).
T

rilobiteshad similar
structures, which
may be evidence that
the trilo- bites were

ancestral to the
crustaceans.
CLASS
MALACOSTRAC
A
Malacostraca (mal-ahkos
trah-kah) (Gr.
malakos,

soft
ostreion,
shell) is the largest
class of crustaceans. It
includes crabs,
lobsters,craysh,
shrimp, mysids,
shrimplike
krill, isopods, and
amphipods.

FIGURE 14.15
Order Acarina.
Dermatophagoides farinae
(

200) is common in
homesand grain storage
areas. It is believed to be a
major cause of dust
allergies.

Miller

Harley: Zoology,Fifth
EditionII. Animal

Like Protistsand
Animalia14. The
Arthropods:Blueprint for
Success
The McGraw

HillCompanies, 2001

CHAPTER14The Art
hropods

217
The order Decapoda
(dek-i-pod
ah) is the largest order
of crustaceans and
includes shrimp,
craysh, lobsters, and
crabs.Shrimp have a
laterally compressed,
muscular abdomen

andpleopods for
swimming. Lobsters,
crabs, and craysh are
adaptedto crawling on
the surface of the
substrate (gure 14.19).
The ab-domen of crabs
is greatly reduced and is
held exed beneath
thecephalothorax.Cray
sh illustrate general

crustacean structure and


func-tion. They are
convenient to study
because of their relative
abun-dance and large
size. The body of a
craysh is divided into
two re-gions. A
cephalothorax is
derived
from developmental

fusion of asensory and


feeding tagma (the
head) with a locomotor
tagma(the thorax). The
exoskeleton of the
cephalothorax extends
lat-erally and ventrally
to form a shieldlike
carapace. The abdomen
isposterior to the
cephalothorax, has

locomotor and visceral


func-tions, and in
craysh, takes the form
of a muscular
tail.Paired
appendages are present
in both body regions
(gure14.20). The rst
two pairs of
cephalothoracic
appendages are therst

and second antennae.


The third through fth
pairs of ap-pendages are
associated with the
mouth. During
crustacean evolu-tion,
the third pair of
appendages became
modied into
chewingor grinding
structures called

mandibles.
The fourth and fth
pairsof appendages,
called
maxillae,
are for food handling.
The secondmaxilla
bears a gill and a thin,
bladelike structure,
called ascaphognathite
(gill bailer), for

circulating water over


the gills.The sixth
through the eighth
cephalothoracic
appendages arecalled
maxillipeds and
are derived from the
thoracic tagma. Theyare
accessory sensory
and food-handling
appendages. The

last twopairs of
maxillipeds also bear
gills. Appendages 9 to
13 are tho-racic
appendages called
periopods (walking
legs). The rst periopod, known as
the cheliped, is enlarged
and chelate
(pincherlike)and used in

defense and capturing


food. All but the last
pairof appendages of
the abdomen are called
pleopods
(swimmerets)and used
for swimming. In
females, developing
eggs attach topleopods,
and the embryos are
brooded until after

hatching. Inmales, the


rst two pairs of
pleopods are modied
into gonopods
(a)(b)

FIGURE 14.16
Order Acarina.
(a)Ixodes scapularis,
the tick that transmits the
bacte-ria that cause Lyme
disease.

(b)
The adult (shown here) is
about the sizeof a sesame
seed, and the nymph is the
size of a poppy seed.
People walk-ing in tickinfested regions should
examine themselves
regularly and re-move any
ticks found on their skin
because ticks can transmit
diseases,such as

Rocky Mountain spotted


fever, tularemia, and Lyme
disease.
FIGURE 14.17
Class Pycnogonida.
Sea spiders are often
found in intertidal
regionsfeeding on
cnidarian polyps. This
male sea spider
(Nymphon gracile)

iscarrying eggs on ovigers.

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