C H A P T E R

Crustaceans
20
• PHYLUM ARTHROPODA
• SUBPHYLUM CRUSTACEA

Crustacea

Arthropoda

A Sally Lightfoot crab, Grapsus grapsus, from the Galápagos Islands.

“Insects of the Sea”

The Crustacea (L. crusta, shell) are named after the hard shell that most Age of Mammals. Together, insects and crustaceans compose more
crustaceans bear. Over 67,000 species have been described, and sev- than 80% of all named animal species. Just as insects pervade ter-
eral times that number probably exist. Most familiar to people are the restrial habitats (more than a million named species and countless
edible ones, for example, lobsters, crayfishes, shrimps, and crabs. In trillions of individuals), crustaceans abound in oceans, lakes, and
addition to these “crusty” crustaceans, there is an astonishing array of rivers. Some walk, some burrow, and some (such as barnacles) are
less familiar forms such as copepods, ostracods, water fleas, whale lice, sessile. Some swim upright, others swim upside down, and many
tadpole shrimp, and krill. They fill a wide variety of ecological roles are delicate microscopic forms that drift as plankton in oceans or in
and show enormous variation in morphological characteristics, making lakes. Indeed, it is probable that some of the most abundant animals
a satisfactory description of the group as a whole very difficult. in the world are members of the copepod genus Calanus. In rec-
We live in the Age of Arthropods, notwithstanding our anthro- ognition of their dominance of marine habitats, it is understandable
pocentric attachment to our tradition of calling the current era the that crustaceans have been called “insects of the sea.”

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E
xtant arthropods are divided among four subphyla (see Fig- correct to refer to insects as “terrestrial crustaceans.” Our descrip-
ure 19.2). Crustacea and Hexapoda share five derived fea- tion of crustaceans as “insects of the sea” in the prologue to this
tures and are united in clade Pancrustacea (Figure 20.1). We chapter describes only the ecological role of these animals.
depict crustaceans and hexapods as sister taxa, but some phylog- Crustaceans are divided among five classes (Figure 20.1),
enies using molecular characters support the hypothesis that hexa- although preliminary phylogenies using molecular characters do
pods arose from within the crustacean lineage. If the same pattern not support the monophyly of all classes. We have placed mem-
emerges from studies using other genes, it will be phylogenetically bers of the former phylum Pentastomida in class Maxillopoda,

Pancrustacea
Crustacea

Hexapoda Remipedia Cephalocarida Brachiopoda Maxillopoda Malacostraca

Genital appendages on
Genital appendages
1st abdominal segment
Unique tracheal system on 1st abdominal
segment Maxillopodan naupliar eye
Mouthparts for Abdomen reduced
Abdomen < 4 segments
catching prey to 11 segments
2nd maxillae reduced Thorax < 6 segments
Thorax 8 segments,
Malpighian tubules or absent
No abdominal legs abdomen 6 or 7 segments
plus telson
Maxilliped
No abdominal legs No abdominal
legs Maxillipeds
“Whole-limb” mandibles Reduced carapace
Lateral trunk Abdomen < 8 segments
appendages
Thorax < 11 segments

Loss of 2nd antennae Carapace

Abdomen < 9 segments

6 legs
Reduction in segment number

Tagmata of thorax and abdomen

Biramous 2nd antennae

Stalked compound eyes?
“Gnathobasic” mandibles
Nauplius larva

All head appendages except 1st antennae used for feeding sometime in life
Mandibulate structure of ommatidia
Tripartite brain
Appendages of 3rd postacronal segment are mandibles

Two pairs maxillae on postacronal segments 4 and 5

Figure 20.1
Cladogram showing hypothetical ancestor-descendant relationships of hexapods and classes of crustaceans. Hexapods and crustaceans are
hypothetical sister groups evolving from a common ancestor defined by numerous shared derived characteristics. Characters followed by a
question mark may be ancestral rather than shared derived features. The acron is the anterior region of the head and is not counted as a segment.

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 422 PART THREE Diversity of Animal Life

subclass Pentastomida. Pentastomids, often called tongue worms, Antenna

are parasites of vertebrates, living in lungs or nasal cavities. They
are closely related to fish lice in subclass Branchiura. Rostrum
Cephalothorax Abdomen
Features of crustacean classes and subclasses are discussed 13 segments 6 segments
Eye
following a general introduction to crustacean biology. Antennule Carapace

Thorax
SUBPHYLUM CRUSTACEA
Mandible
General Nature of a Crustacean Maxillae
Crustacea are mainly marine; however, there are many freshwater Maxillipeds
and a few terrestrial species. Crustaceans differ from other arthro-
pods in a variety of ways, but the distinguishing characteristic is that Cheliped Telson
(first leg)
crustaceans are the only arthropods with two pairs of antennae. Uropod
In addition to two pairs of antennae and a pair of mandibles, crus- Chela
taceans have two pairs of maxillae on the head, followed by a pair
of appendages on each body segment. In some crustaceans not all
segments bear appendages. All appendages, except perhaps the Walking legs Swimmerets
first antennae, are ancestrally biramous (two main branches), and
at least some appendages of present-day adults show that condition. Figure 20.2
Organs specialized for respiration, if present, function as gills. Archetypical plan of Malacostraca. The two maxillae and three
maxillipeds have been separated diagrammatically to illustrate the
Most crustaceans have between 16 and 20 segments, but general plan.
some forms have 60 segments or more. A larger number of seg-
ments is an ancestral feature. The more derived condition is to
have fewer segments and increased tagmatization (see p. 406). External Features
Major tagmata are head, thorax, and abdomen. In most Crustacea Bodies of crustaceans are covered with a secreted cuticle com-
one or more thoracic segments are fused with the head to form posed of chitin, protein, and calcareous material. The harder,
a cephalothorax. Tagmata are not homologous throughout the heavy plates of larger crustaceans are particularly high in calcar-
subphylum (or even within some classes) because in different eous deposits. The hard protective covering is soft and thin at
groups different segments may have fused to form what we now the joints between segments, allowing flexibility of movement.
call, for example, a head or a cephalothorax. The carapace, if present, covers much or all of the cephalotho-
By far the largest class of crustaceans is the class Malacos- rax; in decapods such as crayfishes, all head and thoracic seg-
traca, which includes lobsters, crabs, shrimps, beach hoppers, ments are enclosed dorsally by the carapace. Each segment not
sow bugs, and many others. These species show a surprisingly enclosed by the carapace is covered by a dorsal cuticular plate,
constant arrangement of body segments and tagmata, which is or tergum (Figure 20.3A), and a ventral transverse bar, or ster-
considered the ancestral plan of this class (Figure 20.2). This num, lies between the segmental appendages (Figure 20.3B).
typical body plan has a head of five (six embryonically) fused The abdomen terminates in a telson that bears the anus.
segments, a thorax of eight segments, and an abdomen of six Position of the gonopores varies according to sex and
segments (seven in a few species). At the anterior end is a non- group of crustaceans. They may be on or at the base of a pair
segmented rostrum and at the posterior end is a nonsegmented of appendages, at the terminal end of the body, or on segments
telson, which with the last abdominal segment and its uropods without legs. For example, in crayfishes openings of the vasa
forms a tail fan in many forms. deferentia are on the median side at the base of the fifth pair
In many crustaceans the dorsal cuticle of the head may extend of walking legs, and those of the oviducts are at the base of the
posteriorly and around the sides of the animal to cover or be fused third pair. In females an opening to the seminal receptacle is
with some or all of the thoracic and abdominal segments. This cov- usually located in the midventral line between the fourth and
ering is called a carapace (Figure 20.2). In some groups the cara- fifth pairs of walking legs.
pace forms clamshell-like valves that cover most or all of the body.
In decapods (including lobsters, shrimp, crabs, and others), the Appendages Members of classes Malacostraca (for example,
carapace covers the entire cephalothorax but not the abdomen. crayfishes) and Remipedia typically have a pair of jointed append-
ages on each segment (Figure 20.3B), although abdominal seg-
ments in the other classes typically do not bear appendages.
Form and Function Considerable specialization is evident in appendages of derived
Because of their size and easy availability, large crustaceans such crustaceans such as crayfishes. The basic, biramous plan is illus-
as crayfishes have been well studied and are commonly included trated by a crayfish appendage such as a maxilliped, a thoracic
in introductory laboratory courses. Hence many of the comments limb modified to become a feeding appendage (Figure 20.4). The
here apply specifically to crayfishes and their relatives. basal portion, or protopod, bears a lateral exopod and a medial

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 www.mhhe.com/hickmanipz14e CHAPTER 20 Crustaceans 423

Antenna Antenna

Chela Antennules Antennules Chela

Cheliped
Exopod of first
maxilliped
Rostrum
Second maxilliped
Eye
Mouth
Second
walking leg Third maxilliped

Third Second
walking leg walking leg
Carapace Third walking leg
Cervical groove
Fourth Fourth walking leg
walking leg
First (copulatory)
swimmeret of male

Second
swimmeret
Fifth
walking leg
Swimmerets 3 to 5

Tergum Sternum
Abdomen Anus
Telson
Telson
Uropod Uropod

Figure 20.3
External structure of crayfishes. A, Dorsal view. B, Ventral view.

endopod. The protopod is composed of two parts (basis and

coxa), whereas the exopod and endopod have from one to sev-
eral parts each. Variations on the basic form exist (Figure 20.5).
Some appendages, such as the walking legs of crayfishes, have Gill
become secondarily uniramous. Medial or lateral processes,
called endites and exites, respectively, sometimes occur on
crustacean limbs. An exite on the protopod is called an epipod, Coxa
which is often modified as a gill. Table 20.1 shows how various Protopod
appendages have become modified from the presumed ancestral Basis
biramous plan and now perform disparate functions.
Exopod
Terminology applied by various workers to crustacean appendages Endopod
is not uniform. At least two systems are in wide use. Alternative
terms to those we have used, for example, are protopodite, exopo-
dite, endopodite, basipodite, coxopodite, and epipodite. The first
and second pairs of antennae may be termed antennules and
Figure 20.4
Parts of a biramous crustacean appendage
antennae, and first and second maxillae are often called maxillules (third maxilliped of a crayfish). The two
and maxillae. A rose by any other name . . . branches of the appendage are the exopod and
the endopod; both extend from the protopod.

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 424 PART THREE Diversity of Animal Life

Antennule

Antenna Cheliped

Second
Mandible walking
leg

First
maxilla Fourth
walking
leg
Second
maxilla
FEMALE
MALE
FIRST
First
maxilliped
SECOND
Swimmerets

Second
maxilliped THIRD

Third Uropod
maxilliped

Figure 20.5
Appendages of a crayfish showing variations on the basic biramous plan, as found in a swimmeret. Protopod, brown; endopod, blue; exopod, yellow.

Structures that share a similar basic plan and have descended compartments remaining are the end sacs of excretory organs
from a common ancestral form are said to be homologous, and the space around the gonads.
whether they have the same function or not. Since specialized
walking legs, mouthparts, chelipeds, and swimmerets have Muscular System Striated muscles form a considerable part
all developed from a common biramous appendage (that has of the body of most Crustacea. Muscles are usually arranged
become modified to perform different functions), they are all in antagonistic groups: flexors, which draw a part toward the
homologous to each other—a condition called serial homol- body, and extensors, which extend the part outward. The abdo-
ogy. Limbs ancestrally were all very similar, but during struc- men of a crayfish has powerful flexors (Figure 20.6), which are
tural evolution some branches have been reduced, some lost, used when the animal swims backward with a burst of speed to
some greatly altered, and some new parts added. Crayfishes and escape from predators.
their allies possess the most elaborate serial homology in the
animal kingdom, with 17 distinct but serially homologous types Respiratory System Gas exchange in smaller crustaceans
of appendages (Table 20.1). For example, compare the size of occurs across thin areas of cuticle (for example, in the append-
the chela of the cheliped to the tiny claw (chela) of the second ages) or the entire body, and specialized structures for gas
walking leg in Figure 20.5. exchange may be absent. Larger crustaceans have gills, which are
delicate, featherlike projections with very thin cuticle. In deca-
pods the sides of the carapace enclose the gill cavity, which is
Internal Features open anteriorly and ventrally (Figure 20.7). Gills may project from
The muscular and nervous systems in the thorax and abdomen the pleural wall into the gill cavity, from the articulation of tho-
clearly show segmentation, but there are marked modifications racic legs with the body, or from thoracic coxae. The latter two
in other systems. Most changes involve concentration of parts types are typical of crayfishes. The “bailer,” a part of the second
in a particular region or else reduction or complete loss of maxilla, draws water over the gill filaments, into the gill cavity at
parts. the bases of the legs, and out of the gill cavity at the anterior.

Hemocoel The major body space in arthropods is not a Circulatory System Crustaceans and other arthropods have
coelom, but a persistent blastocoel that becomes a blood-filled an “open” or lacunar type of circulatory system. This means that
hemocoel (see p. 396). In crustaceans the only coelomic there are no veins and no separation of blood from interstitial

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TABLE 20.1
Crayfish Appendages
Appendage Protopod Endopod Exopod Function
First antenna 3 segments, Many-jointed feeler Many-jointed feeler Touch, taste,
(antennule) statocyst in base equilibrium
Second antenna 2 segments, excretory Long, many-jointed Thin, pointed blade Touch, taste
(antenna) pore in base feeler
Mandible 2 segments, heavy jaw and 2 distal segments Absent Crushing food
base of palp of palp
First maxilla 2 segments, with Small unjointed Absent Food handling
(maxillule) 2 thin endites lamella
Second maxilla 2 segments, with 1 small pointed Part of scaphognathite Drawing currents
(maxilla) 2 endites and 1 segment (bailer) of water into gills
scaphognathite (epipod)
First maxilliped 2 medial plates 2 small segments 1 basal segment, plus Touch, taste,
and epipod many-jointed filament food handling
Second maxilliped 2 segments plus gill (epipod) 5 short segments 2 slender segments Touch, taste,
food handling
Third maxilliped 2 segments plus gill (epipod) 5 larger segments 2 slender segments Touch, taste,
food handling
First walking leg 2 segments plus gill (epipod) 5 segments with Absent Offense and defense
(cheliped) heavy pincer (chela)
Second walking leg 2 segments plus gill (epipod) 5 segments plus Absent Walking and prehension
small pincer
Third walking leg 2 segments plus gill 5 segments plus Absent Walking and prehension
(epipod); genital small pincer
pore in female
Fourth walking leg 2 segments plus gill (epipod) 5 segments, Absent Walking
no pincer
Fifth walking leg 2 segments; genital 5 segments, Absent Walking
pore in male; no gill no pincer
First swimmeret In female reduced or absent; In male, transferring
in male fused with sperm to female
endopod to form tube
Second swimmeret
Male Structure modified Structure modified
for transfer of for transfer of
sperm to female sperm to female
Female 2 segments Jointed filament Jointed filament Creating water currents;
carrying eggs and young
Third, fourth, 2 short segments Jointed filament Jointed filament Creating water currents;
and fifth in female carrying eggs
swimmerets and young
Uropod 1 short, broad segment Flat, oval plate Flat, oval plate; Swimming; egg protection
divided into 2 parts in female
with hinge

fluid, as there is in animals with closed systems. Hemolymph in the arteries prevent a backflow of hemolymph. Small arteries
(blood) leaves the heart through arteries, circulates through the empty into tissue sinuses, which in turn often discharge into a
hemocoel, and returns to venous sinuses, or spaces, instead of large sternal sinus (Figure 20.7).
veins before it enters the heart. From there, afferent sinus channels carry hemolymph to the
A dorsal heart is the chief propulsive organ. It is a single- gills, if present, for oxygen and carbon dioxide exchange. Hemo-
chambered sac of striated muscle. Hemolymph enters the heart lymph then returns to the pericardial sinus by efferent channels
from the surrounding pericardial sinus through paired ostia, (Figure 20.7).
with valves that prevent backflow into the sinus (Figure 20.7). Hemolymph in arthropods may be colorless, reddish, or
From the heart hemolymph enters one or more arteries. Valves bluish, as in many Crustacea. Hemocyanin, a copper-containing

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 426 PART THREE Diversity of Animal Life

Cephalothorax Abdomen

Antenna Antennule Rostrum Brain Heart Pericardium

(supraesophageal
Ostium Testis Intestine
ganglion)
Flexor muscles
Eye
Stomach

Uropod

Figure 20.6
Internal structure of a
male crayfish. Antennal Mouth Subesophageal Digestive Ganglion Vas Copulatory Swimmeret Anus Telson
gland ganglion gland deferens swimmeret

Heart in this group. Crustaceans do not have Malpighian tubules, the

Ostium
Pericardium excretory organs of spiders and insects.
Pericardial The end sac of the antennal gland consists of a small vesi-
sinus
cle (saccule) and a spongy mass called a labyrinth. The laby-
Gonad rinth connects by an excretory tubule to a dorsal bladder,
which opens to the exterior by a pore on the ventral surface of
the basal antennal segment (Figure 20.8). Hydrostatic pressure
Gut within the hemocoel provides force for filtration of fluid into the
end sac. Filtrate is excreted as urine after the resorption of salts,
amino acids, glucose, and some water.
Excretion of nitrogenous wastes (mostly ammonia) takes place
Sternal by diffusion across thin areas of cuticle, especially gills. The so-
artery called “excretory organs” function principally to regulate ionic and
Gill chamber osmotic composition of body fluids. Freshwater crustaceans, such
Sternal
sinus Edge of
as crayfishes, are constantly threatened with dilution of their blood
carapace

Nerve cord Digestive gland

Coxa

Figure 20.7
Diagrammatical cross section through heart region of a crayfish
showing direction of blood flow in this “open” blood system. Heart Bladder
pumps blood to body tissues through arteries, which empty into tissue
sinuses. Returning blood enters sternal sinus, then goes through gills Labyrinth
for gas exchange, and finally back to pericardial sinus by efferent
channels. Note absence of veins.

respiratory pigment, or hemoglobin, an iron-containing pig-

ment, may be carried in solution. Hemolymph has the prop- Bladder
erty of clotting, which prevents its loss in minor injuries. Some
ameboid cells release a thrombinlike coagulant that precipi-
tates clotting. Tubule

End sac
Excretory System Excretory organs of adult crustaceans are
a pair of tubular structures located in the ventral part of their
Labyrinth
head anterior to the esophagus (Figure 20.6). They are called
antennal glands or maxillary glands, depending on whether Figure 20.8
they open at the base of the antennae or at the base of the Scheme of antennal gland (green gland) of crayfishes. (In natural
second maxillae. A few adult crustaceans have both. Excretory position organ is much folded.) Some crustaceans lack a labyrinth, and
organs of decapods are antennal glands, also called green glands the excretory tubule (nephridial canal) is a much-coiled tube.

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 www.mhhe.com/hickmanipz14e CHAPTER 20 Crustaceans 427

by water, which diffuses across the gills and other water-permeable Light rays
surfaces. Antennal glands, by forming a dilute, low-salt urine, act
as an effective “flood-control” device. Some Na
and Cl are lost
in the urine, but this loss is compensated by active absorption of
dissolved salt by the gills. In marine crustaceans, such as lobsters Cornea
and crabs, the antennal glands functions to adjust salt composition
Crystalline
of hemolymph by selective modification of salt content from urine. cone
In these forms urine remains isosmotic to the blood. Distal retinal
pigment cells
Nervous and Sensory Systems Nervous systems of crus-
taceans and annelids have much in common, although those Reflecting
pigment cells
of crustaceans have more fusion of ganglia (Figure 20.6). The
brain consists of a pair of supraesophageal ganglia that sup- Photoreceptor
plies nerves to the eyes and two pairs of antennae. It is joined by cells
Proximal retinal
connectives to the subesophageal ganglion, a fusion of at least pigment cells
five pairs of ganglia that supply nerves to mouth, appendages, Nerves
esophagus, and antennal glands. The double ventral nerve cord Day-adapted Night-adapted
has a pair of ganglia for each segment and nerves serving the
appendages, muscles, and other parts. In addition to this central Figure 20.9
Portion of compound eye of an arthropod showing migration of
system, there may be a sympathetic nervous system associated
pigment in ommatidia for day and night vision. Five ommatidia
with the digestive tract. are represented in each diagram. In daytime each ommatidium is
Crustaceans have well-developed sense organs. The largest surrounded by a dark pigment collar so that each ommatidium is
sense organs of crayfishes are eyes and statocysts. Widely dis- stimulated only by light rays that enter its own cornea (mosaic vision);
tributed over the body are tactile hairs, delicate projections of in nighttime, pigment forms incomplete collars and light rays can
spread to adjacent ommatidia (continuous, or superposition, image).
cuticle that are especially abundant on chelae, mouthparts, and
telson. Chemical senses of taste and smell reside in receptors on
antennae, mouthparts, and other places. of the field of vision (a mosaic, or apposition, image). In dim
A saclike statocyst, opening to the surface by a dorsal pore, light distal and proximal pigments separate so that light rays, with
is found on the basal segment of each first antenna of crayfishes. the aid of reflecting pigment cells, have a chance to spread to
Statocysts contain a ridge that bears sensory setae formed from adjacent ommatidia and to form a continuous, or superposition,
the chitinous lining and grains of sand that serve as statoliths. image. This second type of vision is less precise but takes maxi-
Whenever the animal changes its position, corresponding changes mum advantage of the limited amount of light received.
in the position of the grains on the sensory setae are relayed as
stimuli to the brain, and the animal can adjust itself accordingly.
Each molt (ecdysis) of cuticle results in loss of the cuticular lining
Reproduction, Life Cycles,
of statocysts and the sand grains that they contain. New grains and Endocrine Function
are acquired through the dorsal pore after ecdysis. Most crustaceans have separate sexes, and there are various spe-
Eyes in many crustaceans are compound, composed of many cializations for copulation among different groups. Barnacles are
photoreceptor units called ommatidia (Figure 20.9). Covering the monoecious but generally practice cross-fertilization. In some
rounded surface of each eye is a transparent area of cuticle, the ostracods and harpacticoid copepods males are scarce, and repro-
cornea, which is divided into many small squares or hexagons duction is usually parthenogenetic. Most crustaceans brood their
known as facets. These facets form the outer faces of the omma- eggs in some manner: branchiopods and barnacles have special
tidia. Each ommatidium behaves like a tiny eye and contains sev- brood chambers, copepods have brood sacs attached to the sides
eral kinds of cells arranged in a columnar fashion (Figure 20.9). of their abdomen (see Figure 20.16), and many malacostracans
Black pigment cells are found between adjacent ommatidia and carry eggs and young attached to their abdominal appendages.
the movement of pigment in an arthropod compound eye permits Crayfishes have direct development: there is no larval form.
it to adjust to different amounts of light. There are three sets of A tiny juvenile with the same form as the adult and a complete
pigment cells in each ommatidium: distal retinal, proximal retinal, set of appendages and segments hatches from the egg. However,
and reflecting; these are so arranged that they can form a collar development is indirect in most crustaceans, and larvae quite
or sleeve around each ommatidium. For strong light (day adap- unlike adults in structure and appearance hatch from eggs. Change
tation) the distal retinal pigment moves inward and meets the from larva ultimately to an adult is called metamorphosis.
outward-moving proximal retinal pigment so that a complete pig- The ancestral and most widely occurring larva in Crustacea is the
ment sleeve forms around the ommatidium (Figure 20.9). In this nauplius (Figures 20.10 and 20.22). Nauplii bear only three pairs
condition only rays that strike the cornea directly will reach the of appendages: uniramous first antennules, biramous antennae,
photoreceptor cells (retinuli), for each ommatidium is shielded and biramous mandibles. All function as swimming appendages
from others. Thus each ommatidium will see only a limited area at this stage. Subsequent development may involve a gradual

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 428 PART THREE Diversity of Animal Life

Intermolt condition Epicuticle

Exocuticle

Egg
Endocuticle

Adult Epidermis
Nauplius

STEP 1:
In the premolt stage,
the old procuticle
separates from the
epidermis, which
secretes a new
Protozoea epicuticle. New epicuticle

Postlarval
stage

STEP 2:
Mysis Still in the premolt
stage, new exocuticle Dissolving
is secreted as molting endocuticle
Figure 20.10 fluid dissolves the old
Life cycle of a Gulf shrimp, Farfantepenaeus. Penaeids spawn at endocuticle. Solution New
depths of 40 to 90 m. Young larval forms are planktonic and move products are exocuticle
inshore to water of lower salinity to develop as benthic juveniles and reabsorbed.
adults. Older shrimp return to deeper water offshore.

change to adult body form, and appendages and segments are

Discarded old
added through a series of molts. However, transformation to the epicuticle and
adult form may involve more abrupt changes. For example, meta- exocuticle
STEP 3:
morphosis of a barnacle proceeds from a free-swimming nauplius At ecdysis, the old
to a larva with a bivalve carapace called a cyprid and finally to a epicuticle and
sessile adult with calcareous plates. exocuticle are
discarded.

Molting and Ecdysis Molting, the physiological process of

making a larger cuticle, and ecdysis (ekdı̄-sis) (Gr. ekdyein, to
strip off), the shedding of the cuticle, are necessary for the body
to increase in size because the exoskeleton is nonliving and does
not grow as the animal grows. Much of a crustacean’s functioning,
including its reproduction, behavior, and many metabolic pro- STEP 4:
cesses, is directly affected by the physiology of the molting cycle. In postecdysis, new
Cuticle, which is secreted by underlying epidermis, has sev- cuticle is stretched
and unfolded, and
eral layers (Figures 19.3 and 20.11). The outermost is epicuticle, endocuticle is secreted. New
a very thin layer of lipid-impregnated protein. The bulk of cuti- endocuticle
cle is the several layers of procuticle: (1) exocuticle, which lies
just beneath the epicuticle and contains protein, calcium salts,
and chitin; (2) endocuticle, which itself is composed of (3) a
principal layer, which contains more chitin and less protein
and is heavily calcified, and (4) an uncalcified membranous Figure 20.11
layer, a relatively thin layer of chitin and protein. Cuticle secretion and resorption in ecdysis.
Molting animals grow in the intermolt phases, or instars, with
soft tissues increasing in size until there is no free space within the During the molting process and some time before actual
cuticle. When the body fills the cuticle, the animal enters the pre- ecdysis, epidermal cells enlarge considerably. They separate
molt phase. Growth occurs over a much longer time period than from the membranous layer, secrete a new epicuticle, and begin
is apparent from examining the external size of the animal. secreting a new exocuticle (Figure 20.11). Enzymes are released

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 www.mhhe.com/hickmanipz14e CHAPTER 20 Crustaceans 429

into the area above the new epicuticle. These enzymes begin to they are homologous to prothoracic glands of insects, which
dissolve old endocuticle, and soluble products are resorbed and produce the hormone ecdysone. Action of molting hormone is
stored within the body of the crustacean. Some calcium salts to initiate processes leading to ecdysis. Once initiated, the cycle
are stored as gastroliths (mineral accretions) in the walls of proceeds automatically without further action of hormones from
the stomach. Finally, only exocuticle and epicuticle of the old either X- or Y-organs.
cuticle remain, underlain by new epicuticle and new exocuti-
cle. The animal swallows water, which it absorbs through its Other Endocrine Functions Removal of eyestalks acceler-
gut, and its blood volume increases greatly. Internal pressure ates molting; in addition, crustaceans whose eyestalks have been
causes the cuticle to split along preformed lines of weakness in removed can no longer adjust their body coloration to match
the cuticle, and the animal pulls itself out of its old exoskeleton the background conditions. It was discovered long ago that this
(Figure 20.12). Following this is a stretching of the still soft new defect was caused not by loss of vision but by loss of hormones
cuticle, deposition of new endocuticle, redeposition of salvaged in the eyestalks. The body color of crustaceans is largely a result
inorganic salts and other constituents, and hardening of the new of pigments in special branched cells (chromatophores) in the
cuticle. During the period of molting, the animal is defenseless epidermis. Concentration of pigment granules in the center of
and remains hidden and quiescent. the cells causes lightening, and dispersal of pigment throughout
When a crustacean is young, ecdysis must occur frequently the cells causes darkening. Pigment behavior is controlled by
to allow growth, and the molting cycle is relatively short. As the hormones from neurosecretory cells in the eyestalk, as is migra-
animal approaches maturity, intermolt periods become progres- tion of retinal pigment for light and dark adaptation in the eyes
sively longer, and in some species molting ceases entirely. Dur- (see Figure 20.9).
ing intermolt periods, increase in tissue mass occurs as living
tissue replaces water.
Neurosecretory cells are nerve cells that are modified for secretion
of hormones. They are widespread in invertebrates and also occur
Hormonal Control of the Ecdysis Cycle Although
in vertebrates. Cells in the vertebrate hypothalamus and posterior
ecdysis is hormonally controlled, the cycle is often initiated by
pituitary are good examples (see p. 759).
environmental stimuli perceived by the central nervous system.
Such stimuli may include temperature, day length, and humidity
(in the case of land crabs) or a combination of environmental Release of neurosecretory material from the pericardial
signals. The signal from the central nervous system decreases organs in the wall of the pericardium causes an increase in the
production of a molt-inhibiting hormone by the X-organ. rate and amplitude of the heartbeat.
(The X-organ is a group of neurosecretory cells in the medulla Androgenic glands, first discovered in an amphipod
terminalis of the brain.) In crayfishes and other decapods, the (Orchestia, a common beach hopper), occur in male malacos-
medulla terminalis is found in the eyestalk. The hormone is car- tracans. Unlike most other endocrine organs in crustaceans,
ried in axons of the X-organ to the sinus gland (which itself is these are not neurosecretory organs. Their secretion stimulates
probably not glandular in function), also in the eyestalk, where expression of male sexual characteristics. Young malacostracans
it is released into the hemolymph. have rudimentary androgenic glands, but in females these glands
A drop in level of molt-inhibiting hormone promotes release fail to develop. If they are artificially implanted in a female, her
of a molting hormone from the Y-organs. Y-organs lie beneath ovaries transform to testes and begin to produce sperm, and
the epidermis near the adductor muscles of the mandibles, and her appendages begin to acquire male characteristics at the next

Rupture of membrane between Old carapace

abdomen and carapace separates and rises

Old

New

Left
cheliped Abdomen
emerging
Carapace Abdomen
A B C

Figure 20.12
Molting sequence in a lobster, Homarus americanus. A, Membrane between carapace and abdomen ruptures, and carapace begins slow elevation.
This step may take up to 2 hours. B and C, Head, thorax, and finally abdomen withdraw. This process usually takes no more than 15 minutes.
Immediately after ecdysis, chelipeds are desiccated and body is very soft. Lobster continues rapid absorption of water so that within 12 hours the
body increases about 20% in length and 50% in weight. Tissue water will be replaced by protein in succeeding weeks.

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 430 PART THREE Diversity of Animal Life

molt. In isopods the androgenic glands are found in testes; in all in the second part; food particles then pass into the intestine
other malacostracans they are between muscles of the coxopods for chemical digestion.
of the last thoracic legs and partly attached near ends of the vasa
deferentia. Although females do not possess organs similar to
androgenic glands, their ovaries produce one or two hormones
A BRIEF SURVEY OF CRUSTACEANS
that influence secondary sexual characteristics. Crustaceans are an extensive group of over 67,000 species world-
Hormones that influence other body processes in Crustacea wide with many subdivisions. They have many structures, habi-
also may be present, and evidence suggests that a neurosecre- tats, and modes of living. Some are much larger than crayfishes;
tory substance produced in the eyestalk regulates the level of others are smaller, even microscopic. Some are highly developed
blood sugar. and specialized; others have simpler organization.
Readers should realize that the following summary of crus-
taceans and the classification on page 438 are misleadingly brief.
Feeding Habits Although we mention all classes, a complete presentation of taxa
Feeding habits and adaptations for feeding vary greatly among in the hierarchy below class level would require coverage well
crustaceans. Many forms can shift from one type of feeding to beyond the scope of this textbook.
another depending on environment and food availability, but
all use the same fundamental set of mouthparts. Mandibles and
maxillae function to ingest food; maxillipeds hold and crush Class Remipedia
food. Walking legs, particularly chelipeds, serve in food capture Remipedia (Figure 20.14A) is a very small class of Crustacea.
in predaceous forms. The 10 species described so far have come from caves with
Many crustaceans, both large and small, are predatory, and connections to the sea. Remipedes have some ancestral crusta-
some have interesting adaptations for killing prey. For example, cean features. There are 25 to 38 trunk segments (thorax and
one kind of mantis shrimp carries, on one of its walking legs, a abdomen), all bearing paired, biramous, swimming append-
specialized digit that can be drawn into a groove and released sud- ages that are essentially alike. Antennules are biramous. Both
denly to pierce passing prey. Pistol shrimps (Alpheus spp.) have pairs of maxillae and a pair of maxillipeds, however, are pre-
an enormously enlarged chela that can be cocked like the hammer hensile and apparently adapted for feeding. The shape of the
of a gun and snapped shut at great speed, forming a cavitation swimming appendages is similar to that found in Copepoda,
bubble that implodes with a force sufficient to stun their prey. but unlike copepods and cephalocarids, swimming legs are
Food sources of suspension feeders range from plankton directed laterally rather than ventrally.
and detritus to bacteria. Predators consume larvae, worms, crus-
taceans, snails, and fishes. Scavengers eat dead animal and plant
matter. Suspension feeders, such as fairy shrimps, water fleas,
and barnacles, use their legs, which bear a thick fringe of setae,
to create water currents that sweep food particles through the
setae. Mud shrimps (Upogebia spp.) use long setae on their first
two pairs of thoracic appendages to strain food from water circu- A Remipede
lated through their burrow by movements of their swimmerets.
Crayfi shes have a two-part stomach ( Figure 20.13 ). The
first part contains a gastric mill in which food, already shred
by their mandibles, can be further ground by three calcareous
teeth into particles fine enough to pass through a filter of setae

Lateral teeth
of gastric mill Dorsal tooth Pyloric stomach

Cardiac Dorsal cecum

stomach Intestine

Ventral cecum B Cephalocarid

Gastrolith
Filtering setae
Esophagus Ventral tooth

Figure 20.13
Malacostracan stomach showing gastric “mill” and directions of food Figure 20.14
movements. Mill has chitinous ridges, or teeth, for mastication, and A, A remipede crustacean of class Remipedia.
setae for straining food before it passes into the pyloric stomach. B, A cephalocarid crustacean of class Cephalocarida.

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 www.mhhe.com/hickmanipz14e CHAPTER 20 Crustaceans 431

Class Cephalocarida produced, and eggs that must be fertilized are produced by
normal meiosis (production of overwintering, fertilized eggs is
Cephalocarida (Figure 20.14B) is also a small group, with only termed ephipia). Fertilized eggs are highly resistant to cold and
nine species known. Cephalocarids occur along both coasts of desiccation, and they are very important to the survival of the
the United States, in the West Indies, and in Japan. They are 2 to overwintering population and for passive transfer to new habi-
3 mm long and live in bottom sediments from the intertidal zone tats. Most cladocerans have direct development, whereas other
to a depth of 300 m. Some features are ancestral: thoracic limbs branchiopods have gradual metamorphosis.
are very similar to each other, and second maxillae are simi-
lar to thoracic limbs. Cephalocarids have no eyes, carapace, or
abdominal appendages. True hermaphrodites, they are unique
among Arthropoda in discharging both eggs and sperm through
Class Ostracoda
a common duct. Members of Ostracoda are, like diplostracans, enclosed in a
bivalve carapace and resemble tiny clams, 0.25 to 10 mm long
(Figure 20.16A). They are commonly called mussel shrimp or
Class Branchiopoda seed shrimp; they have a worldwide distribution and are impor-
tant in aquatic food webs. Ostracods show considerable fusion
There are over 10,000 species of Branchiopoda, which represent of trunk segments, obscuring division between the thorax and
a crustacean type with some ancestral characters. Three orders abdomen. The trunk has one to three pairs of limbs, with the
are recognized: Anostraca (fairy shrimp and brine shrimp, number of thoracic appendages reduced to two or none. Feed-
Figure 20.15B), with no carapace; Notostraca (tadpole shrimp, ing and locomotion are principally by use of the head append-
Figure 20.15A), whose carapace forms a large dorsal shield; and ages. Most ostracods are benthic or climb on plants, but some
Diplostraca (water fleas, Figure 20.15C), which typically have are planktonic or burrowing, and a few are parasitic. Feeding
a carapace that encloses the body but not the head, or a cara- habits are diverse; there are particle, plant, and carrion feed-
pace that encloses the entire body (clam shrimps). Branchiopods ers and predators. They are widespread in both marine and
have flattened and leaflike phyllopodia—legs that serve as the freshwater habitats. Most of the 6,000 known species are dioe-
chief respiratory organs (hence the name branchiopods). Most cious, but some are parthenogenetic. Some bizarre male mussel
branchiopods also use their legs for suspension feeding, and in shrimps emit light and may synchronize their flashing to attract
groups other than cladocerans, they use their legs for locomo- females. Development is by gradual metamorphosis. There are
tion as well. thousands of extant species and over 10,000 ostracod fossil
Most branchiopods are freshwater forms. Most important
and abundant are water fleas (cladocerans), which often form
a large proportion of freshwater zooplankton. Their interesting
means of reproduction is reminiscent of that of some rotifers
(see Chapter 15). During summer cladocerans often produce
only females, by parthenogenesis, rapidly increasing the popu-
lation. With onset of unfavorable conditions, some males are

A Ostracod

B Fairy shrimp
(order Anostraca)
B Mystacocarid

A Tadpole shrimp C Copepod Figure 20.16

(order Notostraca) C Daphnia A, An ostracod of class Ostracoda.
(order Diplostraca, B, A mystacocarid crustacean
suborder Cladocera) of subclass Mystacocarida, class
Maxillopoda. C, A copepod
Figure 20.15 with attached ovisacs; subclass
Animals in A, B, and C are members of class Branchiopoda. Copepoda, class Maxillopoda.

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 432 PART THREE Diversity of Animal Life

species whose presence in certain rock strata often serve as

important indicators of oil deposits.

Class Maxillopoda
Class Maxillopoda (10,000 species worldwide) includes a number
of crustacean groups traditionally considered classes that form a
monophyletic group within Crustacea. They basically have five
cephalic, six thoracic, and usually four abdominal segments plus Figure 20.17
a telson, but reductions of the above are common. There are no A tantulocarid. This curious little parasite is
typical appendages on the abdomen. The eye of the nauplius shown attached to the first antenna of its
(when present) has a unique structure and is called a maxil- copepod host at left; subclass Tantulocarida,
lopodan eye. class Maxillopoda.

marine invertebrates and marine and freshwater fish, and can

Subclass Mystacocarida
be of economic importance. Some species of free-living cope-
Mystacocarida is a class of tiny crustaceans (less than 0.5 mm pods serve as intermediate hosts for parasites of humans, such as
long) that live in interstitial water between sand grains of marine Diphyllobothrium (a tapeworm) and Dracunculus (a nematode),
beaches (Figure 20.16B). Only 10 species have been described, as well as of other animals.
but mystacocarids are widely distributed through many parts of Development in copepods is indirect, and some highly
the world. modified parasites show striking metamorphoses.

Subclass Copepoda Subclass Tantulocarida

This group is third only to Malacostraca in number of species, Tantulocarida (Figure 20.17) is the most recently described class
and their collective biomass exceeds billions of metric tons (here considered a subclass) of crustaceans (1983). Only about
throughout the marine and fresh waters of the world. Copepods 12 species are known so far. They are tiny (0.15 to 0.2 mm) cope-
are small (usually a few millimeters or less in length) and rather pod-like ectoparasites of other deep-sea benthic crustaceans.
elongate, tapering toward the posterior. They lack a carapace They have no recognizable head appendages except one pair of
and retain a simple, median, nauplius (maxillopodan) eye in antennae on sexual females. Their life cycle is not known with
adults (Figure 20.16C). They have a single pair of uniramous certainty, but present evidence suggests that there is a parthe-
maxillipeds and four pairs of rather flattened, biramous, thoracic nogenetic cycle and a bisexual cycle with fertilization. Tantulus
swimming appendages. The fifth pair of legs is reduced. The larvae penetrate the cuticle of their hosts by a mouth tube. Then
posterior part of the body is usually separated from the anterior, their abdomen and all thoracic limbs are lost during metamor-
appendage-bearing portion by a major articulation. Antennules phosis to an adult. Alone among maxillopodans, juveniles bear
are often longer than other appendages and used in swimming. six to seven abdominal segments, but other evidence supports
Copepoda have become very diverse and evolutionarily enter- inclusion in this class.
prising, with large numbers of symbiotic as well as free-living
species. Many parasites are highly modified, and adults may be
so highly modified (and may depart so far from the description Subclass Branchiura
just given) that they can hardly be recognized as arthropods, let Branchiurans are a small group of primarily fish ectoparasites
alone crustaceans. whose mouthparts are modified for sucking ( Figure 20.18 ).
Ecologically, free-living copepods are of extreme importance, Members of this group are usually between 5 and 10 mm long
often dominating the primary consumer level (p. 834) in aquatic and may be found on marine or fresh-
communities. In many marine localities the copepod Calanus is water fish. They typically have a broad,
the most abundant organism in zooplankton and has the greatest shieldlike carapace, compound eyes,
proportion of total biomass (p. 835). In other localities it may be four biramous thoracic appendages for
surpassed in biomass only by euphausids (p. 436). Calanus is swimming, and a short, unsegmented
an important dietary component of such economically and eco- abdomen. Second maxillae have become
logically important fish as herring, menhaden, and sardines. This modified as suction cups, enabling the
genus is also important to the larvae of larger fish and (along parasites to move on their fish host or
with euphausids) is an important food item for some whales and even from fish to fish. Heavily infested
sharks that are filter feeders. Other genera commonly occur in fish may get fungal infections and die. Figure 20.18
marine zooplankton, and some forms such as Cyclops and Diap- There is no nauplius, and young resem- Fish louse; subclass
tomus may form an important component of freshwater plankton. ble adults except in size and degree of Branchiura, class
Many species of copepods are parasites of a wide variety of other development of appendages. Maxillopoda.

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 www.mhhe.com/hickmanipz14e CHAPTER 20 Crustaceans 433

Subclass Pentastomida Figure 20.20

Members of former phylum Pentastomida (pen-ta-stomi-da) Anterior end of a
pentastome. Note
(Gr. pente, five,
stoma, mouth), or tongue worms, consist
both the mouth
of about 130 species of wormlike parasites of the respiratory (arrow), between the
system of vertebrates. Adult pentastomids live mostly in lungs middle hooks, and
of reptiles, such as snakes, lizards, and crocodiles, but one spe- the apical sensory
cies, Reighardia sternae, lives in air sacs of terns and gulls, and papillae.
another, Linguatula serrata (Gr. lingua, tongue), lives in the
nasopharynx of canines and felines (and occasionally humans).
Although more common in tropical areas, they also occur in
North America, Europe, and Australia.
Adults range from 1 to 13 cm in length. Transverse rings give
their bodies a segmented appearance (Figure 20.19). Their body
is covered with a nonchitinous and highly porous cuticle that is
molted periodically during larval stages. The anterior end may nymphs. After growth and several molts, a nymph finally becomes
bear five short protuberances (hence the name Pentastomida). encapsulated and dormant. When eaten by a final host, a juvenile
Four of these bear chitinous claws, and the fifth bears the mouth finds its way to a lung, feeds on blood and tissue, and matures.
( Figure 20.20 ). There is a simple straight digestive system, Several species have been found encysted in humans, the
adapted for sucking blood from the host. The nervous system, most common being Armillifer armillatus (L. armilla, ring,
similar to that other arthropods, has paired ganglia along the bracelet,
fero, to bear), but usually they cause few symptoms.
ventral nerve cord. The only sense organs appear to be papillae. Linguatula serrata is a cause of nasopharyngeal pentastomiasis,
There are no circulatory, excretory, or respiratory organs. or “halzoun,” a disease of humans in the Middle East and India.
Sexes are separate, and females are usually larger than males. A
female may produce several million eggs, which pass up the trachea Subclass Cirripedia
of the host, are swallowed, and exit with feces. Larvae hatch as oval,
tailed creatures with four stumpy legs. Most pentastomid life cycles Cirripedia includes barnacles (order Thoracica), which are usu-
require an intermediate vertebrate host such as a fish, a reptile, or, ally enclosed in a shell of calcareous plates, as well as three
rarely, a mammal, that is eaten by the definitive vertebrate host. smaller orders of burrowing or parasitic forms. Barnacles are ses-
After ingestion by an intermediate host, larvae penetrate the intes- sile as adults and may be attached to the substrate by a stalk
tine, migrate randomly in the body, and finally metamorphose into (gooseneck barnacles) (Figure 20.21B) or directly (acorn bar-
nacles) (Figure 20.21A). Typically their carapace (mantle) sur-
rounds their body and secretes a shell of calcareous plates. The
Hooks Mouth head is reduced, they have no abdomen, and the thoracic legs
are long, many-jointed cirri with hairlike setae. The cirri are
extended through an opening between the calcareous plates to
filter out small particles on which the animal feeds (Figure 20.21).
Enteron Although all barnacles are marine, they are often found in the
Seminal
receptacle intertidal zone and are therefore exposed to drying and some-
times freshwater for some periods of time. For example, Semi-
Ovary balanus balanoides can tolerate below-freezing temperatures in
the Arctic tidal zone and can survive exposed on its rocky sub-
strate for up to nine hours in the summer. During these periods
the aperture between the plates closes to a very narrow slit.

Barnacles frequently foul ship bottoms by settling and growing

Vagina
there. So great may be their number that the speed of the ship may
be reduced by 30% to 40%, necessitating drydocking the ship to
Anus remove them. They may also live atop whales (see Figure 20.26).

Most nonparasitic barnacles are hermaphroditic and undergo

Figure 20.19 a striking metamorphosis during development. Most hatch as
Two pentastomids. A, Linguatula, found in nasal passages of nauplii, which soon become cyprid larvae, so called because
carnivorous mammals. Female is shown with some internal structures.
B, Female Armillifer, a pentastomid with pronounced body rings. In
of their resemblance to an ostracod genus Cypris. They have
parts of Africa and Asia, humans are parasitized by immature stages; a bivalve carapace and compound eyes. Cyprids attach to the
adults (10 cm long or more) live in lungs of snakes. Human infection substrate by means of their first antennae, which have adhesive
may occur from eating snakes or from contaminated food or water. glands, and begin their metamorphosis. This involves several

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 434 PART THREE Diversity of Animal Life

A B

Figure 20.21
Barnacles; order Thoracica, subclass Cirripedia, class Maxillopoda. A, Acorn barnacles, Balanus balanoides, on an intertidal rock await the return of
the tide. B, Common gooseneck 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.

dramatic changes, including secretion of calcareous plates, loss Class Malacostraca

of eyes, and transformation of swimming appendages to cirri.
Members of order Rhizocephala, such as Sacculina, are Malacostraca, with over 20,000 species worldwide, is the larg-
highly modified parasites of crabs. These barnacles are dioe- est class of Crustacea and shows great diversity. The diversity is
cious. Like other cirripedes, they start life as nauplii and then reflected by the higher classification of the group, which includes
become cyprid larvae, but when they find a host, females of most three subclasses, 14 orders, and many suborders, infraorders,
species metamorphose into a kentrogon (Gr. kentron, a point, and superfamilies. We confine our coverage to a few of the most
spine,
gonos, progeny) which injects cells of the parasite into important orders. We described the characteristic body plan of
the hemocoel of its crab host (Figure 20.22). Eventually, rootlike malacostracans on page 422.
absorptive processes grow throughout the crab’s body, and the
parasite reproductive structures become externalized between Order Isopoda
cephalothorax and reflexed abdomen of the crab. Males at the
Isopods are one of the few crustacean groups to have successfully
cyprid stage attach to the external female brood chamber.
invaded terrestrial habitats in addition to freshwater and seawater
habitats and the only crustaceans to have become truly terrestrial.
The exact position at which reproductive structures become exter- They are typically dorsoventrally flattened, lack a carapace,
nalized from the crab’s body is of great adaptive value for rhizo- and have sessile compound eyes. Maxillipeds are the first pair of
cephalan parasites. Because a crab’s egg mass (if it had one) would thoracic limbs; other thoracic limbs lack exopods and are simi-
be borne in this position, a crab treats the parasite as if it were a lar. Abdominal appendages bear gills or lunglike organs called
mass of the crab’s own eggs. It protects, ventilates, and grooms pseudotracheae and, except for uropods, also are similar to each
its parasite and actually assists in the parasite’s reproduction by other (hence the name isopods). Many species have the ability
performing spawning behavior at the appropriate time. The crab’s to roll into a tight ball for protection.
grooming is necessary for continued good health of the parasite. Common land forms are the sow bugs, or pill bugs (Porcel-
But what if the rhizocephalan’s larva is so unlucky as to infect a lio and Armadillidium, Figure 20.23A), which live under stones
male crab? No problem. During the parasite’s internal growth in the and in damp places. Although they are terrestrial, they lack an
male crab, it castrates its host, and the crab becomes structurally efficient cuticular covering and other adaptations possessed by
and behaviorally like a female! insects to conserve water; therefore they must live in moist envi-
ronments (for example, under wet logs or rocks). Caecidotea

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 www.mhhe.com/hickmanipz14e CHAPTER 20 Crustaceans 435

Seta on crab Nauplius larvae

of parasite
released

Kentrogon larva
penetrates at base of
seta, injects cells

Male cyprid
enters
externalized
female

External portion
of Sacculina on
host crab
Embryonic cell
mass attaches
to midgut and
begins to grow Figure 20.22
Sacculina tissue
Life cycle of Sacculina (order
growing on crab
midgut Rhizocephala, subclass Cirripedia, class
Maxillopoda), parasite of crabs (Carcinus).

Figure 20.23
A, Four pill bugs, Armadillidium vulgare
(order Isopoda, class Malacostraca),
common terrestrial forms. B, Freshwater
sow bug, Caecidotea sp., an aquatic isopod. B
Figure 20.24
An isopod parasite (Anilocra sp.) on a coney (Cephalopholis fulvus)
inhabiting a Caribbean coral reef (order Isopoda, class Malacostraca).
(Figure 20.23B) is a common freshwater form found under rocks
and among aquatic plants. Ligia is a common marine form that
scurries across the beach or rocky shore. Some isopods are para- Order Amphipoda
sites of fishes (Figure 20.24) or crustaceans. Amphipods resemble isopods in that they lack a carapace, and
Development is essentially direct but may be highly meta- have sessile compound eyes and only one pair of maxillipeds
morphic in specialized parasites. (Figure 20.25). However, they are usually compressed laterally,

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 436 PART THREE Diversity of Animal Life

C
A B

Figure 20.25
Marine amphipods. A, Free-swimming amphipod, Anisogammarus sp. B, Skeleton shrimp, Caprella sp., shown on a bryozoan colony, resemble
praying mantids. C, Phronima, a marine pelagic amphipod, takes over the tunic of a salp (subphylum Urochordata, see Chapter 23). Swimming by
means of its abdominal swimmerets, which protrude from the opening of the barrel-shaped tunic, the amphipod maneuvers to catch its prey. The
tunic is not seen (order Amphipoda, class Malacostraca).

and their gills are in the typical thoracic position. Furthermore,

their thoracic and abdominal limbs are each arranged in two
or more groups that differ in form and function. For example,
one group of abdominal legs may be for swimming and another
group for jumping. There are many marine amphipods, includ-
ing some beach-dwelling forms (for example, Orchestia, a beach
hopper), numerous freshwater genera (Hyalella and Gamma-
rus), and a few parasites (Figure 20.26). Development is direct
and without a true metamorphosis.

Order Euphausiacea Figure 20.27

Meganyctiphanes (order Euphausiacea, class Malacostraca)
Euphausiacea is a group of only about 90 species, but they are “northern krill.”
important as oceanic plankton known as “krill” (Figure 20.27).

A B
Figure 20.26
A, Head and mouth of a healthy California grey whale, Eschrichtius robustus, bearing its characteristic heavy load of barnacles (order Thoracica,
subclass Cirripedia, class Maxillopoda) and cyamid parasites (order Amphipoda, class Malacostraca) (arrows). Note yellowish plates of baleen in mouth
(p. 640). B, Cyamid parasites of grey whale. Unlike most amphipods, these are dorsoventrally flattened. They have sharp, grasping claws on their legs.

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A B C

D E

Figure 20.28
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, A spiny lobster, Panulirus argus (shown
here), and the northern lobster, Homarus americanus, are consumed with gusto by many people (order Decapoda, class Malacostraca).

They are about 3 to 6 cm long, have a carapace that is fused with spider crabs, whose chelae span 4 m from tip to tip. Crayfishes, lob-
all thoracic segments but does not entirely enclose their gills. sters, crabs, and “true” shrimp belong in this group (Figures 20.28
They have no maxillipeds, but have thoracic limbs with exo- and 20.29). There are about 18,000 species of decapods, and the
pods. Most are bioluminescent, with a light-producing substance order is extremely diverse. They are very important ecologically
in an organ called a photophore. Some species may occur in and economically, and numerous species are relished as food.
enormous swarms, covering up to 45 m2 and extending up to
500 m in one direction. They form a major portion of the diet
of baleen whales and many fishes. Eggs hatch as nauplii, and Figure 20.29
development is indirect and metamorphic. Sponge crab,
Dromidia antillensis.
This crab is one
of several species
Order Decapoda that deliberately
Decapods have three pairs of maxillipeds and five pairs of walking mask themselves
legs. In crabs the first pair of walking legs is modified to form pin- with material from
their immediate
cers (chelae), but the second and third pairs may also be chelate, environment (order
as in crayfishes, lobsters, and most shrimps. Decapods range in size Decapoda, class
from a few millimeters to the largest of all arthropods, Japanese Malacostraca).

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 438 PART THREE Diversity of Animal Life

Classification of Subphylum Crustacea

Higher classification of Crustacea is complex and subject to change Subclass Tantulocarida (tantülōkarədə) (L. tantulus,
as new data become available. This listing relies on several sources, so small,
caris, shrimp). No recognizable cephalic
omitting many smaller taxa. appendages except antennae on sexual female; solid median
Class Remipedia (remə-pədēə) (L. remipedes, oar-footed). No cephalic stylet; six free thoracic segments, each with pair of
carapace; one-segmented protopods; biramous antennules and appendages, anterior five biramous; six abdominal segments;
antennae; all trunk appendages similar; cephalic appendages minute copepod-like ectoparasites. Examples: Basipodella,
large and raptorial; maxilliped segment fused to head; trunk Deoterthron.
unregionalized. Example: Speleonectes. Subclass Branchiura (bran-kēyurə) (Gr. branchia, gills,
Class Cephalocarida (sefə-lō karədə) (Gr. kephalē, head,
ura, tail): fish lice. Body oval, head and most of trunk

karis, shrimp,
ida, pl. suffix). No carapace; phyllopodia, covered by flattened carapace, incompletely fused to first
one-segmented protopods; uniramous antennules and biramous thoracic segment; thorax with four pairs of appendages,
antennae; compound eyes lacking; no abdominal appendages; biramous; abdomen unsegmented, bilobed; eyes compound;
maxilliped similar to thoracic leg. Example: Hutchinsoniella. antennae and antennules reduced; maxillules often forming
Class Branchiopoda (bran-kēäpōdə) (Gr. branchia, gills,
suctoral discs. Examples: Argulus, Chonopeltis.
pous, podos, foot). Phyllopodia; carapace present or absent; no Subclass Pentastomida (pen-ta-stomi-da) (Gr. pente, five,
maxillipeds; antennules reduced; compound eyes present; no
stoma, mouth): pentastomids. Wormlike unsegmented
abdominal appendages; maxillae reduced. body with five short anterior protuberances, four bear claws
Order Anostraca (ənästrəkə) (Gr. an-, prefix meaning without, and the fifth bears the sucking mouth. Examples: Armillifer,

ostrakon, shell): fairy shrimp and brine shrimp. No Linguatula.
carapace; no abdominal appendages; uniramous antennae. Subclass Cirripedia (sirə-pēdēə) (L. cirrus, curl of hair,

Examples: Artemia, Branchinecta. pes, pedis, foot): barnacles. Sessile or parasitic as adults; head
Order Notostraca (nōtästrəkə) (Gr. nōtos, the back,
reduced and abdomen rudimentary; paired compound eyes
ostrakon, shell): tadpole shrimp. Carapace forming large dorsal absent; body segmentation indistinct; usually hermaphroditic;
shield; abdominal appendages present, reduced posteriorly; in free-living forms carapace becomes mantle, which secretes
antennae vestigial. Examples: Triops, Lepidurus. calcareous plates; antennules become organs of attachment,
Order Diplostraca (diplōstrəkə) (Gr. diploos, double, then disappear. Examples: Balanus, Policipes, Sacculina.

ostrakon, shell): water fleas (cladocerans) and clam Class Malacostraca (malə-kä-strəkə) (Gr. malakos, soft,

shrimps (conchostracans). Carapace folded, usually enclosing ostrakon, shell). Usually with eight segments in thorax and six plus
trunk but not head (cladocerans) or enclosing entire body telson in abdomen; all segments with appendages; antennules often
(conchostracans); biramous antennae. Examples: Daphnia, biramous; first one to three thoracic appendages often maxillipeds;
Leptodora, Lynceus. carapace covering head and part or all of thorax, sometimes absent;
Class Ostracoda (ästrəkōdə) (Gr. ostrakodes, having a shell): gills usually thoracic epipods.
ostracods. Bivalve carapace entirely encloses body; body Order Isopoda (īso-pōdə) (Gr. isos, equal,
pous, podos,
unsegmented or indistinctly segmented; no more than two pairs of foot): isopods. No carapace; antennules usually uniramous,
trunk appendages. Examples: Cypris, Cypridina, Gigantocypris. sometimes vestigial; eyes sessile (not stalked); gills on
Class Maxillopoda (maksilapōdə) (L. maxilla, the jawbone,
abdominal appendages; body commonly dorsoventrally
pous, podos, a foot). Usually five cephalic, six thoracic, and four flattened; second thoracic appendages usually not prehensile.
abdominal segments plus a telson, but reductions common; no Examples: Armadillidium, Caecidotea, Ligia, Porcellio.
typical appendages on abdomen; naupliar eye of unique structure Order Amphipoda (am-fi-pōdə) (Gr. amphis, on both sides,
(maxillopodan eye); carapace present or absent.
pous, podos, foot): amphipods. No carapace; antennules
Subclass Mystacocarida (mista·kō-karədə) (Gr. mystax, often biramous; eyes usually sessile; gills on thoracic coxae;
mustache,
karis, shrimp,
ida, pl. suffix): mustache second and third thoracic limbs usually prehensile; typically
shrimps. No carapace; body of cephalon and ten-segmented bilaterally compressed body form. Examples: Orchestia,
trunk; telson with clawlike caudal rami; cephalic appendages Hyalella, Gammarus.
nearly identical, but antennae and mandibles biramous, other Order Euphausiacea (yü-foz-ē-āshēə) (Gr. eu, well,
phausi,
head appendages uniramous; second through fifth trunk shining bright,
L. acea, suffix, pertaining to): krill. Carapace
segments with short, single-segment appendages. Example: fused to all thoracic segments but not entirely enclosing gills,
Derocheilocaris. no maxillipeds; all thoracic limbs with exopods. Example:
Subclass Copepoda (kōpe-pədə) (Gr. kōpē, oar,
pous, Meganyctiphanes.
podos, foot): copepods. No carapace; thorax typically of seven Order Decapoda (dəka-pōdə) (Gr. deka, ten,
pous, podos,
segments, of which first and sometimes second fuse with head foot): shrimps, crabs, lobsters. All thoracic segments fused
to form cephalothorax; antennules uniramous; antennae bi- or with and covered by carapace; eyes on stalks; first three pairs
uniramous; four to five pairs swimming legs; parasitic forms of thoracic appendages modified to maxillipeds. Examples:
often highly modified. Examples: Cyclops, Diaptomus, Calanus, Farfantepenaeus (= Penaeus), Cancer, Pagurus, Grapsus,
Ergasilus, Lernaea, Salmincola, Caligus. Homarus, Panulirus.

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 www.mhhe.com/hickmanipz14e CHAPTER 20 Crustaceans 439

Crabs, especially, exist in a great variety of forms. Although Dll can be observed in two groups of cells, each of which will
resembling crayfishes, they differ from the latter in having a become a branch of the limb. In a uniramous limb primordium,
broader cephalothorax and reduced abdomen. Familiar exam- there is only one such group of cells, and in primodia of phyl-
ples along the seashore are hermit crabs (Figure 20.28B), which lopodous limbs (as in class Branchiopoda), there are as many
live in snail shells (because their abdomens are not protected by groups expressing Dll as there are limb branches.
the same heavy exoskeleton as are the anterior parts); fiddler The wormlike pentastomids were placed in Ecdysozoa
crabs, Uca (Figure 20.28C), which burrow in sand just below the near arthropods because their larval form resembles tardigrade
high-tide level and emerge to run over the sand while the tide larvae, their cuticle is molted, and there are other similarities in
is out; spider crabs such as Libinia; interesting decorator crabs sperm morphology and larval appendages. Phylogenies based
Dromidia, and others, which cover their carapaces with sponges on sequences of ribosomal RNA genes indicate that pentas-
and sea anemones for protective camouflage (Figure 20.29). tomids are crustaceans. A recent study of gene arrangements
and base sequences of mitochondrial DNA confirmed this
result. Pentastomids are now considered highly derived crus-
PHYLOGENY AND ADAPTIVE taceans, placed in class Maxillopoda near fish lice (subclass
DIVERSIFICATION Branchiura).

Phylogeny
Among Crustacea, Remipedia seem to have the most ancestral Adaptive Diversification
characters (see Figure 20.1): They have a long body, with no The level of adaptive diversification demonstrated by the crus-
tagmatization behind the head, a double ventral nerve cord, and taceans is great, with the exploitation of virtually all aquatic
serially arranged digestive ceca. Fossils of a puzzling arthropod resources. They are unquestionably the dominant arthropod
from the Mississippian period seem to be the sister group of group in marine environments, and they share dominance of
remipedians, and their morphology suggested one mechanism freshwater habitats with insects. Invasions of terrestrial environ-
for the origin of biramous appendages. They have two pairs of ments have been much more limited, with isopods being the
uniramous limbs on each segment. Thus, it was suggested that only notable success. There are a few other terrestrial examples,
each crustacean segment represents two ancestral segments that such as land crabs. The most diverse class is Malacostraca, and
fused (“diplopodous condition,” as seen in Diplopoda, p. 414), the most abundant groups are Copepoda and Ostracoda. Mem-
and that biramous appendages derived from fusion of both limbs bers of both taxa include planktonic suspension feeders and
on an ancestral diplopodous segment. However, it is now known numerous scavengers. Copepods have been particularly success-
that modulation in expression of the Distal-less (Dll) gene deter- ful as parasites of both vertebrates and invertebrates, and it is
mines location of distal ends of anthropod limbs. In each pri- clear that present parasitic copepods are products of numerous
mordial (embryonic) biramous appendage, the gene product of invasions of such niches.

SUMMARY
Crustacea is a large, primarily aquatic subphylum. In addition to a important as zooplankton. Within class Maxillopoda, members of
pair of mandibles, crustaceans have common two pairs of antennae subclass Copepoda lack a carapace and abdominal appendages.
and two pairs of maxillae. Their tagmata are a head and trunk or They are abundant and are among the most important of the pri-
a head, thorax, and abdomen. Many have a carapace. Crustaceans’ mary consumers in many freshwater and marine ecosystems. Many
appendages are ancestrally biramous. are parasitic. Most members of subclass Cirripedia (barnacles) are
All arthropods must periodically cast off their old cuticle (ecdy- sessile as adults, secrete a shell of calcareous plates, and filter-feed
sis) and grow in size before the newly secreted cuticle hardens. Pre- by means of their thoracic appendages. Subclass Branchiura con-
molt and postmolt periods are hormonally controlled, as are several tains fish lice. Closely related to fish lice are tongue worms; they
other processes, such as change in body color and expression of are parasitic in the lungs and nasal cavities of vertebrates. These
sexual characteristics. members of former phylum Pentastomida now comprise subclass
Feeding habits vary greatly in Crustacea, and there are many Pentastomida in class Maxillopoda.
forms of predators, scavengers, suspension feeders, and parasites. Malacostraca is the largest and most diverse crustacean class,
Respiration is through the body surface or by gills, and excretory and the most important orders are Isopoda, Amphipoda, Euphausia-
organs take the form of maxillary or antennal glands. Circulation, as cea, and Decapoda. All have both abdominal and thoracic append-
in other arthropods, is through an open system of sinuses (hemo- ages. Isopods lack a carapace and are usually dorsoventrally flat-
coel), and a dorsal, tubular heart is the chief pumping organ. Most tened. Amphipods also lack a carapace but are usually laterally
crustaceans have compound eyes composed of units called omma- flattened. Euphausiaceans are important oceanic plankton called
tidia. Sexes are usually separate. krill. Decapods include crabs, shrimps, lobsters, crayfishes, and oth-
Class Branchiopoda is characterized by phyllopodia and ers; they have five pairs of walking legs (including chelipeds) on
contains, among others, order Diplostraca, which is ecologically their thorax.

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 440 PART THREE Diversity of Animal Life

SELECTED REFERENCES
Bliss, D. E. (editor-in-chief). 1982–1985. The biology of Crustacea, vols. 1– hypothesis of a parthenogenetic cycle alternating with a cycle that
10. New York, Academic Press, Inc. This series is a standard reference includes fertilization in these bizarre little creatures.
for all aspects of crustacean biology. Lavrov, D. L., W. M. Brown, and J. L. Boore. 2004. Phylogenetic position
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gene translocation, indicative of a recent common ancestor, is shared probably closely related to fish lice.
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