Phylum Arthropoda: Subphylum Crustacea
Biological Sciences 102 Animal Biology Notes & Vocabulary
Phylum Arthropoda: Subphylum Crustacea (more than 67,000 species)
See page 402 in the text for classification
Body Plan Features Characteristic of Members of the Phylum Arthropoda:
1. developed striated muscle for rapid movement
2. an exoskeleton, or cuticle, containing the tough nitrogenous polysaccharide chitin for
support and protection
3. gills for very efficient gaseous exchange in crustaceans
4. greater specialization of body organs, especially specialization of form and function
among the appendages
5. "open" circulatory system
Body
Plan Features Retained by Arthropods (seen in previously studied phyla):
bilateral symmetry
eucoelomic (true coelom)
high degree of cephalization = well developed head with sensory organs
specialized segmentation (metamerism) = tagmata
triploblastic structure (endoderm, mesoderm and ectoderm)
all organ systems are present
Adaptations of the Subphylum Crustacea (kr-st'sh) (L. crusta, shell)
Body
Plan Features and Characteristics of Members of the Subphylum Crustacea:
most crustaceans have between 16 and 20 segments some have many more
different segments in different groups may have fused to form a head or cephalothorax
head fused to thoracic body segments to form a cephalothorax
abdomen is formed of the posterior segments
telson (tail) with uropod
first walking leg is chelate (cheliped)
two pairs of antennae (one pair of antennae and one pair of antennules)
two pairs of maxillae on the head
one pair of mandibles (crustaceans are the aquatic mandibulates)
usually one pair of appendages on each body segment
all appendages except the first pair of antennae in present-day crustaceans are
biramous (two-branched)
efficient gills for respiration
body is covered with secreted cuticle of protein, chitin and calcareous material
cuticle is thinner around joints for movement
tergum is the dorsal cuticle plate on each segment not covered by carapace
sternum is the ventral cuticle on each segment
gonopore position varies usually near the 5th pair of walking legs in males & the 3rd
pair of walking legs in females; opening to the seminal receptacle is near the 4th and 5th
pairs of walking legs in females
considerable specialization of appendages has occurred in many derived groups such as
the crayfishes
mainly marine, many freshwater, a few are terrestrial
Class Malacostraca is the largest group (lobsters, crabs, shrimps, beach hoppers and
many others)
walking legs, mouthparts, chelipeds and swimmerets have all become modified for
different functions from a common biramous appendages (they are serially
homologous)
crustaceans have segmented nervous and muscular systems
Phylum Arthropoda: Subphylum Crustacea
Biological Sciences 102 Animal Biology Notes & Vocabulary
hemocoel occupies the major body space from schizocoelous coelom formation and
coelom is very reduced similar to molluscs
Appendage Parts protopod is formed of a basis and coxa; exopods and
endopods have one to several parts
protopod = basal portion
exopod = lateral part of protopod
endopod = medial part of protopod
some appendages have become secondarily uniramous (walking legs)
epipods are modified processes on a protopod that often form gills
striated muscles are arranged in antagonistic groups (eg. flexors and extensors)
small crustaceans utilize thin areas in cuticle for respiration while large
crustaceans have gills; part of the second maxillae forms a bailer that draws water
over the gills into the gill cavity at the base of the legs and out at the anterior end
open circulatory system with blood leaving the dorsal heart through short arteries
to the hemocoel and back to the heart through venous sinuses; arteries possess valves
to prevent backflow
hemolymph may be colorless, reddish or bluish due to hemocyanin (copper containing
respiratory pigment or hemoglobin (iron containing respiratory pigment); hemolymph
can clot to prevent loss in minor injuries and ameboid cells release clotting proteins
green/antennal or maxillary glands depending on where they open at are paired
tubular structures for excretion; crustaceans lack Malpighian tubules
freshwater crustaceans have green glands that form a dilute low-salt urine for
osmotic balance; some salt is lost with the urine but is replaced by salt absorption
through the gills (little nitrogenous waste is actually excreted by the green glands)
excretion of primarily ammonia occurs primarily by diffusion across thin areas of
the cuticle and gills
marine crustaceans have kidneys for salt and water balance (osmoregulation)
nervous system is composed of a pair of supraesophageal ganglia with nerves to the
eyes and two pairs of antennae which is joined to a subesophageal ganglion with
nerves to the mouth, appendages, esophagus and antennal glands; paired ventral nerve
cords with a pair of ganglia in each segment and peripheral nerves to appendages,
muscles, etc.
sense organs include: compound eyes formed of ommatidia, tactile hairs (especially
on chelae and mouthparts and telson); chemoreceptors for taste and smell on
antennae, mouthparts and other places; statocysts open on dorsal side at the base of
each first antennae (in crayfish) with chitinous lining and sand grains that act as
statoliths which are replaced after each molt
Each hexagonal ommatidia of the eye is covered by a transparent part of the cuticle =
cornea; each ommatidia acts as a tiny eye with pigment cells between them that
form a collar to separate ommatidia; in bright light, each ommatidia only sees a
limited area of the field of view to form a mosaic image; at night, distal and
proximal pigment cells can separate so light hits multiple retinal cells in multiple
ommatidia to form a superimposed continuous image that is not mosaic, but less
precise this makes the most of the limited available light at night
most crustaceans have separate sexes with specializations for copulation and ways of
brooding their eggs
barnacles are monoecious but utilize cross-fertilization with other individuals via a
very long penis (relative to body size) as sessile adults
ostracods and some copepods are often parthenogenetic
the ancestral most common type of larva in crustaceans is a nauplius larva with three
pairs of appendages, uniramous first antennules, biramous antennae and biramous
mandibles all used for swimming with gradual changes to an adult form through a
series of molts or only a fews molts (eg. barnacles)
Phylum Arthropoda: Subphylum Crustacea
Biological Sciences 102 Animal Biology Notes & Vocabulary
Molting and Ecdysis in Crustaceans
molting = the process of making a larger cuticle
ecdysis = the shedding of the old cuticle
much behavior, metabolism and reproductive physiology is tied to the molting cycle
the cuticle is formed of:
epicuticle = outermost thin layer of protein and lipid
procuticle = formed of exocuticle and endocuticle
exocuticle = beneath the epicuticle made of protein, chitin and calcium salts
endocuticle = made of prinicipal layer which is mostly chitin and calcium and less
protein and the uncalcified membranous layer of chitin and protein
Molting Cycle
1. before ecdysis the epidermal cells enlarge and separate from the membranous layer,
secrete a new epicuticle and a new exocuticle
2. enzymes are released above the new epicuticle to dissolve the old endocuticle and
soluble products are reabsorbed and stored in the body; some calcium salts are stored in
the gastroliths in the stomach
3. as the animal swallows water its blood volume increases and internal pressure splits the
cuticle along preformed weak lines (usually between the cephalothorax and abdomen) and
the animal pulls itself out of its old exoskeleton
4. the new soft cuticle is stretched, new endocuticle is deposited and new cuticle is
hardened with stored salts and other constituents
while molting, the animal is defenseless and hides
young crustaceans have shorter molting cycles than older mature crustaceans; in some
species molting ceases after sexual maturity is reached
Ecdysis is controlled by hormones
environmental stimuli such as temperature, daylength and humidity changes (for land
crabs) signal a decrease in the production of molt inhibiting hormone (MIH) from the
X-organ.
the X-organ is a group of neurosecretory cells in the medulla terminalis of the brain
which in decapods is found in the eyestalk
MIH is carried in axons of the X-organ to the sinus gland in the eyestalk and released into
the hemolymph
a decrease in MIH allows an increase in the release of molting hormone from the Yorgans that lie beneath the epidermis near the mandible adductor muscles (same as
the prothoracic glands of insects)
molting hormone stimulates the processes of ecdysis and once stimulated the process
proceeds without further hormone or glandular action
molting hormone is a steroid hormone similar to ecdysone in insects
if the eyestalks of a crustacean are removed, molting is accelerated and body
coloration can no longer match the background (no visual cues for camouflage via
chromatophores in the epidermis = also hormonally controlled)
some malacostracans have androgenic glands (in testes or near last thoracic legs) that
regulate male sexual characteristics
Feeding Habits of Crustaceans
much variation in feeding habits of crustaceans and some can shift feeding strategies (eg.
pistol shrimp use their enlarged chela to snap shut very fast and form a cavitation bubble
that can stun prey)
chelipeds are used in prey capture and mandibles and maxillae and other mouthparts used
in ingesting food
crustaceans can be suspension feeders, predators or scavengers
Phylum Arthropoda: Subphylum Crustacea
Biological Sciences 102 Animal Biology Notes & Vocabulary
Special Adaptations within the Subphylum Crustacea
Class Malacostraca (about 20,000 species)
below is the basic body plan of an ancestral malacostracan which is typical of
most members of this class seen today
head with five fused segments fused to a thorax of eight segments
abdomen has 6 (sometimes 7) segments
anterior end is a non-segmented rostrum
posterior end is a non-segmented telson with uropods that form the fan tail
carapace is the dorsal cuticle of the head extended posteriorly and around the sides to
cover the thoracic and abdominal segments (in decapods, all of the cephalothorax, but
none of the abdominal segments are covered)
see table 19.1 in the text for specifics and functions of crayfish appendages
crayfish have direct development with no larval form that is seen in many other
crustaceans
crayfish have two-portioned stomach: first part has gastric mill with three
calcareous teeth for grinding food then through filtering setae to the second part before
entering the intestines
Order
Decapoda (shrimps, lobsters, crabs, crayfish = about 18,000 species)
freshwater or marine
have a carapace
compound eyes on eyestalks
three pairs of maxillipeds
five pairs of walking legs
five pairs of swimmerets
first walking legs is often chelated (has pincers)
few millimeters to meters (Japanese spider crab has 4 m chelae)
Order
Isopoda (isopods)
freshwater, marine and only truly terrestrial crustaceans
dorsoventrally flattened
lack carapace
compound eyes with no eyestalk
one pair of maxillipeds
thoracic limbs lack exopods
abdominal appendages bear gills or lunglike pseudotracheae
many can roll into a ball for protection
direct development with some metamorphosis in parasitic forms
Order
Amphipoda (eg. beach hoppers)
freshwater and marine (eg. beach hoppers)
laterally flattened
lack carapace
compound eyes with no eyestalk
one pair of maxillipeds
gills in thorax
thoracic and abdominal limps arranged in two or more groups for different
functions (eg. one for swimming and one for jumping)
direct development
Phylum Arthropoda: Subphylum Crustacea
Biological Sciences 102 Animal Biology Notes & Vocabulary
Order
Euphausiacea (krill)
important part of marine plantkton
about 3 to 6 cm long
have a carapace fused with all thoracic segments
no maxillipeds
gills not entirely covered by carapace
most are bioluminescent
can swarm in large masses in the ocean
major portion of the diet of baleen whales
indirect development (metamorphosis from larval nauplii)
Class Branchipoda (fairy shrimp, brine shrimp, tadpole shrimp, water fleas)
with or without carapace
flattened leaflike phyllopodia = legs for respiration that are often used for filter
feeding
mostly freshwater
water fleas (eg. Daphnia) form a large part of the freshwater zooplankton
reproduction often by parthenogenesis in the summer and sexual before overwintering
fertilized eggs are produced
gradual metamorphosis or direct development
Class
Ostracoda (mussel shrimp or seed shrimp)
look like tiny clams
feed and move using head appendages
most live on bottom or on plants, some burrow, some parasitic
particle and carrion feeders, herbivores or predatory
mostly dioecious, some parthenogenetic
gradual metamorphosis
males sometimes emit light to attract females
Class Maxillopoda (Mystacorarida, Copepoda, Tantulocarida, Branchiura, Cirripedia)
this may be a polyphyletic group divided into individual classes
Subclass or Class Copepoda (copepods)
comprise a large and ecologically important portion of the marine and
freshwater zooplankton
usually only a few millimeters or less long
often paddle shaped tapering from anterior to posterior
lack a carapace
simple, single median eye
single pair of antennules
single pair of maxillipeds
four pairs of flattened, biramous thoracic swimming appendages
free-living or symbiotic (many parasitic)
indirect development
Phylum Arthropoda: Subphylum Crustacea
Biological Sciences 102 Animal Biology Notes & Vocabulary
Class Maxillopoda (Mystacorarida, Copepoda, Tantulocarida, Branchiura, Cirripedia)
Subclass or Class Cirrepedia (barnacles)
enclosed in a shell of calcareous plates
sessile as adults and may have stalk (eg. gooseneck barnacles)
carapace surrounds body and secretes plates
head is reduced, no abdomen, and thoracic legs are long cirri with hairlike
setae which are used for filter feeding
all marine and often intertidal (aperture between plates closes when exposed)
hermaphroditic with significant metamorphosis
nauplii hatch to become cyprid larvae with a bivalve carapace and
compound eyes that attach to substrate with their first antennae via adhesive
glands; calcareous plates are secreted, eyes are lost and appendages transform
during metamorphosis
Order Rhizocephalans such as Sacculina are crab parasites; start life as nauplii that
become cyprid larvae that metamorphose into a kentrogon which injects cells of the
parasite into the hemocoel of its crab host; root-like processes grow throughout the
crabs body and the parasite reproductive structures become externalzed between the
cephalothorax and abdomen
