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Phylum Arthropoda: Subphylum Crustacea

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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
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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 (krŭ-stā'shə) (L. crusta, shell)
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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

appendages. 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. each ommatidia acts as a “tiny eye” with pigment cells between them that form a collar to separate ommatidia. reddish or bluish due to hemocyanin (copper containing respiratory pigment or hemoglobin (iron containing respiratory pigment). at night. in bright light. 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. 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. 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. uniramous first antennules. esophagus and antennal glands. 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. 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. flexors and extensors) small crustaceans utilize thin areas in cuticle for respiration while large crustaceans have gills. crustaceans lack Malpighian tubules freshwater crustaceans have green glands that form a dilute low-salt urine for osmotic balance. arteries possess valves to prevent backflow hemolymph may be colorless. each ommatidia only “sees” a limited area of the field of view to form a mosaic image. paired ventral nerve cords with a pair of ganglia in each segment and peripheral nerves to appendages. sense organs include: compound eyes formed of ommatidia. tactile hairs (especially on chelae and mouthparts and telson). chemoreceptors for taste and smell on antennae. etc.Phylum Arthropoda: Subphylum Crustacea 2 Biological Sciences 102 – Animal Biology – Notes & Vocabulary ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ 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. barnacles) . muscles. mouthparts and other places. exopods and endopods have one to several parts ¾ protopod = basal portion ¾ exopod = lateral part of protopod ¾ endopod = medial part of protopod ¾ some appendages have become secondarily uniramous (walking legs) ¾ epipods are modified processes on a protopod that often form gills striated muscles are arranged in antagonistic groups (eg.

the new soft cuticle is stretched. 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. in some species molting ceases after sexual maturity is reached ¾ ¾ Ecdysis is controlled by hormones environmental stimuli such as temperature. 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. enzymes are released above the new epicuticle to dissolve the old endocuticle and soluble products are reabsorbed and stored in the body. secrete a new epicuticle and a new exocuticle 2. predators or scavengers . some calcium salts are stored in the gastroliths in the stomach 3. pistol shrimp use their enlarged chela to snap shut very fast and form a cavitation bubble that can stun prey) ¾ chelipeds are used in prey capture and mandibles and maxillae and other mouthparts used in ingesting food ¾ crustaceans can be suspension feeders. 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 ¾ ¾ ¾ ¾ ¾ ¾ ¾ Feeding Habits of Crustaceans ¾ much variation in feeding habits of crustaceans and some can shift feeding strategies (eg. the animal is defenseless and hides young crustaceans have shorter molting cycles than older mature crustaceans. chitin and calcium salts ¾ endocuticle = made of prinicipal layer which is mostly chitin and calcium and less protein and the uncalcified membranous layer of chitin and protein ¾ ¾ ¾ ¾ ¾ Molting Cycle 1. daylength and humidity changes (for land crabs) signal a decrease in the production of molt inhibiting hormone (MIH) from the X-organ. metabolism and reproductive physiology is tied to the molting cycle ¾ the cuticle is formed of: ¾ epicuticle = outermost thin layer of protein and lipid ¾ procuticle = formed of exocuticle and endocuticle ¾ exocuticle = beneath the epicuticle made of protein.Phylum Arthropoda: Subphylum Crustacea 3 Biological Sciences 102 – Animal Biology – Notes & Vocabulary Molting and Ecdysis in Crustaceans ¾ molting = the process of making a larger cuticle ¾ ecdysis = the shedding of the old cuticle ¾ much behavior. before ecdysis the epidermal cells enlarge and separate from the membranous layer. new endocuticle is deposited and new cuticle is hardened with stored salts and other constituents ¾ ¾ while molting.

lobsters.000 species) ¾ below is the basic body plan of an ancestral malacostracan which is typical of most members of this class seen today ¾ head with five fused segments fused to a thorax of eight segments ¾ abdomen has 6 (sometimes 7) segments ¾ anterior end is a non-segmented rostrum ¾ posterior end is a non-segmented telson with uropods that form the fan tail ¾ carapace is the dorsal cuticle of the head extended posteriorly and around the sides to cover the thoracic and abdominal segments (in decapods. 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 ¾ ¾ ¾ ¾ ¾ ¾ ¾ Amphipoda (eg.1 in the text for specifics and functions of crayfish appendages ¾ crayfish have direct development with no larval form that is seen in many other crustaceans ¾ 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 ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ Decapoda (shrimps. one for swimming and one for jumping) ¾ direct development . crabs. all of the cephalothorax.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 ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ Isopoda (isopods) freshwater. but none of the abdominal segments are covered) ¾ see table 19. crayfish = about 18. 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. beach hoppers) freshwater and marine (eg.Phylum Arthropoda: Subphylum Crustacea 4 Biological Sciences 102 – Animal Biology – Notes & Vocabulary Special Adaptations within the Subphylum Crustacea Class Malacostraca (about 20.

herbivores or predatory mostly dioecious. Cirripedia) ¾ this may be a polyphyletic group divided into individual classes ¾ Subclass or Class Copepoda (copepods) ¾ comprise a large and ecologically important portion of the marine and freshwater zooplankton ¾ usually only a few millimeters or less long ¾ often paddle shaped tapering from anterior to posterior ¾ lack a carapace ¾ simple. tadpole shrimp. Branchiura. single median eye ¾ single pair of antennules ¾ single pair of maxillipeds ¾ four pairs of flattened. biramous thoracic swimming appendages ¾ free-living or symbiotic (many parasitic) ¾ indirect development . Daphnia) form a large part of the freshwater zooplankton ¾ reproduction often by parthenogenesis in the summer and sexual before overwintering fertilized eggs are produced ¾ gradual metamorphosis or direct development Class ¾ ¾ ¾ ¾ ¾ ¾ ¾ Ostracoda (mussel shrimp or seed shrimp) look like tiny clams feed and move using head appendages most live on bottom or on plants.Phylum Arthropoda: Subphylum Crustacea 5 Biological Sciences 102 – Animal Biology – Notes & Vocabulary Order ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ ¾ 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. some parasitic particle and carrion feeders. Tantulocarida. brine shrimp. some parthenogenetic gradual metamorphosis males sometimes emit light to attract females “Class” Maxillopoda (Mystacorarida. some burrow. Copepoda. water fleas) ¾ with or without carapace ¾ flattened leaflike phyllopodia = legs for respiration that are often used for filter feeding ¾ mostly freshwater ¾ water fleas (eg.

and thoracic legs are long cirri with hairlike setae which are used for filter feeding ¾ all marine and often intertidal (aperture between plates closes when exposed) ¾ hermaphroditic with significant metamorphosis ¾ nauplii hatch to become cyprid larvae with a bivalve carapace and compound eyes that attach to substrate with their first antennae via adhesive glands. eyes are lost and appendages transform during metamorphosis ¾ Order Rhizocephalans such as Sacculina are crab parasites. Copepoda. no abdomen. Cirripedia) ¾ Subclass or Class Cirrepedia (barnacles) ¾ enclosed in a shell of calcareous plates ¾ sessile as adults and may have stalk (eg. root-like processes grow throughout the crab’s body and the parasite reproductive structures become externalzed between the cephalothorax and abdomen .Phylum Arthropoda: Subphylum Crustacea 6 Biological Sciences 102 – Animal Biology – Notes & Vocabulary “Class” Maxillopoda (Mystacorarida. calcareous plates are secreted. gooseneck barnacles) ¾ carapace surrounds body and secretes plates ¾ head is reduced. Branchiura. Tantulocarida. 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.