20. Crustaceans

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    Crustaceans

    Chapter 20

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    Subphylum Crustacea

    Crustaceans,

    subphylum

    Crustacea typically

    have biramous,branched,

    appendages that are

    extensively

    specialized forfeeding and

    locomotion.

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    Subphylum Crustacea

    Crustacea is

    divided into 5

    classes.

    Currentmolecular

    phylogenies do

    not support the

    monophyly of allclasses.

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    Subphylum Crustacea - External

    Features

    Secreted cuticle is made of chitin, protein, and

    calcareous material.

    Heavy plates have more calcareous deposits -

    joints are soft and thin, allowing flexibility.

    Dorsal tergum and ventral sternum are plates

    on each somite lacking a carapace.

    Anterior end is a nonsegmented rostrum andthe posterior end is the unsegmented telson.

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    Subphylum Crustacea

    Crustaceans are the onlyarthropods that have two pairsof antennae.

    They also have a pair ofmandibles (jaw-likeappendages) and two pairs ofmaxillae on the head.

    Each body segment usually has

    one pair of appendages. Ancestrally biramous except for the

    first antennae.

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    Subphylum Crustacea -

    Appendages

    Members of Malacostraca andRemipedia have appendages on eachsomite.

    Other classes may not bear appendages onabdominal somites.

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    Subphylum Crustacea -

    Appendages

    Appendages have become specialized by

    evolving into a wide variety of walking legs,

    mouthparts, swimmerets, etc. from

    modification of the basic biramous appendage.

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    Subphylum Crustacea

    The ancestral condition in arthropods is

    to have many body segments.

    Fewer segments and increasedtagmatization is the derived condition.

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    Subphylum Crustacea - Internal

    Features

    Muscular and nervous systems and

    segmentation exhibit metamerism of

    annelid-like ancestors.

    Hemocoel - persistent blastocoel that

    becomes filled with blood.

    Coelomic compartments remain as end

    sacs of excretory organs and gonads.

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    Subphylum Crustacea - Muscular

    System

    Striated muscles make up a major

    portion of crustacean body.

    Most muscles arranged as antagonisticgroups.

    Flexors draw a limb toward the body and

    extensors straighten a limb out.

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    Subphylum Crustacea - Muscular

    System

    Abdominal flexors of a crayfish allow it to

    swim backward.

    Strong muscles located on each side ofstomach control the mandibles.

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    Subphylum Crustacea - Respiratory

    System

    Smaller crustaceansmay exchangegases across thinner

    areas of cuticle. Larger crustaceans

    use featherlike gillsfor gas exchange.

    Bailer of 2ndmaxilla draws waterover gill filaments.

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    Subphylum Crustacea - Circulatory

    Open circulatory system Dorsal heart - single-chambered

    sac of striated muscle.

    Valves in the arteries preventbackflow of hemolymph.

    Hemolymph conducted to gills, ifpresent, for oxygen and carbondioxide exchange.

    Hemolymph may be colorless,reddish, or bluish.

    Hemocyanin (blue) and/orhemoglobin (red) are respiratorypigments.

    Contains ameboid cells thatmay help prevent clotting.

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    Subphylum Crustacea - Excretory

    System Antennal ormaxillary glands are

    called green glands in decapods.

    End sac of antennal gland has a

    small vesicle and a spongy labyrinth.

    Labyrinth connects by an excretorytubule to dorsal bladder that opens

    to exterior pore.

    Resorption of salts and amino acids

    occurs as the filtrate passes the

    excretory tubule and bladder.

    Mainly regulates the ionic and

    osmotic composition of body

    fluids.

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    Subphylum Crustacea - Excretory

    System

    Nitrogenous wastes are excreted across

    thin areas of cuticle in the gills.

    Freshwater crustaceans constantlythreatened by over-dilution with water.

    Gills must actively absorb Na+ and Cl-.

    Marine crustaceans have urine that is

    isosmotic with blood.

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    Subphylum Crustacea - Nervous

    System

    Pair ofsupra-esophageal ganglia connects to eyes and twopairs of antennae.

    Neuron connectives join this brain to the subesophagealganglion.

    Supplies nerves to mouth, appendages, esophagus, and

    antennal glands. Double ventral nerve cord has a pair of ganglia for each somite

    to control appendages

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    Subphylum Crustacea - Sensory

    System

    Eyes and statocysts are the largest sensoryorgans.

    Tactile hairs occur on the body, especially on

    chelae, mouthparts and telson. Chemical sensing of taste and smell occurs in

    hairs on antennae and mouth.

    Statocyst opens at base of first antenna in

    crayfish. Statocyst lined with sensory hairs that detect

    position of grains of sand.

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    Subphylum Crustacea - Sensory

    System Compound eyes are made of

    many units called ommatidia.

    Cornea focuses light down thecolumnar ommatidium.

    Distal retinal, proximal retinal,and reflecting pigment cellsform a sleeve around eachommatidium.

    Each ommatidium detects arestricted area of objects, a

    mosaic, in bright light. In dim light, the distal and

    proximal pigments separateand produce a continuousimage.

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    Subphylum Crustacea - Diversity of

    Reproduction

    Barnacles are monoecious but generally

    cross-fertilize.

    In some ostracods, males are scarce and

    reproduction is by parthenogenesis.

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    Subphylum Crustacea - Diversity of

    Reproduction

    Most crustaceans brood eggs in brood chambers, in

    brood sacs attached to the abdomen, or attached to

    abdominal appendages.

    Crayfishes develop directly without a larval form.

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    Subphylum Crustacea - Diversity of

    Reproduction

    Most

    crustaceans

    have a larva

    unlike the adult

    in form, and

    undergo

    metamorphosis.

    Gulf shrimp

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    Subphylum Crustacea - Diversity of

    Reproduction The nauplius is a common

    larval form with uniramousfirst antennae, and biramoussecond antennae andmandibles that all aid in

    swimming. Appendages and somites are

    added in a series of molts.

    Metamorphosis of a barnacleproceeds from a free-

    swimming nauplius to acypris larva with a bivalvecarapace and finally to asessile adult with plates.

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    Subphylum Crustacea - Ecdysis

    Ecdysis is necessary for a

    crustacean to increase in size

    the exoskeleton does not

    grow. Physiology of molting affects

    reproduction, behavior, and

    many metabolic processes.

    Underlying epidermis

    secretes cuticle.

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    Subphylum Crustacea - Ecdysis

    Molting animals grow in the intermolt phases, orinstars.

    Soft tissue increases in size until there is no spacewithin the cuticle.

    When body fills the cuticle, animal is in the premoltphase.

    Molting occurs often in young animals and may ceasein adults.

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    Subphylum Crustacea - Ecdysis

    Hormonal Control of Ecdysis:

    Temperature, day length, or other stimuli

    trigger central nervous system to begin

    ecdysis.

    Central nervous system decreases

    production of molt-inhibiting hormone by the

    X-organ. Promotes release of molting hormone from

    the Y-organs which promotes ecdysis.

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    Subphylum Crustacea - Endocrine

    Functions

    Removing eyestalks accelerates molting

    and prevents color changes to match

    background.

    Hormones from neurosecretory cells in

    eyestalk control dispersal of cell pigment.

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    Subphylum Crustacea - Feeding

    Habits

    Same fundamental mouthparts in variouscrustaceans are adapted to a wide arrayof feeding habits.

    Suspension feeders generate watercurrents in order to feed on plankton,detritus ,and bacteria.

    Predators consume larvae, worms,

    crustaceans, snails, and fishes. Scavengers eat dead animal and plant

    matter.

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    Subphylum Crustacea - Feeding

    Habits

    Crayfishes have a two-part stomach.

    Gastric mill grinds up food in 1st

    compartment.

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    Class Remipedia

    Only 10 described species inClass Remipedia.

    All found in caves connected tothe sea.

    Primitive features include 2538

    segments with similar, paired,biramous, swimmingappendages.

    Antennules also biramous.

    Maxillae and maxillipeds areprehensile and specialized for

    feeding. Swimming legs are directed

    laterally rather than ventrally as isfound in copepods andcephalocarids.

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    Class Cephalocarida

    Only 9 species described inClass Cephalocarida.

    Live in coastal bottomsediments from intertidal

    zones to 300 meters depth. Thoracic limbs and 2nd

    maxillae are very similar.

    Lack eyes, a carapace, andabdominal appendages.

    True hermaphrodites andunique in discharging eggsand sperm through sameduct.

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    Class Branchiopoda

    Includes three orders:

    Anostraca fairy

    shrimp and brine shrimp,

    no carapace. Notostraca tadpole

    shrimp, carapace forms

    a large dorsal shield.

    Diplostraca waterfleas carapace

    encloses body but not

    head.

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    Class Branchiopoda

    Phyllopodia legs that serve as

    respiratory organs.

    Legs may be used for filter feeding and

    locomotion as well.

    Mostly freshwater forms.

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    Class Branchiopoda

    Water fleas (like Daphnia) produce

    females parthenogenetically in summer.

    Males are produced when unfavorable

    conditions arise and overwintering

    fertilized eggs are produced that are

    resistant to cold and desiccation.

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    Class Ostracoda

    Ostracods are enclosed in a two part

    carapace and look a bit like a clam.

    Marine or freshwater. Mostly benthic.

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    Class Maxillopoda

    axillopods generally

    have 5 cephalic, 6

    thoracic, and 4

    abdominal segmentsplus a telson.

    Reductions common.

    The nauplius of

    maxillopods has a

    maxillopodan eye

    unique to this group.

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    Class Maxillopoda Subclass

    Copepoda

    Planktonic crustaceansinclude many species ofcopepods (subclass

    Copepoda) which are amongthe most numerous of allanimals.

    They lack a carapace.

    Retain the simplemaxillopodan eye in adults.

    Antennules used in swimming.

    Very diverse.

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    Class Maxillopoda Subclass

    Copepoda

    Parasitic forms highly modified and reduced -often unrecognizable as arthropods.

    Free-living copepods may be the dominantconsumer.

    Marine copepod Calanus is most abundantorganism in zooplankton by biomass.

    Cyclops and Diaptomus important elements offreshwater plankton.

    Some free-living copepods are intermediatehosts of human parasitic tapeworms andnematodes.

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    Class Maxillopoda Subclass

    Tantulocarida

    Subclass Tantulocarida

    - only recently

    described.

    Approximately 12species.

    Tiny copepod-like

    ectoparasites of deep-

    sea benthic

    crustaceans.

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    Class Maxillopoda Subclass

    Branchiura

    Members of thesubclass Branchiuralack gills.

    Most areectoparasites ofmarine andfreshwater fish.

    510 mm long. Development is

    direct.

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    Class Maxillopoda Subclass

    Pentastomida

    Subclass Pentastomida -tongue worms.

    Consist of about 90species of parasites of

    vertebrate respiratorysystems.

    Most infect reptile lungs,a few infect air sacs ofbirds or mammals.

    Range from 1 to 13 cm inlength.

    Chitinous cuticle regularlymolted.

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    Class Maxillopoda Subclass

    Cirripedia

    Barnacles subclass cirripedia are a

    group of mostly sessile crustaceans whose

    cuticle is hardened into a shell.

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    Class Maxillopoda Subclass

    Cirripedia

    Their legs are long,

    many jointed cirri that

    extend out through the

    calcareous plates tofilter feed.

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    Class Maxillopoda Subclass

    Cirripedia

    Barnacles are

    hermaphroditic.

    Most hatch as a nauplius

    larva then become a cypridlarva (resembles the

    ostracod Cypris).

    Cyprids attach to the

    substrates and beginsecreting calcareous plates.

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    Class Maxillopoda Subclass

    Cirripedia

    Parasitic forms may have a kentrogon

    stage that injects cells into the hemocoel

    of host.

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    Class Malacostraca

    alacostracans

    usually have a

    head with 5 fused

    segments, athorax with 8

    segments and an

    abdomen with 6.

    Anteriorrostrum

    Posteriortelson

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    Class Malacostraca Order Isopoda

    Order Isopoda including pill bugs. Only truly terrestrial crustaceans.

    Also have marine and freshwater forms.

    Dorsoventrally flattened, lack a carapace, and

    have sessile compound eyes.

    Compressed dorsoventrally.

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    Class Malacostraca Order

    Amphipoda

    OrderAmphipoda many marine, terrestrial& freshwater forms.

    Amphipods resemble isopods: Lack a carapace, have sessile compound eyes,

    and one pair of maxillipeds. However, they are compressed laterally.

    Development is direct.

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    Class Malacostraca Order

    Euphausiacea

    OrderEuphausiacea contains approximately 90species.

    Includes important ocean plankton called krill.

    Most are bioluminescent with a light-producing organ

    called a photophore. Form a major component of the diet of baleen whales

    and of many fishes.

    Eggs hatch as nauplii.

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    Class Malacostraca

    Harder, heavy plates in larger crustaceans

    due to calcareous deposits in addition to

    chitin.

    The carapace covers much or all of thecephalothorax.

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    Phylogeny

    Remipedia appear to be the most

    primitive of Crustacea.

    Two pairs of uniramous limbs on each

    segment.

    One theory is that each modern somite

    represents two ancestral somites that

    fused together, forming the biramousappendage.

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    Adaptive Diversification

    Crustaceans are unquestionably the dominant

    arthropod in marine environments.

    They also share dominance in freshwater

    environments with the insects. The class Malacostraca is most diverse and

    members of Copepoda are most abundant.

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    Classification

    Class Remipedia

    Class Cephalocarida

    Class Branchiopoda Order Anostraca

    Order Notostraca

    Order Cladocera

    Ph l d Ad ti

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    Phylogeny and Adaptive

    Diversification

    Class Ostracoda

    Class Maxillopoda

    Subclass Copepoda Subclass Tantulocarida

    Subclass Branchiura

    Subclass Pentastomida

    Subclass Cirripedia

    Ph l d Ad ti

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    Phylogeny and Adaptive

    Diversification

    Class Malacostraca

    Order Isopoda

    Order Amphipoda

    Order Euphausiacea

    Order Decapoda