Chapter 9
Key Concepts
• Molluscs have soft bodies that are usually covered by a shell.
• Molluscs are important herbivores and carnivores in the marine environment.
• Polychaete diversity stems from the evolution of a segmented body that allows increased motility.
• In addition to being important consumer organisms, polychaetes are the primary prey of many marine animals and play an important role in recycling nutrients.
• Arthropods have external skeletons, jointed appendages, and sophisticated sense organs.
• Crustaceans make up a majority of the zooplankton that are a major link between phytoplankton and higher-order consumers in oceanic food webs.
• Nematodes are abundant and important members of the meiofauna.
• Arrowworms are carnivorous zooplankton.
• Echinoderms exhibit radial symmetry as adults.
• Echinoderms have internal skeletons and a unique water vascular system that functions in locomotion, food gathering, and circulation.
• Acorn worms are benthic suspension feeders and deposit feeders.
• Invertebrate chordates include tunicates, salps, larvaceans and lancelets.
Molluscs
• Phylum Mollusca
• One of the most successful groups
• Have soft bodies, usually covered by a calcium carbonate shell
• Wide range of sizes, lifestyles and relationships to humans
Molluscan Body
• 2 major parts:
– head-foot
– visceral mass
• circulatory, digestive, respiratory, excretory, reproductive systems
• Mantle
– protective tissue covering soft parts
• mantle cavity
• Radula
– a ribbon of tissue containing teeth
Molluscan Shell
• Secreted by the mantle
• Normally comprises 3 layers:
– periostracum
– prismatic layer
– nacreous layer
Chitons
• 8 shell plates
• Attach tightly to rocks, in intertidal zone
• Most scrape algae off the rocks
Gastropods
• Class Gastropoda
• unshelled or a univalve shell
– operculum
• Feeding and nutrition
– exhibit wide variety of feeding styles
• Naked gastropods – Nudibranchs
– cerata
– stinging cells as defensive weapons
– bright colors indicate toxicity to predators
• Reproduction and development
– separate sexes
– internal fertilization
– some shed eggs directly into the sea
– 2 types of free-swimming larva
• trochophore
• veliger
– some are hermaphroditic
– some without planktonic larval stage
Bivalves
• shells divided into 2 valves
• includes:
– clams
– oysters
– mussels
– scallops
– shipworms
• Bivalve anatomy
– no head or radula
– laterally compressed bodies
– shell halves attached dorsally
• umbo
• adductor muscles
– inhalant and exhalant openings
– Siphons – facilitate filter feeding while buried
• Reproduction in bivalves
– majority have separate sexes
– fertilization usually occurs in the water column
– trochophore and veliger stages
– some hermaphroditic
Cephalopods
• Named after the foot
• tentacles projects used in prey capture, defense, reproduction and sometimes locomotion
• most lack shells
• Types of cephalopods
– nautiloids
• large, coiled shells
– siphuncle
• head has 60-90 tentacles
• jet propulsion
• diurnal migration
• eat hermit crabs and scavenge on the bottom
• food is stored in crop
– coleoids
• cuttlefish
• squids
• octopods
• coleoids
• swim by jet propulsion or by fin undulation
• have the most advanced, complex nervous system among invertebrates
• Color and shape in cephalopods
– body movements and color changes are used in communication
– chromatophores
• Feeding and nutrition
– carnivores
– beak-like jaws
– diet varies with habitat
• Reproduction in cephalopods
– sexes are separate
– often includes courtship display
– modified arm used to transfer spermatophore
– some females incubate eggs
– usually reproduce once and then die
Sipunculids
• solitary benthic worms
• live in burrows
• peanut worms
• suspension or deposit feeders
• separate sexes, external fertilization
Annelids: The Segmented Worms
• bodies are divided into segments
– hydrostatic skeleton
– setae
• Types of marine annelids
– polychaetes
– echiurans
– pogonophorans
Polychaetes
• Most common marine annelids
• Traditionally divided into 2 groups:
– errant polychaetes
– sedentary polychaetes
• Feeding and digestion
– predators, filter or suspension feeders
– digestive tract is usually a straight tube
– deposit feeders
• selective and nonselective deposit feeders
• Reproduction in polychaetes
– asexual reproduction via budding or fragmentation
– most reproduce only sexually
– gametes are released into the water column
– epitoky
– swarming
Echiurans
• Mostly deposit feeders
– typically have a flat proboscis to collect particles
• separate sexes, shed gametes into the water column, and have a planktonic larval stage
Pogonophorans
• Live in buried tubes
• cylindrical body with a ring of tentacles
• Lack mouth or digestive tract
• May absorb nutrients dissolved in seawater or obtain nourishment from chemosynthetic bacteria
Nematodes
• Most numerous animals on earth
• Critical role as scavengers
• Some are parasitic
• Most are hermaphroditic
Ecological Role of Marine Worms
• Nutrient cycling
• Predator-prey relationships
• Symbiotic relationships
Arthropods
• 75% of identified species
• molting exoskeleton
• divided into segments
• jointed appendages
• developed nervous systems
• 2 major groups of marine arthropods:
– chelicerates
– mandibulates
Chelicerates
• Primitive group
• 6 pairs of appendages
• Horseshoe crabs
– 3 body regions
• cephalothorax
• abdomen
• telson
– carapace
• Sea spiders
– 4+ pairs of legs
– male carries eggs
– palps
– feed on juices from cnidarians and other soft-bodied invertebrates, using a long sucking proboscis
Mandibulates
• Crustaceans—marine mandibulates
– 3 main body regions:
• head
• thorax
• abdomen
– appendages:
• 2 pairs of sensory antennae
• mandibles and maxillae
Decapods
• 5 pairs of walking legs
• chelipeds
• wide range in size
• Specialized behaviors
– hermit crabs
– decorator crabs
– common blue crabs
• Nutrition and digestion
– chelipeds
– appendages are used for scavenging
– some are deposit or filter feeders
• Reproduction
– sexes usually separate
– males have modified appendages modified
• spermatophores
• copulatory pleopods
– most brood their eggs
– larval stages:
• zoea larval stage
• nauplius larva
Mantis Shrimp
• specialized predators
• 2nd pair of thoracic appendages
• reproduction
– some pair for life
– larvae hatch from egg mass
– retain planktonic form for up to 3 months
Krill
• Pelagic, 3-6 cm long
• Filter feeders
• Most are bioluminescent
– photophore
– swarms
• Food source
Amphipods
• Laterally compressed, with posterior 3 pairs of appendages directed backward
• Many are burrowers
• Most are detritus feeders or scavengers
• Young resemble adults upon hatching
Copepods
• Largest group of small crustaceans
• Abundant member of the zooplankton
• Mostly suspension feeders
• Males use spermatophores
• Eggs are shed and hatched into water
Barnacles
• Only sessile crustaceans
• Most have calcium carbonate shell
• Attach to a hard surface
• Filter feed using cirripeds
• Reproduction
– hermaphroditic
– long, extensible penis
– nauplius larvae develop into cyprid larvae
Ecological Roles of Arthropods
• Arthropods import are source of food
– copepods form link in marine food chain
– krill are consumed in large quantities
• Arthropods as symbionts
– cleaning shrimps
– ecto and endo parasites for fish
– barnacles are commensal
• Role of arthropods in recycling and fouling
Arrowworms
• Phylum Chaetognatha
– common and planktonic
– carnivorous, voracious feeders
– 4 – 14 grasping spines
Echinoderms: Animals with Spiny Skins
• Sea stars, sea urchins and sea cucumbers
• Bilateral symmetry in larvae, radial symmetry in adults
Echinoderm Structure
• Endoskeleton
– ossicles
– pedicellariae
• Water vascular system
– madreporite
– tube feet
Sea Stars
• Central disk with 5+ rays
• ambulacral grooves
• Aboral surface
• Feeding in sea stars
– most are carnivores or scavengers
– sea stars envelope and open bivalves, evert a portion of the stomach, and insert it into the bivalves to digest them
• Reproduction and regeneration
– sea stars can regenerate rays
– most have separate sexes, eggs and sperm released into the water
Ophiuroids
• Benthic with 5 spiny arms
• Lack pedicellariae
• Tube feet lack suckers
• Feeding in ophiuroids
– brittle stars usually filter feed
– deposit feeders use their podia
– basket stars suspension feed
Ophiuroids
• Reproduction and regeneration in ophiuroids
– autotomize
– asexual reproduction by division
– planktonic larvae metamorphose into adults before settling
Sea Urchins and their Relatives
• Body enclosed by test
• Benthic
• Moveable spines
• Feeding in echinoids
– regular echinoids – grazers, Aristotle’s lantern
– feeding in irregular urchins - deposit feeders
Sea Cucumbers
• Class Holothuroidea
• Respiratory trees
• Sexes are generally separate
• Eggs may be brooded or incubated
• Larvae are planktonic
• Feeding in sea cucumbers
– deposit or suspension feeders
– oral tentacles
• Defensive behavior
– Cuvierian tubules
– can eviscerate
Crinoids
• Feather stars, sessile with grasping cirri
• Suspension feeders
• Can regenerate lost arms
• Separate sexes shed
• larvae have free-swimming stage
Ecological Roles of Echinoderms
• Spiny skins deter most predators
• Predators of molluscs, cnidarians, crustaceans
• Urchins eat kelp, coral
• Sea cucumber poison, holothurin, has potential as a medicine
Hemichordates
• Sessile, burrow in sediments
• Large proboscis, used to dig burrows
Invertebrate Chordates
• 4 key anatomical characteristics
1) notochord
2) pharyngeal gill slits
3) post anal tail
4) dorsal, hollow nerve tube
Tunicates
• Subphylum Urochordata
• Mostly sessile, widely distributed
• Named for their tunic
• Types:
– sea squirts
– salps
– larvaceans
Sea Squirts
• Class Ascidiacea
• Tends to expel a water when disturbed
• Filter feeders
• 2 projecting tubes
– incurrent siphon
– excurrent siphon
• Lifestyles: solitary, colonial, compound
• Filter feeders
• Budding occurs in colonial ascidians
• Most are hermaphrodites
• Short lived, tadpole-like larvae
Salps and Larvaceans
• Salps
– free-swimming tunicates with incurrent and excurrent siphons
• Larvaceans
– free-swimming; produce delicate enclosures of mucus used in feeding
Cephalochordates
• Fish-like chordates
• eel-like in form and behavior
• Benthic
• Suspension feed
• Separate sexes
• Internal fertilization
• Complex life cycles