Segmented Worms

Chapter 17
Annelids and Allied Taxa

l Phylum Annelida
l Class
Polychaeta
l Class
Oligochaeta
l Class Hirudinea
l Phylum Echiura
l Phylum
Sipuncula
Annelids and Allied Taxa

l Members of phyla
Echiura and
Sipuncula are
benthic marine
animals with
unsegmented
bodies.
Phylum Annelida

l Annelids are
protostome coelomates
in superphylum
Lophotrochozoa.
l Spiral, determinate
cleavage.
l Nervous system more
centralized & circulatory
system more complex
than in previous phyla.
Phylum Annelida
• Annelids are segmented worms.
• They have bodies composed of a series of
fused rings.
• Earthworms, leeches, clam worms.
Phylum Annelida

l The evolutionary innovation shown by

annelids is segmentation
(metamerism).
l Segmentation evolved separately in
annelids, arthropods, and chordates.
l The body is divided into a series of
segments, each having similar
components of all major organ systems.
l Built in fail-safe.
l Allows for specialization.
Phylum Annelida
Many annelids have
chitinous bristles called
setae.
Help in locomotion
Anchor worm in place
Deter predators
Phylum Annelida

l Annelids can be found worldwide in

marine, freshwater, and terrestrial
habitats.
Phylum Annelida – Body Plan

Prostomium – anterior part

followed by segmented
body.
Pygidium – terminal
portion.
Phylum Annelida – Body Plan

l Peritonia (layers of
mesodermal
epithelium) of
adjacent segments
meet to form septa.
l Fluid-filled coelom
acts as a
hydrostatic
skeleton.
Phylogeny
l Traditionally, annelids are divided among 3
classes:
l Class Polychaeta
l Class Oligochaeta
l Class Hirudinida
l Polychaeta is a paraphyletic class.
l Oligochaeta and Hirudinida form a
monophyletic group called Clitellata.
l Characterized by reproductive structure called a
clitellum.
l Class Oligochaeta is a paraphyletic group
because ancestors of leeches arose from
within it.
Class Polychaeta

l Polychaeta is the
largest, most
diverse class.
l May be brightly
colored, variable
shape.
Class Polychaeta

l Polychaetes have
some features other
annelids do not:
l A well developed
head.
l Paired appendages,
parapodia, that
function as gills and
aid in locomotion.
l No clitellum.
l Many setae
Class Polychaeta

l Polychaetes are
mostly marine and
mostly benthic.
l May live under
rocks, burrow into
sediment, or build
their own tubes.
l Some are
planktonic.
Class Polychaeta

l Sedentary and
errant (free-
moving) forms.
l Sedentary forms
often have
elaborate devices
for feeding and
respiration.
l Filter or deposit
feeders.
Class Polychaeta

l Errant forms
include pelagic
and benthic
types and are
often predators
or scavengers.
Class Polychaeta - Reproduction
l Gonads are temporary structures in polychaetes.
l Sexes usually separate.
l Fertilization is external.
l Early larva is a trochophore.
Circulation and Respiration
l Most have parapodia and gills for gaseous
exchange.
l Others use the body surface.
l Circulation varies.
l In Nereis a dorsal vessel carries blood forward and
a ventral vessel carries blood posteriorly.
l Blood flows across between these major vessels in
networks around the parapodia and intestine.
l In some, septa are incomplete and coelomic
fluid serves circulatory function.
l Many polychaetes have respiratory pigments -
Hemoglobin, chlorocruorin or hemerythrin.
Excretion

l Excretory organs vary, from protonephridia to

metanephridia, and mixed forms.
l One pair per metamere.
l Inner end (nephrostome) opens into the
coelomic cavity.
l Coelomic fluid enters the nephrostome.
l Selective resorption occurs along the
nephridial duct.
Nervous System and Sense Organs

l Double ventral nerve cord runs length of the

worm with ganglia in each metamere.
l Sense organs include:
l Eyes, nuchal organs and statocysts.
l Eyes vary from simple eyespots to well-developed
image-resolving eyes similar to mollusc eyes.
l Nuchal organs are ciliated sensory pits that are
probably chemoreceptive.
l Some burrowing and tube-building polychaetes use
statocysts to orient their body.
Representative Polychaetes
l Clam Worms: Nereis
l Errant polychaetes
l Live in mucus-lined burrows
near low tide level.
l Come out of hiding places at
night to search for food.
l Prostomium bears a pair of
palps sensitive to touch and
taste, a pair of short sensory
tentacles, and two small dorsal
eyes sensitive to light.
l Peristomium has a ventral
mouth, a pair of jaws, and four
pairs of sensory tentacles.
Representative Polychaetes

l Scale worms
l Flattened bodies
are covered with
broad scales.
l Some are large, all
are carnivores and
some are
commensals in
burrows of other
organisms.
Representative Polychaetes

l Fireworms
l Have hollow, brittle
setae that contain
poisonous
secretions.
l Feed on cnidarians.
Representative Polychaetes

l Tubeworms
l Tube-dwellers
l May line their
burrows with mucus
l Use cilia or mucus
to obtain food
Representative Polychaetes

l Fanworms or
Featherduster worms
l Unfurl tentacular
crowns to feed.
l Food moved from
radioles to mouth
by ciliary action.
Representative Polychaetes

l Parchment Worms
l Lives in a U-shaped
tube.
l Modified segments
pump water through
tube.
Clade Siboglinidae (Pogonophorans)

l Formerly members of
phylum Pogonophora
(beardworms).
l Discovered in 1900.
l 150 species described.
l Most are small, less
than 1 mm in diameter.
l Giant beardworms that
live in deepwater
hydrothermal vents are
3 m long and 5 cm in
diameter.
Clade Siboglinidae
(Pogonophorans)
l Most live in mud on ocean floor at depths of 100 to
10,000 m.
l Sessile animals that secrete and live in long chitinous
tubes.
l Tubes have general upright orientation in bottom
sediments.
l Tubes are generally three or four times the length of the
animal.
Clade Siboglinidae
(Pogonophorans)
l Long cylindrical body covered with cuticle.
l Divided into a short anterior forepart, a long slender
trunk, and a small, segmented opisthosoma.
l Tentacles are hollow extensions of the coelom and
bear minute pinnules.
Clade Siboglinidae
(Pogonophorans)
l No mouth or digestive tract.
l Nutrients such as glucose and amino
acids absorbed from seawater through
pinnules and microvilli of tentacles.
Clade Siboglinidae
(Pogonophorans)
l Most energy derived from a mutualistic
relationship with chemoautrophic
bacteria that oxidizes hydrogen sulfide.
l Trophosome, derived embryonically from
midgut, houses the bacteria.
Clade Siboglinidae
(Pogonophorans)
l Sexes are separate.
l Research suggests that cleavage is unequal
and atypical.
l Appears to be spiral.
l Coelom formed by schizocoely.
l Embryo
l Worm-shaped and ciliated.
l Poor swimmer.
l Probably carried by water currents until it settles.
Clade Clitellata
l Class Oligochaeta and Class Hirudinida
l Form reproductive structure called a clitellum.
l Ring of secretory cells found in a band around the body.
l Permanent in oligochaetes but visible only during reproductive
season in leeches.
l Members are derived annelids that lack parapodia.
l Hermaphroditic (monoecious) animals that exhibit
direct development.
l Young develop inside a cocoon secreted by the
clitellum, and emerge as small worms.
Class Oligochaeta

l Class Oligochaeta includes earthworms

and many freshwater worms.
l They possess setae, but not as much as
polychaetes.
Class Oligochaeta
l Earthworms are the
most familiar
oligochaetes, found in
moist, rich soil.
l They can burrow deep
underground and remain
dormant in a slime
chamber during dry
weather.
l Setae help prevent
slipping while burrowing.
Class Oligochaeta

l Darwin wrote about earthworms in The

Formation of Vegetable Mould Through
the Action of Worms.
l He noted the beneficial activities of worms,
aeration, moving nutrients up from subsoil,
adding nitrogenous products, breakdown of
organic matter in dead leaves etc.
l An earthworm can ingest its own weight in
soil every 24 hours.
Class Oligochaeta - Reproduction

l Earthworms are hermaphroditic – male and

female organs in the same animal.
l When mating, two worms are held together by
mucus secreted by the clitellum.
Class Oligochaeta - Reproduction

l After mating, a
cocoon forms
around the
clitellum, as it
passes forward it
gathers both
gametes, and
fertilization
occurs inside.
Class Oligochaeta - Reproduction

l Development occurs
inside the cocoon
and young worms
hatch out.
l Development is
direct, no larval
stage.
Class Oligochaeta - Feeding

l Food is stored in a thin-walled crop.

l Muscular gizzard grinds food into small
pieces.
l Digestion and absorption occur in intestine.
Class Oligochaeta - Excretion
l Each somite, except the 1st
three and terminal one, have a
pair of metanephridia.
l A ciliated funnel, the
nephrostome, draws in wastes
and leads through the septum.
l These coil until the nephridial
duct ends at a bladder that
empties outside at
nephridiopore.
l Wastes from both the coelom
and the blood capillary beds
are discharged.
l Aquatic oligochaetes excrete
toxic ammonia.
Class Oligochaeta - Circulation and
Respiration
l Coelomic fluid and blood transport food, wastes, and
respiratory gases.
l Blood circulates in a closed system with five main
trunks running lengthwise in the body.
l Dorsal vessel contains valves and functions as a true
heart.
l Pumps blood anteriorly into 5 pairs of aortic arches.
l Aortic arches ensure steady pressure in ventral vessel.
Class Oligochaeta - Nervous
System and Sense Organs
l Central nervous system and
peripheral nerves.
l Pair of cerebral ganglia
connect around the pharynx to
the ganglia of the ventral nerve
cord.
l Neurosecretory cells in brain
and ganglia secrete
neurohormones.
l Regulate reproduction,
secondary sex characteristics,
and regeneration.
l Lack eyes but have many
photoreceptors in the
epidermis.
l Free nerve endings in
tegument are probably tactile
structures.
Class Oligochaeta - General
Behavior
l Avoid bright light (negative phototaxis).
l Chemical stimuli are important in
locating food.
l Limited learning ability - primarily trial-
and-error learning.
Class Hirudinea

l Class Hirudinea
includes the leeches.
l Primarily freshwater,
a few marine &
terrestrial.
l More common in
tropical climates.
Class Hirudinea

l Many leeches live as carnivores on small

invertebrates.
l Some are temporary parasites.
l Some are permanent parasites – they never
leave their host.
Class Hirudinea

l Leeches are hermaphroditic

and have a clitellum (only
appears during breeding
season), like oligochaetes.
l Leeches do not have setae.
l They’ve developed suckers
for attachment and a
specialized gut for storing
large amounts of blood.
Class Hirudinea - Respiration and
Excretion
l Some fish leeches have gills.
l All other leeches exchange
gases across epidermis.
l 10 to 17 pairs of nephridia.
l Coelomocytes and other
special cells may assist in
excretion.
Class Hirudinea - Nervous and
Sensory Systems
l Two “brains”
l Anterior fused ganglia form a ring around
the pharynx.
l Seven pairs of posterior fused ganglia.

l 21 pairs of segmental ganglia in between

along a double nerve cord.
l Epidermis contains free sensory nerve
endings and photoreceptor cells.
l Pigment-cup ocelli are present.
Class Hirudinea - Circulation

l Coelom reduced by invasion of

connective tissue.
l Forms system of coelomic sinuses and
channels.
l Some have a typical oligochaete
circulatory system.
l Coelomic system is auxiliary.
l Some lack blood vessels and coelomic
sinuses serve as only vascular system.
Class Hirudinea

l Leeches are highly sensitive to stimuli

associated with the presence of prey.
l Those that feed on mammals are attracted
by warmth.
Phylum Echiura
l Approximately 140
species of marine
worms that burrow into
mud or sand.
l Live in empty snail
shells or sand-dollar
tests, or rocky crevices.
l Found in all oceans.
l Length varies from a few
millimeters to 40 or 50
cm.
Phylum Echiura – Form and
Function
l Sausage-shaped.
l Inextensible proboscis anterior to
the mouth.
l Often called spoon worms”.
l Simple nervous system with a
ventral nerve running length of the
body.
l Ciliated groove on the proboscis
allows them to gather detritus over
the mud while lying buried.
l Muscular body wall is covered by a
cuticle and epidermis which may
be smooth or covered by papillae.
Phylum Echiura
l Large coelom.
l Digestive tract long and coiled.
l Pair of anal sacs may serve an
excretory and osmoregulatory
function.
l Most have a closed circulatory
system with colorless blood.
l Hemoglobin found in certain
cells and in coelomic
corpuscles.
l Respiration probably occurs in
hindgut which is continually filled
and emptied by cloacal irrigation.
Phylum Echiura - Reproduction

l Sexes are separate.

l Gonads produced by special regions in
peritoneum in each sex.
l Fertilization usually external.
l Early cleavage and trochophore stages
similar to annelids.
Phylum Sipuncula
l Approximately 250 species
of benthic marine worms.
l Sedentary, living in burrows
of mud or sand, snail shells,
coral crevices, or among
vegetation.
l More than ½ restricted to
tropical zones.
l Some are tiny, slender
worms, but most range from
3 to 10 cm in length.
l Some are known as “peanut
worms” because when
disturbed, they contract to a
peanut shape.
Phylum Sipuncula - Form and
Function
l No segmentation or setae.
l Slender, retractable introvert or
proboscis at anterior end.
l Walls of the trunk are muscular.
Phylum Sipuncula - Nutrition
l Some appear to be detritivores
and others suspension feeders.
l Some nutrition may come from
dissolved organic matter in the
surrounding water.
l From burrow or hiding place,
they extend tentacles to explore
and feed.
l Collected organic matter moved
from mucus on tentacles to
mouth by ciliary action.
l Large fluid-filled coelom.
l Digestive tract is U-shaped.
Phylum Sipuncula - Respiration

l Lack a circulatory and respiratory system.

l Gas exchange appears to occur across
the introvert and tentacles.

17-59
Phylum Sipuncula - Nervous and
Sensory Systems
l Bilobed cerebral
ganglion behind
tentacles.
l Ventral cord
extends the length
of body.
Phylum Sipuncula - Reproduction

l Sexes are separate.

l Sex organs develop seasonally within
the connective tissue covering the
origins of the retractor muscles.
l Sex cells are released through the
nephridia..
l Asexual reproduction occurs by
transverse fission.
Phylogeny

l Similarities in the early development of

molluscs, annelids, and some primitive
arthropods indicate that these three
groups are probably closely related.
l Trochophore larva
l Spiral cleavage

l Schizocoelous coelom formation

Evolutionary Significance of
Metamerism
l No satisfactory explanation for origins of metamerism
and coelom has gained acceptance.
l Coelom may have been advantageous as a
hydrostatic skeleton.
l Coelomic fluid would have acted as a circulatory fluid
and reduce need for flame cells everywhere.
l Coelom could store gametes for timed release.
l Would require nervous and endocrine control.
l Unlikely that segmentation is homologous among
annelids, arthropods, and chordates.
l Current evidence supports the hypothesis that
segmentation arose independently multiple times.
Evolutionary Significance of
Metamerism
l Selective advantage of a segmented body
for annelids appears to lie in the efficiency
of burrowing.
l However, does not explain segmentation in
arthropods given the rigidity of the
exoskeleton.
Phylogeny and Adaptive
Diversification
l Molluscs and annelids share many
developmental features so are presumed to be
closely related.
l However, shared features are likely to be a
retained ancestral feature for lophotrochozoan
protostomes.
l Pogonophorans and vestimentiferans were
once placed in Annelida but are now in clade
Siboglinidae.
l Molecular analyses place sipunculids and
echiurans closely related to the annelids.
Phylogeny and Adaptive
Diversification
l With more studies, Echiura, like Pogonophora,
may no longer be a valid phylum.
l Placement of Sipuncula is contentious.
l Members are not metameric and lack setae.
l Larval development similar to annelids, molluscs,
and echiurans.
l Molecular data may clarify position within
Lophotrochozoa.
l Presently, depicted as the sister taxon to a clade of
annelids and echiurans.
l Polychaeta is a paraphyletic group.
Phylogeny and Adaptive
Diversification
l Adaptive Diversification
l Oligochaetes are constrained by terrestrial soil
environment.
l Polychaetes inhabit a wide range of habitats.
l Septal arrangement with fluid-filled compartments
has been varied for precise movements.
l Feeding adaptations vary widely, from chitinous
jaws to specialized tentacles.
l Leeches have developed both parasitic and
predatory adaptations.