Chapter 13: PHYLUM ANNELIDA -The remaining organsappendages (when present), coelomic

cavities, nephridia, and gonadsare segmental and repeated in

-Annelida = ringed each segment.
-occupy the large-worm niche in marine, freshwater, and terrestrial
habitats Body Wall
-approximately 12 000 species
-most diverse taxon of large-bodied worms -Annelid body wall:
-evolved in conjunction with segementation(construction of a body
Fibrous collagenous cuticle
from a series of modular sections)
o Cuticular fibers, arranged in a crossed-helical
-all feeding modessuspension feeding, deposit feeding,
pattern, toughen the body wall, resist bulges, and
scavenging, herbivory, carnivoryare represented by annelids
often impart an iridescent sheen to the body
o Polychaetes lack cuticle, but their secreted tubes
FORM AND FUNCTION resemble the cuticle
Segmentation Glandular monolayered epidermis
-Annelid body is composed of three regions: o Mucus-secreting gland cells are common
Prostomium o Mucus consolidates the burrow lining and
o Anterior end of the trunk, which contains the protects surface of the body
brain and bears the sense organs o Chaetae are chitinous bristles that project
Trunk outward from the epidermis to provide traction
o Elongate, consists of a longitudinal series of and perform other tasks
similar body units, the segments, which are Simplest chaetae are unjointed,
separated from each other externally by a tapered bristles (capillary chaetae) in
shallow constriction bilaterally paired bundles, one
o Annelids are named for the body annulations that dorsolateral and one ventrolateral pair
result from the ringlike segments per segment
o Segments lie between the prostomium and Each chaeta arises from a pitlike
pygidium epidermal follicle composed of:
o The first segment, peristomium, lies Follicle cells, which form the
immediately behind the prostomium and ventrally follicle wall, and
surrounds the mouth Chaetoblast cell at the base
Pygidium of the follicle, only one
o Forms the posterior end of the body and includes (single)
the anus Chaeta is secreted around long
microvilli on chaetoblast surface
Microvilli later withdraw,
-The prostomium and pygidium, though segmentlike in appearance,
leaving a bundle of parallel,
are not considered to be segments because they do not develop
hollow canals in the
from the segmental growth zone.
completed chaeta.
-Growth Zone is localized to a region immediately in front of the Base of each chaeta remains in the
pygidium follicle, but the shaft extends beyond it
above the epidermal surface
Cells in this zone are paired ectodermal and mesodermal Anchored by
teloblast cells that divide and differentiate to form each hemidesmosomes to the
new segment. follicle cells, which are linked
to the subepidermal muscles
-Teloblastic Growth is the body growth; it results from the that move the chaeta
successive addition of segments posteriorly Composed of -chitin
Polymerized, chainlike
Youngest body segment lies immediately in front of the molecules are parallel and
pygidium share the same polarity
The oldest (peristomium) immediately behind the Tough and flexible
prostomium Some annelids harden and
stiffen their chaetae with
-All internal structures are segmented, but some organs and tissues sclerotized protein and
also span segments, integrating them into an individual worm inorganic materials such as
CaCO3
-The chief integrating structures are the nervous system, hemal
Connective-tissue dermis (Cutis) of varying thickness
system, musculature and digestive system
Musculature derived from coelomic lining
o Obliquely striated musculature
 o Outer circular Layer
o Inner longitudinal layer
The longitudinal fibers typically are
grouped together to form distinct
muscle bands
o The coelomic lining can be a mesothelium
composed solely of epitheliomuscular cells of the
body-wall musculature
o Musculature can be separated from the coelom
by a peritoneum
-Chaetae of annelids
Different from arthropod setae
o Hair- or bristlelike sensory projections that pivot
on a flexible joint with the exoskeleton
o Deflection stimulates a sensory neuron attached
to setal base
o Develops from an epidermal follicle composed of:
Trichogen cells (one or more) which
secrete the seta
Tormogen cells (one or more) which
produce the pivot joint
Sensory Cell
Thecogen
o May be hollow or solid, but lacks the intenal
bundle of parallel canals associated with the
chaetae
o Composed of -chitin
Adjacent parallel molecules have
opposite polarity
Hard and stiff
Can be further enhanced by sclerotized
protein (as in insects)or CaCO3 (in
many crustaceans)
Nervous System
-Central Nervous System consists of:
Anterior-Dorsal Brain
o Located in the prostomium
o Consist of a pair of dorsal suprapharyngeal
ganglia
o Joined to a pair of ventral subpharyngeal ganglia
in the peristomium, by a pair of circumpharyngeal
connectives
The subpharyngeal ganglia give rise to
the nerve cords that extend the length
of the body
Ventral pair of Longitudinal Nerve Cords
o Bear a pair of ganglia that are joined together by
a transverse nerve (commissure), in each of the
segments
Each pair of trunk ganglia is essentially
a segmental brain
From which a bilateral pair of
segmental nerves enters
the body wall, arches
dorsally, nearly or completely
encircles the body, and
innervates the body-wall
musculature and sensory
structures of that segment.
o Has large-diameter giant axons
Low resistance, thus, allows for rapid
impulse conduction and have shorter
response times for the worm to do
rapid retreat (for defense)
Branches from the giant axon run to
the longitudinal muscles
Functions only in the escape response
and not in normal locomotion
If severed, locomotion is
unaffected but escape
response is blocked.
-Ladderlike Nervous System
-Few annelids have intraepidermal NS, but in most, it is submerged
in the connective tissue below the epidermis
-Polyneural innervation
Innervation pattern of annelid body muscles, similar to
arthropods
One muscle fiber (cell) is innervated by more than one
neuron
Rate (speed) and force of fiber contraction depends on the
summed effects of all the neurons, each of which alone
elicit a different response from the fiber
-Sensory Structures are primarily unicellular receptors
Photoreceptors
Chemoreceptors
Mechanoreceptors
Sense Organs
o Ocelli, Eyes (maybe present in Hirudinea),
Statocysts, Chemosensory Nuchal Organs (all
present in Polychaeta)
o Present in Prostomium, peristomium, and first
few segments
In some polychaetes, they also occur
on trunk segments, as well as on the
pygidium
Coelom and Hemal System
-Bilateral pair of coelomic cavities
Occurs in each segment
o The pair of coelomic cavities are lined by ciliated
mesothelium
Mesothelium on the body-wall side of
the coelom is composed of modified
epitheliomuscular muscles that form
the body-wall muscles
 Contractile mesthelial cellsmay also
form a musculature on the septal faces
(radial and circular muscles), around
the blood vessels (circular muscles)
and around the gut (predominantly
circular muscles)
Maybe specialized locally as
chlorogogen cells, which form a
yellow or brown layer of tissue around
part of the gut wall and certain blood
vessels
Chlorogogen Cells
Plays a vital role in
intermediate metabolism
similar to vertebrate liver
Chief center of glycogen and
fat synthesis and storage
Storage and detoxification of
toxins, hemoglobin synthesis,
protein catabolism, formation
of ammonia and urea
synthesis
Longitudinal Mesenteries separate the left and right
coelomic cavities in each segment
o One dorsal, one ventral to the gut, in the
midsagittal plane of the body
o Structurally similar with the septum
Isolated from neighboring segments by transverse septa
o The placement of septa coincides with the
constrictions on the surface of the body
o Each septum is composed of two layers of
mesothelium:
One from the segment in front
One from the segment behind
A layer of connective tissue is
sandwiched between the two
mesothelium
o Septa isolate the hydrostatic skeleton of each
segment
The force of a segmental muscular
contraction can be directed and applied
solely to that segment
o Well-developed septa occur in annelids that
move by actively using the appendages or whole-
body peristalsis
o Incomplete or Reduced septa occur in
semisessile annelids that move by other means
such as with cilia (small-bodied annelids),
whipping movements, or eversion-retraction of
the pharynx (some burrowers)
Fluid skeleton (hydrostat) against which the muscles act to
change body shape
o Contraction of longitudinal muscles causes the
body to widen
o Contraction of circular muscles causes the body
to elongate
-Coelomic Fluid
Circulated by cilia and the contraction of body-wall muscles
Contains Coelomocytes
o For internal defense and gas transport (for
hemoglobin-containing coelomocytes)
-Well-developed hemal system of blood vessels and hearts
Principal vessels:
o Dorsal blood vessel in the dorsal mesentery
o Ventral blood vessel in the ventral mesentery
Blood Flow
o Anteriorly in dorsal blood vessel
o Posteriorly in ventral blood vessel
o In each segment, blood in the ventral vessel
returns to the dorsal vessel via a capillary
network (plexus) in the body wall
o Blood in the dorsal vessel reaches the ventral
vessel by way of capillary plexus, vessel or
sinuses around th wall or the gut
Muscular heart
o Formed by enlarged, anterior region of the dorsal
vessel of some annelids
o In many earthworms, the dorsoventral gut
vessels in the anterior end are specialized hearts
In annelids with reduced or absent blood vessels, coelomic
fluid performs the task of whole-body transport.
-Gas exchange
Across body wall, appendages and gills
Respiratory pigment is Hemoglobin
o Occur in coelomic fluid, blood, muscle, and nerve
o Packaged with coelomytes in the coelom
o Dissolved in the plasma of blood
Excretory System
-Segmental Nephridia
Either Metanephridial System, or
o Occurs in annelids that have hemal system,
some vessels of which bear the podocyte-
covered filtration sites
Protonephridia
o Present in annelids that lack a hemal system
-The inner ends of the nephridia (funnels or terminal cells,
respectively) are often associated with the anterior face of each
septum, from which they project into the coelomic fluid
-Nephridial tubule penetrates the septum and passes into the next
posterior segment before opening to the exterior
Digestive System
-Straight tube that extends between the mouth and the pygidial anus
-Gut passes through the septa and supported above and below by
the mesenteries
-Gut tube:
Ectodermal foregut
o Specialized into two regions:
Muscular, often protrusible and
eversible pharynx
Ciliated Esophagus
Links pharynx to the midgut
 Endodermal Midgut (stomach and intestine) Monophyletic (accdg. to R.O.
Ectodermal hindgut (rectum) Brinkhurst)
Autapomorphy-the muscular
Reproduction and Development pad or bulb uniquely in the
-All forms of asexual (clonal) reproduction dorsal wall of the pharynx
Fragmentation, budding, transverse division (paratomy and -Annelids and Panarthropods (Onychophora and Arhtropoda) are
stolonization) sister taxa of Articulata, the segmented protostomes
-Primitively Gonochoric Evidences: Segmentation, teloblastic growth, expression of
-Segmental Gonads the gene engrailed at posterior margin in developing
Submesothelial clusters of germ cells that are released and segments
stored in the coelomic cavities Yet, molecular systematists place annelids, sipunculan
-Gametes molluscs and other protostomes with trochophore larva
Spawned through metanephridia under Trochozoa taxon;
-External Fertilization o Panarthropods and other molting animals under
-Zygotes undergo spiral cleavage Ecdysozoa taxon

-Trochophore larva
Apical Tuft EVOLUTION AND SIGNIFICANCE OF
Episphere SEGMENTATION
Prototroch -Hypotheses on the origins of segmentation:
Mouth Gonocoel Theory
Metatroch o Articulate ancestor was not segmented but had
Neurotroch at least one set of organs repeated serially along
Protonephridium the length of its body
Telotroch o These organs enlarged and adopted additional
Anus functions to become the paired segmental
-Metamorphosis coeloms of each segment
Larval episphere (pretrochal region)prostomium Cyclomerism Theory
Body posterior to telotrochpygidium o Ancestor had a few segments (Oligomery)
Growth zone anterior to telotrochSegments that form the thought to be in the form of three pairs, last pair
trunk subdivided to form the segmented trunk
AnusAnus Corm Theory
o Ancestor reproduced clonally to form the zooids,
DIVERSITY OF ANNELIDA similar to some flatworms undergoing paratomy
-Three Major Taxa (Classes in Linnean System): o The zooid lost their independence as they
Polychaeta became more highly integrated into the body as
o Most diverse (8000 spp.) dominates marine segments
environment
Oligochaeta -Selective Advantages
o Including earthworms (3500 spp.) present in Segmentation of the coelomic hydrostat enabled worms to
marine, freshwater and land burrow more efficiently than their nonsegmented burrowing
Hirudinomorpha (Hirudinea) relatives (burrowing hypothesis)
o Leeches (500 spp.) present in marine, freshwater Further specialization of the body; regional specialization of
and land segments (tagmosis)
o Results in restriction of certain segmental
PHYLOGENY OF ANNELIDA structures to only a few segments
-Annelida is divided to two monophyletic sister taxa of equal rank: Gonads are often restricted to few
Polychaeta specialized genital segments
o Have appendages (parapodia) and nuchal o May also result in divergence in structure and
organs function between segments where all segmental
Clitellata structures are retained in all segments
o Absent appendages and nuchal organs o Also results in fusion of segments
o Highly specialized reproductive structures Usually anterior segments, or the
(clitellum, cocoons) associated with a prostomium and peristomium forming
commitment to hermaphroditism and yolk-rich complex head
eggs -Segmentation gene engrailed is expressed similarly in insects and
o Hirudinomorpha (leeches) is monophyletic annelids (leeches)
o Oligochaeta is Suggesting that there is a homologous genetic control of
Paraphyletic (accdg. to P. Ax) segmentation in these two taxa
POLYCHAETAC
- Bristleworms
- Diverse, common, often colourful annelids of the sea
- 8000 species, most are burrowers, but some crawl over the
bottom or on the surfaces of attached organisms
- Others secrete and occupy tubes
- Medium-sized worms of less than 10 cm and 2-10 mm in
diameter
- Eunice aphroditiois body exceeds 3 m in length
FORM AND FUNCTION
- Annelids with legs, called parapodia
o Lateral fleshy outgrowths of the body wall that
may be large and complex or little more than low
nubbins or ridges
o Each segment bears one pair of parapodia
o Basically biramous, each division is supported
internally by an aciculum chitinous skeletal rod
Notopodium - upper division
Neuropodium ventral division
o Parapodial muscles that attach to the acicula
move the parapodia
o Cirri tentacle-like sensory processes, project
from dorsal base of the notopodium and from the
ventral base of the neuropodium
o Notopodia and neuropodia assume various
shapes in different taxa and may be subdivided
into several lobes or even greatly reduced
o Well-developed parapodium is fleshy,
compressed projection of the body wall
- Parapodial lobes (rami) house the chaetal sacs, each of
which secretes a bundle of chaetae
o Two bundles of chaetae are borne by each
parapodium
o New chaetae are produced by chaetael sac as
older chaetae are lost/shed
o Chaetae adopt diverse shapes + chaetal bundles
of a particular species may be composed of more
than one type of chaeta
o Most chaetae are used to improve traction for
locomotion through sediment or over surfaces, so
their tips are variations
o Some are spatulate shovels used for digging,
some are for swimming, others (ucini) bear
numerous microscopic Velcro-like hooks to grip
inner walls of tubes and burrows
- Their segments are generally similar but in some burrowers
and tube dwellers there has been a tendency for trunk to
become specialized into distinct regions (thorax +
abdomen) = result of variations in the parapodia or
presence/absence of gills
o Number of segments range from 10 to over 200
- Polychaete head can have:
o Prostomium only
o Prostomium + peristomium
o Prostomium
o Peristomium and one or more add. Segments
- Prostomial appendages:
o Anterior/anterodorsal antennae
o Anteroventral palps
- in many polychaetes, periostomium beas sensory
tentacular cirri or tentacular palps which are two long
feeding appendages
- polychaete pygidium primitively bears one pair of pygidial
cirri but some have more than one pair
BODY WALL AND TUBES
- body wall is similar to that of annelids in general
- unique among annelids, because many secrete and
occupy a tube
o tube may be open at one or both ends + partially
buried in sediment or attached to surfaces
o worm may permanently occupy its tube enlarging
it with growth (Chaetopterus feather-duster
worms)
o may abandon the tube and crawl for a new
location and secrete another (Diopatra)
o Owenia fusiformis, is able to burrow through the
sediment in its flexilbe tube
- tube material is a fibrous protein that has appearance and
texture of cellophane, parchment or silk
o foreign material is often incorporated into the wall
of the tube for strength, camouflage and food
o some calcify the tube to form a shell (Serpulidae,
Spirorbidae)
- tube is like a lair from which to emerge + catch prey
(Diopatra)
- tube may provide a source of clean, oxygenated water to
its subterranean occupant
- few worms brood their eggs and young within the tube
- Chaetopterus pumps water through its tube for gas
exchange and filter feeding
MUSCULATURE AND LOCOMOTION
- Peristaltic burrowing is common to those having elongate
bodies + reduced parapodia and head appendages and
many similar segments
- Circular musculature is well developed and septa are
complete, restricting the coelomic fluid to individual
segments
- Active burrowers resemble earthworms and are
represented in Scolecida
- Other burrowing polychaetes augment or replace whole-
body peristalsis with an eversible pharynx
 o As it everts, it pushes into the sediment and
anchors
o Animal then pulls itself forward into the sediment
as it retracts the anchored pharynx
o Glycera (blood worm) has abandoned peristalsis
and burrows solely with its prodigious pharynx
which punches into the sediment
o Armandia and Opthelia produce rapid lateral
undulations of the body + swim through loosely
consolidated sand
o Very small sediment dwellers use cila for
locomotion (Dinophilidae and Diurodrilidae)
- Many polychaetes either crawl over surfaces or swim using
well-developed parapodia and chaetae (Nereidae,
Phyllodocidae, Polynoidae, Alciopidae, Tomopteridae)
o Prostomium is equipped with eyes and other
sense organs
o Parapodia well-developed, segments are
generally similar
o Movement = result of the combined action of the
parapodia + body-wall musculature + coelomic
fuid skeleton
o The longitudinal layer is better developed than
the circular layer
o Specialized parapodial muscles are well
developed
o Incomplete septa
- During slow crawling, parapodia and chaetae move like
legs, alternately pushing against the substratum in the
power stroke + lifting above the substratum in the recovery
stroke
o As the power stroke begins, the acicula and
chaetae extend, the parapodium makes contract
with the substratum and then sweeps rearward
o Recovery stroke is inititated as the acicula and
chaetae retract, then the parapodium lifts off the
substratum and swings forward
o Movements of the parapodia are coordinated to
avoid interference + effectively move the worm
o Parapodia on opposite sides of same segment
are 180 degrees out of phase if
the left-side = power stroke
right side = recovery stroke
o the series of parapodia are coordinated in waves,
like falling dominoes that pass from posterior to
anterior (progressive wave)
o when parapodium in the middle of its power stroke
the next anterior parapodium completes its power
stroke as the next posterior parapodium completes
its power stroke = similar to crawling millipede
- Rapid crawling involves parapodia + lateral body
undulations produced by waves of longitudinal-muscle
contraction
o These progress forward along the body +
coincide with the alternating waves of parapodial
activity
o Parapodial power stroke occurs at the crest of an
undulatory wave
o With the crest parapodia fimly planted to the
substratum, contraction of longitudinal muscles
on the opposite, through side of the body imparts
additional thrust to the crest-side
o As the longitudinal muscles shorten + compress
the through side, the crest side expands thrusting
back against the parapodia anchored in the
substratum = pushing the animal forward
- Benthic polychaetes (Naeris) swim using the rapid-crawling
movements
- Members of Alciopidae have enormous eyes
- Species of Tomopteridae (no chaetae) have membranous
parapodial pinnules = most highly specialized of the
pelagic polychaetes
NERVOUS SYSTEM AND SENSE ORGANS
- Brain and ladderlike nerve cords are like the general
annelids
- Polychaete brain = large and lobed if the head bears sense
organs
- Pedal ganglia = additional ganglia; commonly associated
with the segmental nerve cords at the bases of the
parapodia
o These are important centers that control the
complex parapodial movements
- Ventral nerve cord may be paired + ladderlike
o In Owenia, unpaired and nonganglionated nerve
cord occurs in th epidermis
- Chief sense organs:
o Nuchal organs
o Ocelli
o Statocysts
- Nuchal organs pair of ciliated sensory pits or slits
situated posteriolaterally on the prostomium
o Widespread; unique to polychaetes
o chemoreceptive organs; important for detecting
food
o Amphinomidae they expand to form a
convoluted, brainlike crown (caruncle) on the
upper surface of the head
- Eyes (ocelli) which are best developed in errant
polychaetes (Aciculata) are found on the surface of the
prostomium in two, three, or four pairs
o They sometimes occur elsewhere on the body
Feathers of feather-duster worms have
a shadow reflex and instantly withdraw
into their tubes when a shadow is cast
over them
o Prostomial ocelli are pigment cups
 Walls of which are
composed of rodlike
photoreceptors (modified
microvilli) + pigment cells
+ supporting cells
o Determines light intensity and direction
Acliopidae have bulging
eyes that may be capable of
image formation
- Statocysts predominate among sedentary burrowers and
tube dwellers
o Lugworm Arenicola = has one pair of cilia-lined
statocysts embedded in the body wall of the head
each opening to the exterior via lateral canal
Contains spicules + diatom shells +
quartz grains = all coated with chitinous
material
Always burrows head downward
Worm makes a compensating 90 turn
in burrowing if an aquarium containing
a worm is tilted 90
If statocysts are destroyed =
compensating ability is lost
- Sensory appendages of:
o Prostomium (antennae, palps)
o Peristomium (tentacular cirri)
o Parapodia (dorsal + ventral cirri)
all bear numerous sensory cells
- Each appendage is thought to house both
mechanoreceptors + chemoreceptors, but distinct sensory
roles for the individual appendages have not determined
- Some mechanoreceptors may be
o Tactile
o Sensitive to water flow
o Dendrites are embedded in the cuticle, may be
stretch receptors (proprioceptors)
- Some chemoreceptive cells detect pheromones released
prior sexual reproduction
DIGESTIVE SYSTEM
- Gut is typically a straight tube extending from the mouth at
the anterior end of the worm to the anus on the pygidium
- Digestive system is like that of general annelids
o Commonly differentiated into a pharynx (buccal
cavity if no pharynx) + short esophagus +
stomach (in sedentary species) + intestine +
rectum
- Pharynx can be:
o Protrusible, tonguelike muscular bulb
Situated in the midventral wall of the
foregut
o Eversible organ
Teeth of various forms and functions occur in the
pharynx
o In Naeris, two large glandular ceca open into
the esophagus
o The ceca secrete digestive enzymes
- In worms that crawl or burrow = feces are simply released
and abandoned
- Chaetopterus pump water unidirectionally through their
tubes defecate into the downstream exhaust flow
- Runs foul when only one end of the tube/burrow is exposed
to the surface water, therefore polychaetes live upside-
down
- Many invert themselves temporarily to defecate at the
surface
- Feather-duster worms and mason worms live upright in
blind-ended tubes
o They vent their feces without standing on their
heads
o Mason worms bends their body into a U
- Dorsal body of feather-duster worms has a ciliated out of
the tube
- Many species consolidate their feces into high-density fecal
pellets or strings, which tend not to resuspend and reenter
their burrows or tubes
NUTRITION
- Direct deposit feeders ingest sediment directly with the
mouth or a nonmuscular, bulbous, protrusible pharynx
(lugworms - Arenicolidae, bamboo worms- Maldanidae,
Orbiniidae, Capitellidae)
- Indirect deposit feeders feed on organic material in
sediments but collect material first with a specialized
appendage that conveys food to the mouth
o Appendages are extensible prostomial tentacles
(spaghetti worms)
o Palp worms use a pair of long peristomial palps
- Carnivores + herbivores + scavengers are motile (errant)
polychaetes that crawl over surfaces and in crevices or
catch plankton (Alciopidae)
o Some are tube dwellers or active burrowers
o They typically have a well-developed muscular
developed pharynx
o Everted pharynx may be
muscular grasping tube
(paddleworms)
thick rasping lower lip (fireworms)
bear jaws with grasping (Nereidae,
Eunicidae, Lumbrineridae)
venomous teeth (Gylceridae)
- horny jaws are composed of a cross-linked (tanned) protein
- when food is detected pharynx rapidly everts exposing and
opening jaws
o food is then seized then pharynx is retracted
- protractor muscles may be present, but increased coelomic
pressure everts the pharynx
 o when it relaxes + coelomic pressure drops =
pharynx is withdrawn by retractor muscles
- all suspension feeders are sedentary animals
o they occupy tubes in sediment/attached to
shells/rocks and other hard surfaces
o presence of ciliated appendages with large SA
for trapping suspended particles (feathery crown
of feather-duster worms)
o they secrete a mucous net through which water
is pumped + filtered (Chaetopterus)
o Labrorostratus lives in the coelom of other
polychaetes + may be as big as host
o Ectoparasites attach to fins of marine eels
(Ichthyotomidae)
GAS EXCHANGE
- Occurs across general body wall
- Additional specialized gills are present
o Esp. in large-bodied burrowers + tube dwellers
o Always unprotected outgrowth of the body
surface
o Never enclosed in gill chambers
o Absent in tiny polychaetes + those with long
threadlike bodies
- Gills are associated with the parapodia
o Notopodium may be flattened to form branchial
(gill) lobe in Nereids
o Dorsal cirrus of parapodium = modified as a gill
o Gills can arise from base of dorsal cirrus
- Scaleworms gas exchange is largely restriceted to dorsal
body surface
o Roofed by elytra overlapping fishlike scales
Each of which is borne on a short stalk
- Cilia on the dorsal body surface create a current of water
beneath the elytra
o Aphrodite lacks cilia but similar water current is
produced by rocking its elytra
- Gills of many sedentary species are branched feather or
treelike structures restricted to the anterior end of the body
+ independent of parapodia
o Gills of spaghetti worms
Spectacular red absorbent gills
Ciliated + contractile
Pulsate rhythmically
o Feather duster of feather duster worms function
as gills + filter for feeding
- Ciliary beating + muscular contraction = gill ventilation
- Worms that ventilate using peristalsis = exhibit a
spontaneous ventilating rhythm in which it alternates with a
period of rest
- Muscular exertion associated with peristaltic ventilation =
increase the worms O2 consumption by 15-fold but
approx. 20-fold increase O2 uptake
INTERNAL TRANSPORT
- Circulation results from fluid movement in hemal system +
coelom
o Reduced septa (glyceride bloodworms)
Coelomic system is replaces hemal
system + transports substances
throughout the body
- Basic layout of hemal system is similar to general annelid
but complicated
o By addition of parapodial + gill circulations
- Ventral vessel gives rise to:
o parapodial vessels
transport blood to parapodi +body wall
+ nephridia before returning to dorsal
vessel
o several pairs of intestinal vessels
deliver blood from gut to ventral vessel
- capillary plexus in parapodia + gut wall = facilitates
exchange in p. vessels and i.vessels
- gills have afferent + efferent vascular loops
o permitting two-way flow
lugworms
notopodal lobes of Nereids
- feathers of feather-duster worms contain a single vessel
which blood flows tidally in and out
- Glyecerids parapodial gills are irrigated with coelomic
fluid instead of blood
- Blood is colorless in small polychaetes
- Hemoglobin occur in plasma in larger species
- Respiratory pigments are diverse
- They have three to four animal respiratory pigments
o Haemoglobin most common
o Chlorocruorin slight chemical difference
makes it green instead red
o Hemerythrin pink or violet hue (Magelona)
binds an atom of iron at its center
nonheme (lacks porphyrin)
- coelomic fluid is circulated by muscular contractions of the
body wall or by muscular contractions and cilia on the
coelomic lining
o may be colorless or
o contain low-molecular weight haemoglobin in
coelomocytes
pigment sometimes occur only on the
coeleom (Capitellidae) not in the hemal
system
terebellids and opheliids
two different hemoglobins
present: one in blood and
one in the coelom
Glycera + most scaleworms
o Have bright red hemoglobin
in/around their nerve cords
 Scaleworms
Myoglobin occurs in muscles
- Oxygen-binding properties of pigments differ when
haemoglobin is present in hemal + coelom
o Hemal vessel (gill) least tight
o Muscle + nerve most tight
o Coelomic fluid - intermediate
= results in a cascade of oxygen transport from gill
vessels -> coelom -> oxygen-demanding cells
- Hemal system haemoglobin binds oxygen at the gill to be
stripped away by the higher affinity coelomic haemoglobin
has its oxygen removed by the higher affinity nerve and
muscle haemoglobin
- Mitochondrial cytochromes highest affinity; final
destination for respiratory oxygen
- Oxygen availability drops as tide receds + water stagnates
o Polychaetes may suppress their metabolic rate =
become torpid
o Rely on stored oxygen bound to haemoglobin
or both
EXCRETION
- Filtration nephridia polychaete excretory organs
- May play accessory roles in excretion:
o Chlorogogen tissue
o Coelomocytes
o Intestinal wall
- Protonephridia occur in few adult polychaetes that
lack/have reduced hemal system
- Most polychaetes have hemal + metanephridial systems
- Correlation of blood vessels with metanephridia + absence
with protonephridia = how ultrafiltration occurs in two
functional groups
- Polychaetes with metanephridial systems have vascular
sites covered with podocytes + septum associated
metanephridia
- Nereis bears ciliated funnel-like nephrostome
- the long postseptal tubule forms a coiled mass enclosed in
a thin mesothelial sheath
o tubule elongation + coiling = adaptations that
increase SA for tubular secretion or reabsorption
o nephridiopore opens at the base of the
neuropodium on the ventral side
o entire lining of tubule is ciliated
- metanephridia of most other polychaetes differ slightly from
Nereis but may be regionally restriced to excretory
segments
o feather-duster worms one pair of anterior
nephridia
ducts of two nephridia unite in the
ventral midline + open through a single
nephridiopore on the head
worm releases urine directly to
exteneral environment + avoid fouling
- solenocyte each terminal cell in the cluster of cells in
protonephridium
o closely resembles a sponge choanocyte
- many polychaetes can tolerate low salinities
o worms eliminate excesss water and take up salt
(have glomerulus-like network of blood vessels)
o Nereis succinea - gills contain cells for actively
absorbing ions
o Manyunkia speciosa colonizes freshwater +
widely distributed
REPRODUCTION
Regeneration and Clonal Reproduction
- Regeneration is common in burrowers and tube dwellers
- Potential regeneration is greater in worms with uniformly
differentiated trunks than those with specialized thoracic +
abdominal regions except:
o Chaetopterus and Dodecaceria regenerate at
the entire body from a single segment
- Nervous system plays an important role in regeneration
o Neuroendocrine is involved in some way
- Clonal reproduction occurs in some
o Cirratulids (Dodecaceria)
o Syllids, feather duster worms (Sabellidae)
o Palp worms (Spionidae)
- Budding + transvers fission = two common modes of
asexual reproduction
Sexual Reproduction
- Majority of species are gonochoric
- Gonads are distinct
- Segemented paired organs found in the connective tissue
with septa + blood vessels + coelom lining
- Most segments bear gonads in many polychaetes but
some are restricted to genital segments
- Gonads are usually limited to the abdomen in polychaetes
with distinct thoracic + abdominal regions
o Feather duster worms have anterior abdominal
segments = produce eggs; posterior produce
sperm
- Germ cells are shed into the coelom as
gametogonia/primary gametocytes
- Growth and differentiation of the gametes is completed in
the coelomic fluid
 o When worm is mature, coeleom is packed with - Odontosyllis enopla swarms in the summer beginning 3
ripe eggs or sperm = can be seen through the days after the full moon and 56 min after sunset
body wall - Female swims in circles at the surface while emitting
o Pomatoceros bright pink or orange bioluminescent glow that attracts males
o Proceraea fasciata red-banded whitish body - Males are called blinkers swim rapidy to the female
- Gametes are either spawned through the metanephridia or - They then spew gametes in a stunning luminescent finale;
by rupturing of the body wall spent female and male return to the bottom and convert
o Body-wall rupture is common to species that back to atokes
transform into pelagic swarmers
- Palolo has swarming lunar periodicity
Epitokes Nereidae, Syllidae,
o Palolo referred to Palola virdis
Eunicidae
o Occupies rock and coral crevices below the low-
o Rupture is typical to shimmy worms (Nephtyidae)
tide mark -> releases epitokes in October or
and bloodworms (Glyceridae)
November
Protonephridia present as adults
- Female epitokes can exceed 1mm in length
Epitoky - Eunice schemacephala negatively phototactic ; emerges
from the burrow to feed only at night
- Epitoky formation of a pelagic reproductive individual - Swarming takes place in July near last quarter of a lunar
called epitoke from a benthic nonreproductive atoke cycle
o At 3 or 4 o clock am, worm backs out of its
- Coincides with gamete production + sexual maturity
burrow
- Involved changes in nonreproductive structures that adapt
o Caudal epitokal region breaks free
the worm for swimming and allow it to detect mate
o Epitokeheads for the surface and swims slowly,
o Eye enlargement
spiralling
o Parapodia modification + chaetae for swimming
o Segment enlargement s
Endocrine Control of Reproduction
o Segmental musculature enhancement
- One or more epitoke can arise from the atoke either by
- Reproductive events are regulated by hormones
metamorphosis of the entire individual (Nereids), or
o Neurosecretions produced by the brain
- By differentiation + separation (budding) from the posterior o Syllids nervous elements of the sucking foregut
half of the body (Eunicids and Syllids)
- Nereids and Syllids hormone regulates entire
- Gamete-bearing segments of epitokes are modified = body reproductive state (production + epitoke development)
of worm divided into different regions
- Hormone is required for gamete development in worms
o Nereis irrorata and N. succinea
that breed more than once
Have large eyes and reduced
prostomial palps + tentacles
Oviposition
o Palola (Eunice) viridis anterior end is
unmodified and epitokal region consists of chain
- Many shed their eggs freely into the seawater
of egg-filled segments
- After fertilization they become planktonic
Swarming - Some retain their eggs within their tubes or burrows or lay
them in jelly masses
- Epitokes swim to surface simultaneously and shed eggs - Axiotella bamboo worm produces an ovoid egg mas
and sperm resembling a large transparent grape that is attached to the
- Female produces a pheromone that attracts male and chimney of tube
stimulates shedding of sperm - Many brood their eggs
- Sperm then stimulates shedding of eggs o Spinoids and Serpulids brood in their tubes
o Spirorbis cavity of operculum
- Autolytus swims in circles around the female touching her
o Autolytus secreted sac attached to the ventral
with antennae + releasing sperm
surface of the body
- Light cues are often important in synchronizing swarming
o Nereis limnicola gestate eggs in coelom
- Autolytus edwardsi induced by changes in the light
intensity dawn and dusk to leave the bottom and swim to
the surface
- May also be affected by lunar periods
Development and Metamorphosis Population Biology

- Gastrulation takes place by invagination, epiboly or both - Burrowing and tube-dwelling polychaetes occur in
- Embryo rapidly develops to top-shaped trochophore larva enormous numbers on the ocean floor
after gastrulation - Compose a major part of the soft-bottom fauna
o Owenia, Polygordius, Phylodocidaem Serpulidae - Density of polychaetes in Tampa Bay, Florida is 13,425 per
(greatest development attained from these square meters
species) - 40 to 80% of the infauna in upper continental slope + deep
- Trochophores of some are lecithotrophic and their short ocean floor
larval existence is spent near the bottom - Euzonus (Thoracophelia) mucronatus occur in dense
- Trochophore -> juvenile body aggregations on the protected intertidal beaches along the
- Gradual lengthening of the body = most conspicuous pacific coast of US
feature of metamorphosis - Number of worms averages 2500 to 3000 per 30 square
o Pretrochal region + apical plate form the meters
prostomium and the brain
- Worms occupying a strip in a beach 1.5 km long 3 m wide,
Post-telotrochal region = pygidium
and 30 cm thick ingest ~ 13,270 metric tons of sand each
- Metamorphosis may result in the immediate termination of year
planktonic existence - Such populations do not appear to be limited by food
o Metamorphosing stages of Spionids, Sabellariids, resources
Oweniids have greatly enlarged + erectile +
- Predation and other pressures prevent annelid, mollusc
protective anterior chaetae
and other infaunal populations from ever reaching the
- Some have their trochophore stage passed in the egg prior carrying capacity of the habitat
to hatching
- Areas in York River of the Chesapeake Bay were protected
o Less drastic because larval structures never fully
from fish and crabs by wire cages
develop
- = half of polychaetes increased from 2 to many times
o Autolytus elongate larva breaks free from the
brood sac
o Axiotella mucosa and Scoloplos armiger direct
development +juveniles from jelly capsule
assume immediate adult mode
- Developmental patterns:
o Annual species
Live only 1 to 2 years and spwan only
once
Produce large number of small eggs
Well-developed feeding larvae
(planktonic for a week or more)

o Perrenial species
Live and breed for more than a year
Produce small numbe of large yolky
eggs
Nonfeeding benthic larvae
o Multiannual species
Short life spans
Several generations in one year
Produce small batches of large yolky
eggs
Nonfeeding benthic larvae
DIVERISTY OF POLYCHAETA o Proventriculus specialized muscular part of
Scolecida the pharynx used for sucking out the contents of
the prey
- No prostomial appendages - Family Oenonidae (formerly Arabellidae)
- Two or more pairs of cirri on pygidium - Family Lumbrineridae
- Most are burrowers with bulbous protrusible pharynx - Family Ichthyomidae
- Sediment directly ingested
CANALIPALPATA
- sedentary polychaetes

Family Arenicoliae - Sister taxon of Articulata

Family Maldanidae - sedentary species in tubes/ semi-permanent burrows
Family Capitellidae - Longitudinally grooved palps on prostomium
Family Orbiniidae o Used for suspension/deposit feeding
Family Opheliidae - In some, palps have shifted to the peristomium

PALPATA Order Spionida

- Sister taxon of Scolecida - Family Spionidae

- Presence of sensory palps on the prostomium - Family Magelonidae
- Family Chaetopteridae
ACICULATA
Order Terebellida
- Biramous parapodium
- One aciculum at least in the notopodium + neuropodium - Family Terebellidae
- Dorsal + ventral cirri present - Family Ampharetidae
- Four, three or two antennae - Family Cirratulidae
- Formerly called errant - Family Pectinariidae
- Family Flabelligeridae
Order Phyllodocida
Order Sabellida
- Family Phyllodocidae
- Family Tomopteridae - Family Oweniidae
- Family Alciopidae - Family Sabellariidae
- Family Nereidae - Family Sabellidae
o Nereis o Radioles bipinnate tentacles
- Family Glyceridae o Pinnules side branches of each
- Family Nephtyidae o Sabella
o Nephtys
o Shimmy worms Order Pogonophora
- Family Aphroditidae + Polynoidae + Polyodontidae +
Sigalionidae - Opisthoma posterior end of the body; short and bulbous
o Elytron each scale has this highly modified o composed of several short segments
dorsal cirrus o anchor the worm in its tube (together with the
- Family Hesionidae chaetae) allowing it to withdraw rapidly

Order Myzostomida
Order Eunicida
- Family Eunicidae
- Family Amphinomidae
- Family Onuphidae
- Family Syllidae
CLITELLATA
-clitellate annelida or girdle worms
-includes earthworms, small inconspicuous worms, leeches and Body wall and Coelom
relatives
-all LACKS parapodia, head and pygidial appendages Body wall is similar to polychaetes
-have Clitellum, a series of anterior segments enclosed in a thick, Cuticle
glandular epidermis that often forms conspicuous girdle around the Overlies an epidermal layer that contains
body mucous gland
Circular muscle
either encompasses the female gonopores or is located Longitudinal muscle
behind them Four bands
secretes mucus for copulation, nutritious albumen for eggs, Septa
and a cocoon in which eggs and albumen are deposited Relatively complete
Some earthworms with well developed septa, can have
-all are copulating hermaphrodites
-gonads are always restricted to few genital segments with the testes sphincters
anterior to ovaries Located around septal perforation
-zygotes develop in the cocoon and emerge as juveniles Central flow of coelomic fluid from one
-direct development segment to another
-brain has shifted posteriorly from the prostomium to an anterior trunk Coelomic fluid
segment Keeps integument moist
Deter predators
Coelomopore
OLIGOCHAETAc Connects coelomic compartment to the outside
Located mid dorsally in the intersegmental
Largest Oligochaeta burrows
Earthworm Provided with sphincter
Including Australian earthworm, Megascolides Giant Australian worm
australis (3m long) Eject fluid from their pores to a height of 10 cm
Squirter worm
Form and Function Eject fluid to a height of 30 cm
Didymogaster sylvaticus
Similar to thatgeneralized annelid
Well-developed segments Locomotion
Four bundles of chaetae per segment
Small prostomium and small pygidium (both Oligochaetes crawl and burrow by peristaltic contraction)
are devoid of appendages) Body shortens
Oligochaete chaetae Chaetae are extended
Simple and terminate in a needlelike point or Longitudinal muscle contraction
tips that are bifid, pectinate, different from the Body elongates
shaft Chaetae retracts
More complex in genital segments Circular muscle contraction
Chaetal sac Contraction waves can be reversed
Found each side of segment Enables worm to move backward
Which chaetae are secreted Fewer chaetae than in polychaetes
Which chaetae emerge as bundles Species of microscopic aelosomatids
Chaetal bundles Swim by ciliated prostomium
Two are ventral
Two are ventrolateral or dorsolateral Nervous system
1-25 number of chaetae per bundle
In Lumbricus Two nerve cords
Eight chaetae per segment Fused in the ventral midline
Two chaetae in each of the four bundles Inside the muscle fiber of body wall
Fewer chaetae than in polychaetes Brain
Protractor and retractor muscles Shifted posteriorly
Allow chaetae to be extended or withdrawn In Lumbricids
Lies in the third segment above
anterior end of pharynx
In ventral nerve cord
 5 giant axons Used for grinding food
3 large diameter in dorsal part particles
2 smaller diameter in ventral part Lined with chitinous cuticle
Mid dorsal axon Very muscular
Fired from sensory input from the head Crop
Two dorsoventral fibers Occurs in lumbricid
Fired from sensory input from the posterior end earthworm
of the body Storage chamber
Sensory input from the other end of the worm illicit a Esophageal wall
response however the response is more rapid if the Houses calciferous gland
head is grasped or disturbed Secrete calcium carbonate to
Subpharyngeal ganglion esophageal wall in a form of calcite
Center of motor control and vital reflexes crystals
If this is destroyed, movement will cease Calcite crystals are then transported
into the gut and pass out as feces. It
Sense Organs is not reabsorbed
Eliminates CO2 via the gut and
Lacks eyes excess calcium taken in with food
Intergument has dispersed unicellular photoreceptors Ciliated intestine
located in inner part of epidermis, dorsally at anterior Anterior half of the intestine
end Principal site of enzyme secretion
Tubercles Posterior half of the intestine
Form three rings around each segment Absorptive
Numerous on the most anterior segments, Intestinal epithelium
especially in the prostomium Has cellulase (digest plant cell wall)
and chitinase (digest fungal cell wall)
Nutrition and Digestive system Absorbed food material are passed to blood
sinuses
Feed in dead organic matter Typhlosole
In Aelosoma Increase surface area of the intestine
Collect detritus with its prostomium Projects internally from mid dorsal
In Chaetogaster wall
Feed using sucking action of pharynx Chlorogogen cells
Eat amoeba, ciliates, rotifer, and trematode Yellowish mesolecithal cells
larvae
Hemal system
Similar to annelids in general
Oligochate digestive tract
Have integument capillaries
Straight and simple
Heart
Mouth
Supplement dorsal blood vessel
Beneath prostomium
In Lumbricus
Opens into small buccal cavity
Five pairs of heart surrounds the esophagus
Opens into a spacious
In Tubifex
pharynx
Has one pair of circumintestinal heart
Mid dorsal pharyngeal bulb
Principal ingestive organ
Gas Exchange
In aquatic species, it is everted and collects
particles on its adhesive surface Gas exchange is through diffusion of gases through
Pharynx body wall
Acts as pump Mucous gland secretions and fluid through
Pharyngeal glands produces salivary secretion ceolomopores moisten the surface epidermis and
containing mucus and enzymes facilitate gas exhange
Opens into narrow, tubular, esophagus In Dero and Aulophorus
Esophagus may be modified to form gizzard or Have circle of fingerlike gills at posterior end of
crop its body
Gizzard In Branchiura
 Has gill filaments dorsally and ventrally in
In Nais and Aelosoma
posterior quarter of the body Reproduce by paratomy
Members of Tubificidae Differentiation precedes separation of
Tolerates low oxygen or even complete lack of individuals, resulting in chains of
oxygen individuals called zooids
Tubifex tubifex Sexual Reproduction
It can die due to long exposure of Reproductive system
ordinary oxygen tension Hermaphroditic
Genital segments are found in the
Excretion and Diapause anterior half of the worm
Female segment are always behind
Have metanephridial system the male segment
Each segment has one pair of metaphridial tubules per Paired ovaries and testes occur on
segment except at extreme posterior ends the posterior face of a genital septum
Segment following the nephrostome, the tubules is Gametes are released from gonadsat
greatly coiled an early stage of development and
Holonephridia enter seminal vesicles (male) and
Single typical pair of nephridia ovisacs (female)
Found in each segment Gonoducts
Enteronephridia Sperm ducts (male) and
Open into the various parts of the gut oviducts (female)
In terrestrial species, it appears to be an In males, sperm ducts
adaptation for water retention usually fuse before opening
By first releasing the urine in the to the outside via single
digestive tract, much of the remaining male gonopore
water can be reabsorbed in the Prostate glands
intestine en route to anus Glandular tissues
Worms with enteronephridia can tolerate drier associated with male
soils and do not have to burrow so deeply gonoducts
during dry periods Absent from most lumbricid
Found in each segment earthworms
Ammonia remains an important excretory product Seminal Receptacle
Earthworms also secrete urea. Its level of secretion Sac for storing partners
depends on the condition of the worm and the sperm prior to fertilization
environmental situation Testis sac
Salt and water balance is as well regulated by the In Lumbricus
nephridia Special ventral
Urine is hyposmotic compartment that encloses
Reabsorption of salt must take place as fluid passes the testes, sperm duct
through nephridial tubule funnel, and the opening of
Few aquatic oligochaetes are capable of encystment by the seminal vesicle
secreting tough mucous covering that forms the cyst Completely separate from
wall the remaining portion of
coelomic activity making the
Reproduction and Development testes not visible in the
usual dorsal dissection
Clonal Reproduction Aelosomatidae
Common in aeolosomatids, naidids, and Lack distinct gonad and
lumbriculids have large number of
Other oligochaetaes reproduce asexually in the genital segments
summer and sexually in the fall Absent gonoduct
Clonal reproduction is always transverse Sperm use nephridia to exit
division (fission) of the parent worm into two or the body
more new individual
In Lumbriculus
Clitellum
Reproductive structure of clitellates
Reproduces by fragmentation
Fission precedes differentiation
 2 clitellar segments in aquatic forms,
6-7 in Lumbricus, 60 in
Glossoscolecidae
In aquatic species, clitellum is usually
in the same region as the female
gonopes
In lumbricids, gonopore in located at
the posterior of the female gonopore
Copulation
Less continuous breeders in contrast
to polychaetes
During copulation, ventral contact is
established between anterior ends of
the two oppositely facing worms
Most oligochaetes except the
lumbricids, male gonopore directly
appose the seminal receptacle of
another worm
Held in a position by a coat
of mucus secreted by
clitella
Genital chaetae hooks them
together
Genital Chaetae
Specialized ventral
segmental chaetae
Situated along the male
gonopore region or seminal
receptacle
In the lumbricids, male gonopores do
not appose the seminal receptacle
instead the sperm must swim
externally from the anterior male
gonopore to the posterior pores of
the partners seminal receptacle
In lumbricids copulation, the clitellum
of one worm presses against the
seminal receptacle mucus
secreted enveloped in a slimy
tube- attachment by genital
chaetae sperm are released
together with the contraction of
specialized muscles created
grooved in the body wallbetween
male gonopore and clitellum-
sperm moves along the grooves
clitellum- cross over to the other
worm - enter seminal receptacle
Cocoon
Secreted by the clitellum after
copulation for the deposition of eggs
Mucus tube is secreted around the
anterior segments including the
clitellum clitellum secretes chitin-
like material that forms the wall of
cocoon deep glandular layer of
clitellum secretes albumen into the
space between the wall of cocoon
and clitellum
Albumen-filled cocoon slides forward
as the worm moves backwards
As it passes the female gonopore,
eggs are discharged in the cocoon
moves to the seminal receptacles
cross fertilization cocoon slips
over the head of the worm freed
from the body mucous tube
disintegrates cocoon constricts
and seal themselves
Finished cocoon contains zygote and
nutritive albumen
In contrast to polychaetes, hormones
produced by the brain appears to
stimulate reproduction
Development
Direct developers
Have yolky eggs
Terrestrial group have smaller eggs
and lesser yolk because of the
albumen which provides the embryo
nutrients
Larva is absent
Development into juvenile stage
occurs in the cocoon
Juvenile emerges from the end of
cocoon after 8 days or several
months of development
Diversity of Oligochaeta
AelosomatidaeF
Aquatic
Less than 5 mm
Ventrally ciliated prostomium
Chaetae present in most segment
Segmental septa is absent
Sexual reproduction: rare; Clonal reproduction:
common
Aelosoma
NaididaeF
Aquatic
Few segments
Prostomium is not ciliated
Chaetae is absent
Segmental septa is present
Some have long prosboscis
Paratomy is common
Stylaria, Chaerogaster, Nais
TubificidaeF
Tubifex
LubriculidaeF
Many segments ith four pairs of chaetae
Clonal reproduction by fragmentation
Lumbriculus variegatus
HIRUDINOMORPHAC
-Clitellate taxon includes tiny Branchiobdellids
(Branchiobdellida), which cling and creep over the exoskeleton
of crustaceans and leeches (Hirudinea). Leeches are common
in freshwaters and can occur in moist land.
-ALL hirudinomorphs have a terminal or posterior sucker that
adheres to host or the substratum and a dorsal anus located in
front of the sucker. The re-positioning of the ancestral anus
from terminal to dorsal is related to the origin of the sucker
specialization of the posteriormost segment as well as the
pygidium.
-Branchiobdellids and especially a leech called Acanthobdella
peledina, are important in the Hirudinomorph evolution.
Euhirudinea
-marine, freshwater, terrestrial worms commonly known as
leeches.
-popularly considered to be bloodsuckers, but many are non-
bloodsucking carnivores.
-leeches share clitellate traits with Oligochaeta.
-leeches never as small as polychates/oligochaets.
-Giant Haementeria ghiliani
-few tolerate rapid currents,
most prefer shallow environment
FORM and FUNCTION
-body typically dorsoventrally flattened and frequently tapered
at the anterior end.
- Segments at both extremities have been modified to form
suckers.
-Anterior sucker is usually smaller than the posterior and
frequently surrounds the mouth,
-Posterior sucker is disc shaped and turned ventrally
-Segmentation is very much reduced
-Unlike other annelids, leeches have fixed number of 33
segments, but secondary superficial annulations externally
mask the segmentation
-Chetae is ABSENT.
-Head consist of reduced prostomium plus 5 segments. (1-5)
-Dorsally, head bears several ocelli, and ventrally modified to
form the anterior sucker surrounding the mouth.
-Trunk, encompasses the preclitellar region, clitellum and
postclitellar region consist of 21 segments. (6-26)
-Behind the trunk is large, ventral posterior sucker derived
from 7 segments (27-33)
-The number of annulation per segment varies not only in
different regions but also in defferent species.
NERVOUS SYSTEN and SENSE ORGANS
-similar to other annelids but anterior and posterior ganglia are
concentrated into masses because of segmental modifications
forming the suckers.
-the brain, located in segment 5, consist of paired
subpharyngeal gangliea (original paired ganglia of prostomium
+ peristomium).
-ventrally, first 4 pairs of gangliea (2-5) are fused to form
subpharyngeal ganglion.
-entire nervous system enclosed in the unpaired ventral
coelomic channel.
-some Rhynchobdellid leeches can change color dramatically
as a result of pigment movement in large specialized cells
called chromatophores under neural control
-sensory papillae small projecting discs arranged in a
dorsal row or in a complete ring around 1 annulation of each
segment,
-Despite lack of highly organized organs, leeches can detect
low levels of many types of stimuli.
BODY WALL, COELOM, HEMAL SYSTEM,
LOCOMOTION
-consist of cuticle and epidermis
-UNLIKE polychaetes and oligochaetes, Fibrous connective
tissue beneath the epidermis is very thick.
-dorsoventral muscles also present
-Leeches LACK paired segmental coeloms that typify other -leech digestion is peculiar: gut secretes no amylase, lipase, or
annelids. (Acanthobdella peledina is the only exception) endopeptidase

-Much of coelomic mesothelium is specialized as large -presence of only exopeptidase explains the fact that digestion
nutrient-storage cells called Chlorogogen tissue in in bloodsucking leeches is so slow.
rhynchobdellids and bortyroidal tissue in archynchobdellids.

-LOSS of septa, chatae, and segmented coelomic cavities

REPRODUCTION
-Leeches NOT produce asexually NOR can they regenerate
GAS EXCHANGE lost parts.

-Gills occur only in Piscicolidae (fish leeches) -Hermaphrodites

-General body surface procide gas exchange for other leeches -Protandric, not simultaneous. Testes matures before ovary

-Piscicolid gills are latral leaf like or branching outgrwths of the -accessory reproductive organs are incorporated into the
body wall. sperm ducts and oviducts unlike in oligo, separate sem vesicle
and receptacle.
-Respiratory pigment, hemoglobin, found only in
arhynchhobdellid leeches. -INTERNAL fertilization

-hypordermic impregnation

EXCRETORY SYSTEM -following liberation and spermatophore, sperm migrate to the

-10-17 pairs of Metanephredia: 1 pair per segment
-Main duct typically expands into a urinary bladder, before
opening to exterior.
ovisacs via coelomic channels or use a tissue pathway called
target tissue, specialized to receive spermatophore and
conduct sperm

DIGESTIVE SYSTEM AND NUTRITION

-either a protrusible pharynx (called Proboscis)

or a nonprotrusible sucking pharynx with or without jaws.

-Tubular pharynx (rhnchobdellida) joins the ventral mouth by a

short narrow canal.

-pharynx is highly muscular and has triangular lumen and lined

with cuticle

-When feeding, animal protrudes the pharynx from the mouth

forcing it into the tissue of the host.

-immediately witin mouth cavity, of most species are 3 large

oval bladelike jaws, each bearing many small teeth along the
free edge, arranged in a triangle, 1 dorsally and 2 laterally.

-Jaws swing toward and away from each other, activated by

muscles attached to their bases.

-Salivary glands secrete an anticoagulant called hirudin,

-All leeches are carnivores or bloodsucking ectoparasites.

DIVERSITY OF HIRUDINOMORPHA Segmental coeloms lack septa and mesenteries gave
transformed into circulatory channels
BRANCHIOBDELLIDA
Rhynchobdellida
-ectommensals/ ectoparasites oon crayfishes or crabs
-aquatic leeches with protrusible muscular pharynx
-Chatae, prostomium, and pygidium usually are absent
-with coelomic and hemal system
-body has 15 segments, the last forming a sucker
-feed by protruding pharynx
-head (peristomium plus 3 segments) is modifies into a sucker
with the mouth at its center -some such as Glossiphonia and Helobdella
(Glossosiphoniidae F), feed only on invertebrates
-zygotes deposited in cocoons attached to host
-Fish leeches (Piscicolidae F) parasitized both freshwater and
-embryos develop into cryptolarvae as in archychobdellid marine fish
leeches.
Order Arhynchobdellida
HIRUDINEA
Haemopsis terrestris
-sister taxon of Branchobdellida
Haemapdisa
-typcical leeches including Euhirudinea with A and P suckers
Phytobdella
-all have diagonal (crossed helical) body wall muscle and a
sperm storage that does not include seminal receptacles Suborder Gnathobdelliformes

Acantobdellida peledina -chiefly aquatic blood-suckers

-sole species in taxon -The aquatic Hirudo medicinalis (Hirudinidae F) is

the best known leech
-sister taxon of Euhirudinea
Aeromonas hydrophila
-ectoparasitic
Haemopsis
-lacks anterior sucker, instead attaches its oral end to the host
with hooked chatae borne on the first 5 segments. (seg 2-6 Macrobdella
prostomium and peri absent)
Philobdella
-Unique among leeches as having chatae, segmental coelomic
cavities and septa

- despite primitive arrangement of coelomic cavities and Suborder Pharyngobdelliformes

absence of anterior sucker, still moves in inchworm fashion
-Erpobdella
and does not use peristalsis.
Dina
-posterior sucker consists of terminal 4 segments

Euhirudinea

-Sister taxon of Acantobdellida peledina

-350 species of true leeches with A and P suckers

- Body composed of prostomium plus 33 segments

Posterior sucker of 7 segements (27-33)

Chatae absent