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Integrated Principles of Zoology

Eighteenth Edition

Chapter 14

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 Xenacoelomorpha, Platyzoa and Mesozoa

© McGraw Hill ©age fotostock/Alamy 2

 Getting Ahead

Radially symmetrical cnidarians and ctenophores can snare

prey from any direction but cannot chase prey efficiently.

Animals that actively seek food, shelter, and reproductive

mates require directed movements most effectively achieved
by elongated bodies that have a head and tail or bilateral
symmetry.

Cephalization is the concentration of sense organs in the

head region.

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 Body Cavities

Most metazoans have triploblastic bodies—have ectoderm,

endoderm, and mesoderm layers that produce all body
structures.
• Coelomate—have body cavity that develops entirely from
the mesoderm.
• Acoelomate—have no coelom but have digestive cavity.
• Region between epidermis and digestive cavity filled with
parenchyma.

• Pseudocoelomate—has internal body cavity surrounding

the gut but not completely lined with mesoderm.
• Psuedocoel may be filled with fluid or gelatinous matrix.

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 Body Plans

Figure 14.1 Acoelomate, pseudocoelomate, and coelomate body plans.

Access the text alternative for slide images.

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 Phylum Xenacoelomorpha

This is a new phylum.

Two sister clades are found in this phylum:
• Xenoturbellida.
• Acoelomorpha.

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 Xenoturbellids

Wormlike, ciliated animals.

Two furrows.
• Ring furrow: external, ciliated.
• Side furrow: longitudinal.
Xenoturbella is the lone genus.
• Six species.

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 Xenoturbella profunda

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 Features of Xenoturbella

Thick epidermis.
Subepidermal nerve net.
Frontal pore.
Ventral glandular network of unknown function.
Circular and longitudinal muscles.
Ventral mouth.
Blind gut.
Direct development.

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 Acoelomorpha

Small flat worms less than 5 mm in length.

Typically live in marine sediments; few are pelagic and some
live in brackish water.
Mostly free-living but some are symbiotic and a few are
parasitic; almost 350 species.
Members were formerly in Class Turbellaria within phylum
Platyhelminthes.
Have cellular ciliated epidermis with parenchyma layer that
has small amounts of ECM and circular, longitudinal, and
diagonal muscles.

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 Acoelomorph Worms

Figure 14.3 Acoelomorph worms, Waminoa sp., on a bubble coral,

Plerogyra sinuosa.

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 Digestion in Acoelomorphs

Some have digestive system with mouth leading to tube-like

pharynx followed by a sac-like gut, but no anus.
Many acoelomorphs have no gut and the pharynx is absent.
• Mouth leads into either an endodermally derived mass of
cells or syncytial mass.
• Phagocytotic cells digest food intracellularly when food is
passed into these temporary spaces.

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 Structure of an Acoelomorph

Figure 14.4 (A) Generalized acoelomorph flatworm. (B) Midsagittal

section showing gut cavity filled with endodermal cells.
Access the text alternative for slide images.

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 Reproduction in Acoelomorphs

Acoelomorphs are monoecious.

Female reproductive organs produces yolk-filled eggs called
endolecithal eggs.
Following fertilization some or all cleavage events produce a
duet-spiral pattern of new cells which may be one of the
defining morphological feature of acoelomorphs.

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 Features of Acoelomorphs

Have other defining features proposed for acoelomorphs.

• Biochemical patterns of neurotransmitters.
• Cellular ultrastructure such as formation of a network of
interconnecting rootlets from epidermal cilia.
Have distinct anteroposterior axis, but lack a “true” brain.
Have a radial arrangement of nerves in body, not the ladder-
like pattern seen in Platyhelminthes.
Statocysts have different structures than Platyhelminthes.

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 Phylogeny of Acoelomorphs

Phylogenetic studies using molecular characters such as

mitochondrial genome and myosin genes describe
acoelomorphs as early-diverging bilaterally symmetrical
triploblasts.
Have only four or five Hox genes unlike free-living
Platyhelminthes which have seven or eight Hox genes.

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 Clades Within Protostomia

Most triploblastic metazoans are divided into two superphlya:

Protostomia and Deuterostomia.
Protostomes now divided into two large clades: Ecdysozoa
and Lophotrochozoa.
• Ecdysozoa possess a cuticle that is molted as their bodies
grow.
• Lophotrochozoa share either a horse-shoe shaped feeding
structure, the lophophore or have unique larval form called
the trochophore.
Modern molecular phylogenies have grouped acoelomate
and coelomate taxa together within the protostomes.
• These body plans do not form monophyletic groups.
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 Trochophore Larvae

Minute, translucent, and roughly top-shaped.

Have a prominent circlet of cilia and sometimes one or two
accessory circlets.
Occur in the early development of other marine members of
Annelida and Mollusca and are assumed to be ancestral for
these groups.
Trochophore-like larvae also occur in some Platyhelminthes,
Nemertean, Echiura, and Sipunculida groups.

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 Clade Platyzoa

Clade Platyzoa is a unique group of lophotrochozoan

protostomes that contain Platyhelminthes, Gastrotricha, and
Gnathifera.
Gnathifera contains four phyla.
• Gnathostomulida.
• Micrognathozoa.
• Rotifera
• Acanthocephala.

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 Platyzoan Relationships

Figure 14.5 Hypothetical relationships among members of Platyzoa.

Access the text alternative for slide images.

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 Phylum Platyhelminthes

Commonly called flatworms; vary in size from a millimeter to

many meters in length like tapeworms.
Normally slender, leaf-like form but can also be long and
ribbon-like.
Some free-living; others parasitic.
Not a valid monophyletic phylum according to some due to
lack of single unique characteristic.
However, parasitic species have an external body covering
called a syncytial tegument (neodermis), not the cellular
ciliated epidermis of free-living species.

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 Example of a Platyhelminth

Figure 14.6 (A) Stained planarian. (B) Bipalium, a terrestrial flatworm.

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 Platyhelminth Diversity

Platyhelminthes is divided into four classes: Turbellaria,

Trematoda, Monogenea, and Cestoda.
Class Turbellaria.
• Mostly free-living with some symbiotic and parasitic forms.
• Bottom dwellers in marine areas or freshwater streams, pools,
and hot springs.
• Terrestrial flatworms limited to moist places.
Monogenea, Trematoda (flukes), and Cestoda (tapeworms) are all
parasitic.
• Monogeneans mostly ectoparasites.
• Cestodes and Trematodes are endoparasites.
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 Platyhelminthes Relationships

Figure 14.7 Hypothetical relationships among parasitic Platyhelminthes.

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 Turbellarian Form and Function

Most have cellular, ciliated epidermis on a basement

membrane.
Contains rod-shaped rhabdites that swell and form a
protective mucous sheath when discharged with water.
Most turbellarians have dual-gland adhesive organs in the
epidermis.
• Viscid gland cells fasten microvilli of anchor cells to
substrate.
• Secretions of releasing gland cells provide a quick
chemical detachment.

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 Anatomy of a Planarian

Figure 14.8 (A) Whole planarian. (B) Cross section of planarian through
pharyngeal region, showing relationships of body structures.
Access the text alternative for slide images.

© McGraw Hill ©Eric Grave/Science Source 26

 Dual-Gland Adhesive Organ

Figure 14.9 Reconstruction of dual-gland adhesive organ of the

turbellarian Haplopharynx sp.

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 Parasitic Platyhelminth Form and Function

Three parasitic classes have a non-ciliated body covering

called syncytial tegument that has many nuclei enclosed
within a cell membrane.
Many larval forms have temporary ciliated covering that is
shed when a host is contacted to prevent host immune
response.
Neodermis formed after several surface layers of epidermis
are shed, allowing cytoplasmic extensions from cells below
basement membrane to reach the surface.
The three parasitic groups are in clade Neodermata.

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 Tegument of Endoparasites

Resistant to host immune system and to digestive juices

within the host gut which allow tapeworms and other worms
to dwell in it.
Syncytial nature of tegument allows more resistance due to
lack of penetrable junctions between cells.
Tegument can be absorptive and secretory where secreted
enzymes can reduce host digestive system and absorb
nutrients from host gut cavity.
Most tapeworms have no mouth and lack complete digestive
tract.

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 Structure of the Tegument

Figure 14.10 Diagrammatic drawing of the structure of the tegument of a

trematode Fasciola hepatica.
Access the text alternative for slide images.

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 Turbellarian Nutrition and Digestion

Platyhelminthes generally have mouth, pharynx, and intestine.

In turbellarians the pharynx may extend through the ventral mouth.
• Intestine has three branches—one anterior and two posterior.
• Gastrovascular cavity lined with columnar epithelium.
• Generally carnivorous and detect food by chemoreceptors.
• Food trapped in mucous secretions from glands and rhabdites.
• Extend the proboscis to suck up bits of food.
• Enzymes are secreted for extracellular digestion; cells in
gastrodermis complete digestion at intracellular level.
• Undigested food egested out the pharynx.

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 Structure of a Planarian

Figure 14.11 Structure of a planarian. (A) Reproductive and

osmoregulatory systems. (B) Digestive tract and ladder-type nervous
system. (C) Pharynx extended through ventral mouth.

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 Parasitic Platyhelminth Digestion

Monogeneans and trematodes graze on host cells, feeding

on cellular debris and body fluids.
• Mouth opens near the anterior end.
• Pharynx is not extensible.
• Intestine ends blindly, varies in degree of branching but
often is Y-shaped.
Cestodes have no digestive system.
• Cestodes generally rely on the host’s digestive tract and
absorb digested nutrients as small molecules.

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 Structure of a Trematode

Figure 14.12 Structure of human liver fluke Clonorchis sinenesis.

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 Platyhelminth Excretion and Osmoregulation

Flatworms have protonephridia used for osmoregulation and

excretion.
Majority of metabolic wastes removed by diffusion across the
wall.
Have flame cells which are cup-shaped structures that have
flagella extending from the surface.
Beating flagella drive fluids down collecting ducts and
through delicate interlaced projections.
Wall of the duct bears folds or microvilli to resorb ions and
molecules.

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 Variations in Protonephridia

In Planarians, the collecting ducts join and empty at

nephridiopores to regulate water.
Monogeneans have two excretory pores that open laterally
near anterior end.
Trematodes have ducts that empty into excretory bladder
that leads to the outside via a terminal pore.
Cestodes have two main excretory canals on each side that
are continuous along the length of the worm and join on the
last segment and opens to the terminal pore.

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 Platyhelminth Nervous System

Most primitive type of flatworm nervous system, found in

some turbellarians, called subepidermal nerve plexus.
• Resembles the nerve net of cnidarians.
Other flatworms also have one to five pairs of longitudinal
nerve cords under the muscle layer.
Freshwater planarians have one ventral pair of nerve cords
forming a ladder-type pattern and the brain is a bilobed
ganglion anterior to the ventral nerve cords.
• Neurons are organized into sensory, motor, and other
types.

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 Platyhelminth Sense Organs

Active locomotion favored cephalization and evolution of

sense organs.
Ocelli (light-sensitive eyespots) present in turbellarians,
monogeneans, and larval trematodes.
Tactile and chemoreceptive cells are abundant over the body
especially in the ear-shaped auricles on the sides of the
head of planarians.
Some have statocysts for equilibrium and rheoreceptors to
sense the direction of water currents.
Sensory nerve endings found in oral suckers and genital
pores of parasitic groups.

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 Flatworm Reproduction and Regeneration

Many turbellarians reproduce asexually (fission) and sexually.

• Fission involves constricting behind the pharynx and
separating.
• Each half regenerates the missing parts for rapid population
growth.
• Some do not separate immediately, creating chains of zooids.
• Planarians are known for regenerative powers—if the head and
tail are cut off, each end grows the missing part and it retains
polarity.
Trematodes have asexual reproduction in their intermediate hosts,
snails.
Some juvenile cestodes have extensive asexual reproduction.
Nearly all flatworms are monoecious (hermaphroditic) but can
cross-fertilize.
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 Fission in Turbellarians

Figure 14.13 Some small freshwater turbellarians.

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 Flatworm Reproductive Development

Endolecithal eggs with spiral determinate cleavage are typical of

some turbellarians, and likely ancestral for flatworms.
Parasitic flatworms generally have female gametes with little yolk;
yolk is released by separate organs called vitellaria.
• Ectolecithal development occurs when yolk cells surround the
zygote.
• Spiral cleavage pattern cannot be distinguished due to yolk.
• Zygote and yolk cells surrounded by eggshell move into the
uterus and are released through the genital pore.
• Access to the yolk in ectolecithal eggs can be problematic for
the embryo so the outermost epidermal cells grow outwards to
encompass the yolk.
• As epidermal cells are shed, successive inner layers enclose
the yolk.
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 Turbellarian Reproductive Structures

Male structures include one or more testes, connected to

vasa efferentia that connect to one vas deferens.
• Vas deferens runs to a seminal vesicle and leads to a
papilla-like penis or extensible copulatory organ called a
cirrus.
Turbellarians develop male and female organs opening at a
common genital pore.
• After copulation, eggs and yolk cells enclosed in small
cocoon that is attach by a stalk to plants.
• Embryos emerge and resemble little adults.
• Some embryos of marine forms develop to ciliated, free-
swimming larvae similar to trochophores.
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 Parasitic Flatworm Reproductive Structures

Monogeneans hatch free-swimming larvae that attach to

hosts and develop into juveniles.
Larval trematodes emerge as ciliated larvae that penetrate or
are eaten by the intermediate host like snails.
Cestodes hatch only after being consumed by a variety of
intermediate host.

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 Class Turbellaria

Mostly free-living; range from 5 mm to 50 cm long.

Live under objects in marine, freshwater, and terrestrial
habitats.
Turbellarians are distinguished by the presence or absence
of gut, pattern of branching of the gut, and type of pharynx.
• Polyclads have a folded pharynx and a gut with many
branches, with larger polyclads having more highly
branched intestines.
• Members of order Tricladida, like the planaria, are
ectolecithal and have a three-branched intestine.

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 Turbellarian Gut Pattern

Figure 14.14 Intestinal pattern of two orders of turbellarians. (A)

Tricladida. (B) Polycladida.

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 Turbellarian Movement

Turbellarians combine creeping with ciliary movements.

• Very small planaria swim by cilia.
Others move by gliding over a slime track secreted by
marginal adhesive glands.
• Use rhythmical muscular waves that pass backwards from
the head.
Larger polyclads and terrestrial turbellarians crawl with
muscular undulations like snails.

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 Marine Turbellarian

Figure 14.15 Pseudobiceros hancockanus, a marine polyclad

turbellarian.

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 Class Trematoda

All are parasitic flukes and most adults are endoparasites of

vertebrates.
Mostly leaflike with one or more suckers, but lack opisthaptor
of monogenean flukes.
Adaptations for parasitism include:
• Various penetration glands.
• Glands to produce cyst material.
• Hooks and suckers for adhesion.
• Increased reproductive capacity.
• Sense organs poorly developed.
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 Trematode Diversity

Trematodes share many characteristics with ectolecithal

turbellarians.
• Well-developed alimentary canal, reproductive, excretory,
nervous systems, and well-formed musculature and
parenchyma cells.
Subclass Aspidogastrea is least well-known.
• Most have single hosts, usually mollusks.
• Those with secondary hosts usually infect fish or turtles.
Subclass Digenea is largest and most well-known.
• Many species with medical and economic importance.

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 Subclass Digenea

Complex life cycle.

• First intermediate host is a mollusk.
• Definitive or final host is a vertebrate, inside which the
parasites reproduce sexually.
Some species need a 2nd or 3rd intermediate host in the life
cycle.
Parasitize almost all kinds of vertebrate hosts and can inhabit
a wide variety of body parts within the hosts.

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 General Digenean Life Cycle

Larva passes from definitive host in excreta and must reach

water to develop.
Hatches into a free-swimming ciliated larva, the miracidium.
Miracidium penetrates tissues of a snail and is transformed
into a sporocyst.
Sporocyst reproduces asexually to form more sporocysts or
rediae.
Rediae reproduce asexually to form more rediae or
cercariae.
Single egg therefore can produce a multitude of infectious
progeny.
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 Life Cycle of a Digenean

Cercariae emerge from the snail and penetrate a final host, a

2nd intermediate host, or encyst on aquatic vegetation.
Cercaria then develop into metacercariae (juvenile flukes).
Metacercariae are eaten by definitive host and move to final
infection sites and grow into adults.
Numerous infectious digenean parasites impact humans and
domesticated animals.

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 Sheep Liver Fluke

The sheep liver fluke (Fasciola hepatica) was first digenean

life cycle described.
Infects sheep and other ruminants.
Adult flukes live in liver bile passage; eggs released in feces.
Miracidia hatch and penetrate snails to become sporocysts.
After two generations of rediae, the cercaria encyst on
vegetation and await being eaten.
When eaten, metacercariae develop into young flukes and
live in the liver of hosts.

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 Human Liver Fluke

Clonorchis sinensis is most important human liver fluke; also

infects cats, dogs, and pigs.
Common in China, Japan, and Southeast Asia.
Adult fluke is 10 to 20 mm long with an oral and ventral
sucker.
Digestive system includes pharynx, muscular esophagus,
and two long unbranched intestinal ceca.
Excretory system has two protonephridial tubules with
branches lined with flame cells that form a bladder and open
to the outside.

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 Liver Fluke Structure

Nervous system has two cerebral ganglia and longitudinal

cords with transverse connectives like other flatworms.
Reproductive system is hermaphroditic with 80% of body
devoted to it.
• Two branched testes lead to single vas deferens, a
seminal vesicle, an ejaculatory duct, and then to genital
pore.
• Branched ovary leads to short oviduct that is joined by
ducts from seminal receptacle and vitellaria at the ootype.
• From the ootype, surrounded by Mehlis’ gland, the uterus
connects to the genital pore.

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 Clonorchis Life Cycle

Adults live in bile passageways of humans and other fish-

eating mammals.
Eggs containing a complete miracidium are shed into water
with feces.
Eggs hatch only when ingested by snails of specific genera.
Miracidium enters snail tissue and transforms into a
sporocyst.
Sporocyst produces one generation of rediae, which begin
development.
Rediae pass into the snail liver and develop into tadpole-like
cercariae.
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 Life Cycle of Clonorchis

Cercariae escape into water and make contact with fish of

the family Cyprinidae.
Bore under scales and into fish muscles where they shed tail
and encyst as metacercariae.
A mammal eats raw fish and cyst dissolves, releasing young
flukes to migrate up bile duct.
Heavy infection can destroy the liver and result in death.
Control of parasites requires the removal of snails or
thorough cooking of fish.

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 Life Cycle of Clonorchis sinensis

Figure 14.16 Life cycle of Clonorchis sinensis.

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 Schistosoma – Blood Flukes

Over 200 million people infected with schistosomiasis.

Common in Africa, South America, West Indies, and the
Middle and Far East.
Sexes are separate (dioecious) with males being broader,
heavier and have large ventral groove called gynecophoric
canal that is posterior to the ventral sucker.
Gynecophoric canal wraps around long and slender female
during mating session.

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 Schistosomiasis

Three species account for most human schistosomiasis:

• S. mansoni—lives in veins that drain large intestine;
common in Africa, Brazil, West Indies, and many northern
parts of South America; depends on Biomphalaria snails.
• S. japonicum—lives in veins of small intestine; confined to
the Far East and uses Oncomelania snail hosts.
• S. haemotobium—lives in veins of urinary bladder;
prevalent in Africa and uses Bulinus and Physopsis snails
as intermediary hosts.
Control is best achieved through proper hygiene and
avoidance of contaminated areas.

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 Schistosoma Life Cycle

Eggs discharged in human feces or urine.

In water, eggs hatch as ciliated miracidia and search for
snail.
In snail, transform to sporocysts.
Sporocysts produce daughter sporocysts that produce
cercaria.
Cercariae escape snail and swim until they contact human
skin where cercariae pierce the skin and shed their tails.
Enter blood vessels and migrate to the hepatic portal blood
vessels so as to develop in the liver.
No redia or metacercariae stages.

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 Life Cycle of Schistosoma

After development in liver, migrate to target sites.

As females release eggs, they are extruded through venous walls
and gut or bladder lining to exit with feces or urine.
Eggs that do not get extruded flow back to the liver in blood and
can form centers of inflammation.
Eggs cause most of the ill effects in the human host.
• S. mansonii and S. japonicum causes ulcers, abscesses and
bloody diarrhea and abdominal pain; usually more severe cases
of the disease.
• S. haematobium causes bladder ulceration leading to bloody
urine and pain; generally less severe cases of the disease.
• All eggs can impede blood flow and interfere with normal organ
function.
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 Life Cycle of Schistosoma mansoni

Figure 14.17 (A)

Adult male and
female
Schistosoma
mansoni in
copulation.
(B) Life cycle of
Schistosoma
mansoni.

© McGraw Hill ©Larry S. Roberts 63

 Liver Damaged by Schistosoma

Figure 14.18 This cut surface of a liver shows schistosomal hepatic

fibrosis.

© McGraw Hill ©Larry Roberts/McGraw -Hill Education 64

 Schistosome Dermatitis

Schistosome dermatitis is known as swimmer’s itch.

Various species can cause rashes and dermatitis when the
cercariae penetrate an unsuitable host, like humans.
Severity of the rash increases with increasing number of
contacts or sensitization as the cercariae are attacked by the
hosts’ immune system and release allergenic substances.
It affects many tourists at contaminated vacation sites and
infested lakes.

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 Paragonimus—Lung Flukes

Paragonimus westermani is a lung fluke that parasitizes a

variety of mammals.
Eggs are coughed, then swallowed, and eliminated in feces.
Zygotes develop in water and miracidia penetrate a snail
host.
In snail, miracidia produce sporocysts, which become rediae.
Cercariae form in rediae and are shed into the water or
ingested by freshwater crabs that prey on infested snails.
Metacercariae develop in crabs and human infection occurs
by eating uncooked crabmeat.
Infection causes respiratory issues like breathing difficulties
and chronic cough that can lead to fatalities.
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 Other Trematodes

Fasciolopsis buski lives in human and pig intestines.

• Mostly in India and China.
• Larval stages occur in several snail species.
• Cercariae encyst on water chestnuts, an aquatic plant
eaten raw by humans and pigs.
Leucochloridium lives in snails and birds.
• Snails eat vegetation infected with eggs.
• Sporocysts enlarge and enter snail’s head and tentacles,
becoming colorful and pulsating.
• It attracts birds to eat snails and continue the life cycle.

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 Paragonimus

Figure 14.19 Lung fluke Paragonimus westermani.

© McGraw Hill ©Natural History Museum, London/SPL/Science Source 68

 Class Monogenea

Originally placed in Trematoda but now in different classes.

Some now argue they are sister taxa, both having a posterior
attachment with hooks.
Monogeneas are all external parasites of many fish,
especially on gills, but a few are found in bladders of frogs
and turtles.
Generally cause little harm or damage to host but can
become a problematic pathogen in crowded fish farming
areas.

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 Monogenean Features

Have direct life cycle in a single host.

Egg hatches to produce ciliated larvae called
oncomiracidium that attaches to host by posterior hooks.
Posterior hooks may become the posterior attachment organ
of the adult, called the opisthaptor.
Opisthaptors vary widely as hooks, suckers, clamps, and a
combination of forms to withstand the force of water flow
while attached to the gills and skin of fish.

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 Structure of Monogenean

Figure 14.20 A monogenetic

fluke Gyrodactylus
cylindriformis, ventral view.

Access the text alternative for slide images.

© McGraw Hill 71
 Class Cestoda

Tapeworms have long flat bodies.

• Scolex, bearing suckers and hooks, allows attachment to
the host.
• Scolex is followed by a linear series of reproductive units,
proglottids.
Tapeworms lack a digestive system but have well-developed
muscles.
Excretory and nervous systems similar to other flatworms.
Lack sensory organs except for modified cilia that are
sensory endings on the tegument.

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 External Anatomy of a Tapeworm

Figure 14.21 A tapeworm,

showing strobila and
scolex. The scolex is the
organ of attachment.

© McGraw Hill ©Science Photo Library RF/Getty Images 73

 Tapeworm Sensory Ending

Figure 14.22 Schematic drawing

of a longitudinal section through a
sensory ending in the tegument of
Echinococcus granulosus.

Access the text alternative for slide images.

© McGraw Hill 74
 Form of Cestoda

Cestodes, like trematodes and monogeneans, have no

external motile cilia.
However, entire surface of cestodes is covered with small
projections called microtriches similar to microvilli seen in the
vertebrate small intestine.
Microtriches increase the surface area for food absorption
since tapeworms are parasitic and attach to the intestines of
the hosts.
Subclass Eucestoda has the most species.

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 Subclass Eucestoda

Main body of tapeworm is a chain of proglottids called the

strobila.
Proglottids originate in the germinative zone just behind the
scolex.
Some practice self-fertilization, although the norm is cross-
fertilization.
Each proglottid contains a complete male and female
reproductive system.
Shelled embryos form in the uterus and either expelled
through uterine pore or the entire proglottid is shed from the
worm.

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 Tapeworm Diversity

Proglottid formation is not “true” segmentation since

replication of sex organs is not equivalent to metamerism in
annelids and others.

Nearly all cestodes require two hosts and the adult is

parasitic in the digestive tract of the vertebrates.

Over 1000 species of tapeworms known, infecting almost all

vertebrates and having intermediate invertebrate host.

Most tapeworms do little harm to host.

© McGraw Hill 77
 Taenia saginata—Beef Tapeworm

Lives as an adult in the alimentary canal of humans while

juveniles mostly form in the intermuscular tissue of cattle.
Mature adults can reach over 10 meters in length with over
2000 proglottids.
Scolex has four suckers but no hooks and a short neck
connecting to strobili.
Proglottid has muscles and parenchyma with repeating
reproductive and excretory systems and complete male and
female organs like those in trematodes.
Human infection very common; can be avoided by eating
only thoroughly cooked beef since much of the beef supply
(20%) is not inspected by the USDA.
© McGraw Hill 78
 Beef Tapeworm Features

Excretory canals run from scolex along entire body and

connect to excretory duct.
Nerve cords from a nerve ring in the scolex run along
proglottids.
Contains vitellaria in a single compact vitelline gland
posterior to ovaries.
Gravid proglottids break off and usually crawl out of feces
and attach to vegetation.
Proglottids rupture as they dry; embryos are viable for five
months and are picked up by grazing animals.

© McGraw Hill 79
 Life Cycle of Beef Tapeworm

Cattle swallow shelled larvae; these hatch as oncospheres

and use hooks to burrow through intestinal wall to blood or
lymph.
Reach voluntary muscle and encyst to become bladder worms
(juveniles called cysticerci).
• Develop invaginated scolex but remain dormant.
When the infected meat is eaten, the cyst wall dissolves and
the scolex evaginates to attach to intestinal mucosa.
New proglottids develop in 2 to 3 weeks to form mature worm.
Infected individuals expel numerous proglottids daily either in
feces or by crawling out of the anus.
© McGraw Hill 80
 Beef Tapeworm Life Cycle

Figure 14.23 Life

cycle of beef
tapeworm, Taenia
saginata.

© McGraw Hill 81
 Taenia solium—Pork Tapeworm

Adults live in small intestines of humans while juveniles live

in muscles of pigs.
Most common mode of infection occurs when pigs consume
infected human fecal material containing fertilized eggs.
• Inside pig, tapeworm larvae encyst in muscle tissue.
• Humans eating undercooked pork ingest cysts which
hatch and develop into adult tapeworms.
If human ingest fertilized tapeworm eggs, can develop
cysticercosis.
• Larvae migrate to organs and form cysticerci.
• Infection of brain of spinal cord may lead to death.
© McGraw Hill 82
 Brain Damage from Cysticercosis

Figure 14.25 Section through the brain of a person who died of cerebral
cysticercosis, an infection with cysticerci of Taenia solium.

© McGraw Hill ©Ana Flisser 83

 Diphyllobothrium latum—Fish Tapeworm

Adults found in intestines of humans, dogs, cats, and other

mammals.
Immature stages found in crustaceans and fish.
Largest cestode to infect humans, reaching up to 20 meters
in length.
Fish tapeworm infections can occur anywhere people eat raw
fish and is quite common in the Great Lakes region of the
USA.

© McGraw Hill 84
 Echinococcus granulosus—Unilocular Hydatid

A dog tapeworm that causes hydatidosis.

Adults parasitize dogs and other canines; juveniles infest
many mammal species.
Humans may serve as dead-end host.
Juveniles form a special cysticercus, a hydatid cyst, that
grows up to 20 years and form large masses that can affect
major body parts.
• Main cyst maintains a single chamber (unilocular).
• Daughter cysts bud off with thousands of scolices, each
able to produce a worm if eaten by canine.
Treatment is chemotherapies or surgical removal.
© McGraw Hill 85
 Dog Tapeworm Proglottids

Figure 14.24 Mature proglottid of Taenia pisiformis, a dog tapeworm.

© McGraw Hill ©NHPA/M. I. Walker 86

 Dog Tapeworm

Figure 14.26 Echinococcus granulosus, a dog tapeworm. (A) Early hydatid cyst
or bladder-worm stage found in cattle, sheep, hogs, and sometimes humans that
produces hydatid disease. (B) The adult tapeworm lives in the intestine of a dog
or other carnivore.

© McGraw Hill 87
 Phylum Gastrotricha

Gastrotrichs are small ventrally flattened animals.

Look similar to rotifers but lack corona and mastax and have
bristly scaly body.
Usually found gliding along substrates via their ventral cilia.
Found in fresh, brackish, and salt water with many species
being cosmopolitan.
About 450 species, only a few of which can be in both fresh
and marine habitats.

© McGraw Hill 88
 Gastrotrich Form

Have convex dorsal surface bearing bristles, spines or

scales, and a ventral flattened ciliated surface.
Head region is lobed and ciliated while tail region may be
elongated and forked.
Has partial syncytial epidermis beneath cuticle with a dual-
gland system for attachment and release.
No specialized respiratory or circulatory system; uses simple
diffusion.
No body cavity.

© McGraw Hill 89
 Gastrotrich Function

Gastrotrichs have extracellular digestion with complete

digestive system including mouth, muscular pharynx,
stomach-intestine region, and anus.
Protonephridia with solenocytes rather than flame cells.
• Solenocytes enclose a single flagella within a cylinder of
cytoplasmic rods.
Nervous system has brain near pharynx with a pair of lateral
nerve trunks.
Generally lack eyespots; some have pigmented ocelli in
brain.
Some sensory bristles on the head used for tactile response.
Pestle organ on head may be chemoreceptor.
© McGraw Hill 90
 Gastrotrich Reproduction

Gastrotrichs are typically hermaphroditic.

• Male system of some is so rudimentary that they are
functionally parthenogenetic females.
Produce thin-walled eggs but also develop thick-shelled
dormant eggs that can withstand harsh environments for
many years.
Development is direct with growth and maturation being
rapid and juveniles reach sexual maturity within days.

© McGraw Hill 91
 External and Internal Gastrotrich Anatomy

Figure 14.27 (A) Live Chaetonotus simrothic, a common gastrotrich. (B)

Dorsal surface. (C) Internal structure, ventral view.

© McGraw Hill ©Perennou Nuridsany/Science Source 92

 Structure of a Gastrotrich

Figure 14.28 Gastrotrichs in order Macrodasyida. (A) Macrodasys. (B)

Turbanella.

© McGraw Hill 93
 Clade Gnathifera

Consists of 4 lophotrochozoan phyla.

• Gnathostomulida, Micrognathozoa, Rotifera, and
Acanthocephala.
Ancestors possessed complex cuticular jaws with
homologous microstructure.
Living gnathiferans vary in the number of pairs of jaws.
• Acanthocephalans have no jaws.
Most gnathiferans are free-living aquatic animals.
• Acanthocephalans are endoparasites.
Rotifers and acanthocephalans are presumably sister taxa.
• Both have a eutelic syncytial epidermis.
© McGraw Hill 94
 Phylum Gnathostomulida

Jaw worms are found in a variety of areas around the world.

• Mostly in interstitial spaces of very fine sand, sediment,
and silt from the coasts to the deep sea.
• Over 80 species described.
Can endure very low oxygen.
Live in association with a variety of other small forms like
ciliates, tardigrades, and worms.
Glide, swim in loops and spirals and bend the head from side
to side with many sensory cilia on the head.

© McGraw Hill 95
 Gnathostomulid Features

Feed by scraping bacteria and fungi from the substrate with

paired jaws on the pharynx.
Ciliated epidermis, but only 1 cilium per epidermal cell
(unusual in lophotrochozoans).
Body is acoelomate with no circulatory system; probably use
diffusion for excretion and gas exchange.
Not much known about mating behavior and reproductive
system.
• Protandric or simultaneous hermaphrodite that can cross-
fertilize internally forming single zygote.

© McGraw Hill 96
 Structure of a Gnathostomulid

Figure 14.29 (A) Gnathostomula jenneri is a tiny member of the

interstitial fauna between grains of sand or mud.

Access the text alternative for slide images.

© McGraw Hill 97
 Phylum Micrognathozoa

One known species, Limnognathia maerski.

Tiny animals living interstitially, using cilia to move.
Two-part head, thorax and abdomen leading to short tail.
Epidermis has dorsal plates but no ventral ones.
Have a ventral ciliary adhesive pad that produces glue.
Have three pairs of complex jaws with mouth leading to
simple gut and anus.
Two pairs of protonephridia.
Reproductive system is not well understood; only female
organs have been found.
© McGraw Hill 98
 Structure of a Micrognathozoan

Figure 14.30 (A) Limnognathia maerski, a micrognathozoan. (B) Detail of

complex jaws. (C) A living specimen.

© McGraw Hill ©Martin V. Sorensen 99

 Phylum Rotifera

Have ciliated crown (corona) that beats like rotating wheels.

About 2000 species, most between 100 and 500 µm.
Inhabit freshwater lakes and ponds; usually benthic, living on
vegetation and between sand grains (meiofauna).
Pelagic forms common in surface waters; some are epizoic
(live on body of another animal) and some parasitic.
Can have bizarre shapes ranging from globular and saclike
to elongated and vase-like with thick outer epidermis (lorica).

© McGraw Hill 100

 Features of Rotifera

Rotifers can endure long periods of dryness (desiccation).

• During dry periods appear like sand grains.
• Can be revived upon addition of water.
Other species can survive extreme cold temperature (−272
Celsius) and be successfully revived.
Can have a variety of locomotory forms ranging from free-
floating, creeping and swimming, to sessile forms.
Some are colonial while others are solitary.

© McGraw Hill 101

 Structure of a Rotifer

Figure 14.31 (A) Live Philodina sp., a common rotifer. (B) Structure of
Philodina sp.
© McGraw Hill ©John Walsh/Science Source 102
 External Features of Rotifers

Rotiferan body has a head, trunk, and tail (foot); except for
the corona, it is nonciliated and covered in cuticle.
Corona can have sensory bristles (papillae), a midventral
mouth and coronal cilia use for swimming and feeding.
Trunk may be elongated or saclike with sensory antennae.
• Body wall may be superficially ringed, appearing
segmented.
• Have a fibrous layer in the epidermis.
• Some have thick fibrous layer called lorica arranged as
plates or rings.
Foot may have 1 to 4 toes with pedal glands for attachment.
• Reduced in swimming pelagic forms.
© McGraw Hill 103
 Examples of Rotifers

Figure 14.32 Variety of form in rotifers.

© McGraw Hill 104

 Internal Features of Rotifers

Syncytial epidermis secretes cuticle and bands of

subepidermal muscles around the body.
Large pseudocoel filled with fluid, muscles, and mesenchymal
ameboid cells.
Digestive system is complete with pharynx (mastax) fitted with
hard jaws (trophi) for sucking and grinding of food,
• Mastax is often a distinguishing feature.
Salivary and gastric glands likely secrete enzymes for
extracellular digestion; absorption occurs in the stomach.

© McGraw Hill 105

 Rotifer Function

Excretory system has pair of protonephridial tubules with

flame cells that empty to common bladder.
Pulsating motion drains bladder into cloaca; intestines and
oviduct also empty into cloaca.
Protonephridia may be important in osmoregulation.
Has bilobed brain dorsal to the mastax region and has paired
eyespots, sensory bristles, papillae, and antennae.

© McGraw Hill 106

 Rotifer Reproduction

Dioecious with males smaller than females.

• No males known in class Bdelloidea; only present for a few
weeks in class Monogononta.
Females in Bdelloidea and Monogononta have combined
ovaries and yolk glands, called germovitellaria.
• Yolk enters developing ova through cytoplasmic bridges; it
is not stored as separate yolk cells.
In Bdelloidea, all females are parthenogenetic.
• Produce diploid eggs that hatch to diploid females.
In Seisonidea, females produce haploid eggs.
• Must be fertilized and develop into males or females.
© McGraw Hill 107
 Reproduction in Class Monogononta

Most of the year, diploid females produce diploid amictic

eggs.
• Amictic eggs develop parthenogenetically into diploid
females.
Environmental factors like crowding, diet, and photoperiod
may induce amictic eggs to develop into diploid mictic
females.
• These females produce haploid eggs.
• Unfertilized haploid eggs form haploid males.
• Fertilized haploid eggs (mictic eggs) become dormant with
a thick resistant shell to survive over winter and later hatch
as amictic females.
• Dormant mictic eggs can be dispersed by winds and birds.
© McGraw Hill 108
 Monogonont Reproductive Structures

Males have single testis and ciliated sperm duct leading to

genital pore.
• Males usually lack cloaca.
End of sperm duct forms copulatory organ which can
penetrate any part of female body and inject sperm into
pseudocoelom where fertilization occurs.
Females hatch with complete adult features and mature
quickly.
Males often do not grow and are sexually mature at hatching.

© McGraw Hill 109

 Monogonont Life Cycle

Figure 14.33 Reproduction of some rotifers is parthenogenetic during the

part of the year when environmental conditions are suitable.
© McGraw Hill 110
 Phylogeny of Rotifera

Traditionally rotifers are split into three classes:

• Class Seisonidea—marine forms with elongated body but
vestigial corona and epizoic in gills of crustaceans.
• Class Bdelloidea—creeping or swimming form with trochal
discs in corona; parthenogenetic with unknown males.
• Class Monogononta—sessile and swimming forms with
small males and complex egg types.
Recent molecular work has challenged these classifications;
they are subject to debate and possible revision.

© McGraw Hill 111

 Phylum Acanthocephala

Spiny-headed worms have distinctive cylindrical invaginable

proboscis with rows of recurved spines.
• Used to attach to intestines of hosts.
Adults are endoparasites of birds, fish, and mammals while
larvae live in crustaceans and insects.
Females are usually larger than males.
Bilaterally flattened body with many transverse wrinkles.
Variable sizes and cosmopolitan in distribution, with over 1100
species.

© McGraw Hill 112

 Acanthocephalan Form

Syncytial body wall with many minute crypts (depressions) to

increase body surface area.
Tegument has lacunar system of fluid-filled canals to enhance
diffusion across body wall.
No heart but muscular body wall forms tubes connected to
lacunar system and collectively function like heart that uses
lacunar fluid as a circulatory system.
Proboscis can be inverted into a proboscis receptacle.
Two elongated hydraulic sacs, lemnisci, are attached to neck;
function is unknown.

© McGraw Hill 113

 Acanthocephalan Function

No respiratory system.
When present, excretory system has simple protonephridia
with flame cells.
Nervous system and sense organs are reduced.
• Has central ganglia within the proboscis receptacle
connecting to sensory nerves on the proboscis and body.
• Sensory endings on proboscis and genital bursa.
No digestive tract; absorb nutrients through tegument.
• Require host dietary carbohydrates.
• Form a metabolic “sink” for glucose.
© McGraw Hill 114
 Acanthocephalan Reproduction

Acanthocephalans are dioecious.

• Males have paired testes and vas deferens, and a common
ejaculatory duct and copulatory organ.
• Females have ovarian tissue within ligament sacs that breaks
into ovarian balls; these are released to float in the pseudocoel.
• One ligament sac leads to uterine bell that receives developing
embryos and passes them to uterus.
Unique embryo selective apparatus system in uterine bell.
• Fully developed embryos are longer, and are passed on to
uterus.
• Shorter immature embryos are retained for further maturation.
Shelled embryos are released in the feces of host and await entry
to intermediate host.
© McGraw Hill 115
 Hosts of Acanthocephalans

Acanthocephalans are not normally parasitic to humans,

though they do infect pigs and other mammals.
• Can infect humans if we eat infected foods.
One common species uses soil-inhabiting beetle larvae as
intermediate host.
• Acanthocephalan larva (acanthor) burrows into beetle larva
intestines and develops into juvenile (cystacanth) in
hemocoel.
• When the beetle larvae are eaten by a pig, the acanthor
penetrates the intestinal wall with the spiny proboscis to
attach.
• Host response is varied from little inflammation to great
pain.
© McGraw Hill 116
 Structure of Acanthocephalans

Figure 14.34 Details of the Acanthocephalan worm, Polymorphus

botulus.

© McGraw Hill (a-c): ©Wayne Lord and Inga Sidor 117

 Phylogeny of Acanthocephala

Largely organized by shape and structure of spines on the

proboscis.
Traditionally divided into three classes.
• Archiacanthocephala, Eoacanthocephala, and
Palaeacanthocephala.
New molecular data suggests that acanthocephalans may be
a class of highly derived rotifers, possibly sister taxon to
Bdelloidea.

© McGraw Hill 118

 Phylum Mesozoa

Mesozoa were considered a “missing link” between

unicellular eukaryotes and metazoan.

Usually minute, ciliated, and wormlike animals that live as

parasites or symbionts in marine invertebrates.

Arranged in two layers of 20 to 30 cells; layers are not

homologous to germ layers of metazoans.

Two classes, Rhombozoa and Orthonectida, are so different

that some authorities place them in separate phyla.

© McGraw Hill 119

 Rhombozoans

Rhombozoans live in kidneys of benthic cephalopods.

Adults are called vermiforms (or nematogens) and are long
and slender.
Inner, reproductive cells give rise to vermiform larvae.
When overpopulated, reproductive cells develop into gonad-
like structures producing male and female gametes.
Zygotes grow into ciliated infusoriform larvae which are shed
with host urine into the seawater.
• Much of the life cycle is still being determined.

© McGraw Hill 120

 Structure of Rhombozoans

Figure 14.35 Two methods of reproduction by mesozoans. (A) Asexual

development of vermiform larvae. (B) Under crowded conditions in the host
kidney, gametes that produce infusoriform dispersal larvae in the host urine.

© McGraw Hill 121

 Orthonectids

Orthonectids parasitize a variety of invertebrates like brittle

stars, molluscs, and worms.
Reproduce sexually and asexually.
Asexual stage is quite different from rhombozoans.
• Consists of a multinucleated mass called a plasmodium
that divides to form males and females.

© McGraw Hill 122

 Structure of Orthonectids

Figure 14.35 (A) Female and, (B) male orthonectid (Rhopalura).

© McGraw Hill 123

 Phylogeny

Evolutionary relationships are still in flux for these groups.

• Many phylogenies have various disagreements due to
new molecular sequences and cladistic analysis.
Most current phylogenies place members of Acoelomorpha
as sister taxon of all Bilateria.
• Acoelomorpha differ from Platyhelminthes in embryonic
patterns, mesoderm formation, and nervous structures.
• Within Bilateria, molecular evidence suggests protostomes
split from deuterostomes in the Precambrian.
• Protostomes later split into lophotrochozoans and
ecdysozoans; relationships within Lophotrochozoa are still
in flux.
© McGraw Hill 124
 Phylogenetic Uncertainty

We depict a lophotrochozoan clade called Platyzoa

(Platyhelminthes, Gastrotricha, and Gnathifera), but not all
phylogenies support this grouping.
Within Platyhelminthes, class Turbellaria is clearly
paraphyletic.
Within Gnathifera, clade Syndermata (Acanthocephala and
Rotifera) emerges repeatedly from phylogenetic studies
repeatedly.
• Several studies show that acanthocephalans belong within
Rotifera.
Mesozoans are identified as lophotrochozoan protostomes
based on molecular data, but are not placed in Platyzoa.
© McGraw Hill 125
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