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Integrated Principles of Zoology

Eighteenth Edition

Chapter 13

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 Radiate Animals

© McGraw Hill ©William C. Ober/Medical Scientific Illustration 2

 A Fearsome Tiny Weapon

Cnidarians are generally simple animals.

• Relatively more organized than sponges.
• Most are sessile but several are motile.
However, many cnidarians are very effective predators.
• Able to kill and eat prey more highly organized, swift, and
intelligent than themselves.
• They possess tentacles that bristle with tiny, remarkably
sophisticated weapons called nematocysts.

© McGraw Hill 3
 A Nematocyst

A nematocyst is a fearsome, wondrous, tiny weapon.

• Requires only small stimuli to make it fire.
• Tiny thread bursts from a nematocyst that can achieve a
velocity of 2 meters/sec and an acceleration of 40,000×
gravity.
• Instantly penetrates its prey and injects a paralyzing toxin.
• A small animal that brushes against one of the tentacles is
suddenly speared with hundreds or even thousands of
nematocysts and quickly immobilized.

© McGraw Hill 4
 Phylum Cnidaria

Over 9,000 species in phylum Cnidaria.

• Branching, plantlike hydroids.
• Flowerlike sea anemones and jellyfishes.
• Architects of the ocean floor, such as sea whips, sea fans,
and stony corals whose thousands of years of calcareous
house-building activities produced great reefs and coral
islands.
Equipped with unique cells: Cnidocytes that contain a
specialized stinging organelle, the nematocyst.

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 Distribution of Cnidarians

Cnidarians have the longest fossil history with specimens

from over 700 million years ago.
Widespread in marine habitats; most found in shallow water
areas.
Some freshwater; none are terrestrial.
Generally found in warm tropical regions.
Mostly sessile, some free-floating, and a few slow-swimming
species.
Very efficient predators that capture and consume larger,
more complex, swifter swimming creatures.
Can live symbiotically and commensally with other animals.

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 Mutualistic Relationship

Examples of mutualistic relationships between Cnidarians

and other creatures.
• Hydroids and sea anemones can live on snail shells
inhabited by hermit crabs to provide protection from
predators.
• Some ctenophores, molluscs, and flatworms eat hydroids
and use the stinging nematocysts for their own defense.
• Algal cells frequently live as mutuals in the tissues of
hydras and reef-building corals that create extensive
habitats for many other creatures in tropical areas.

© McGraw Hill 7
 Mutualism of Hermit Crab and Cnidarian

Figure 13.1 (A) A hermit crab with its cnidarian mutuals. (B) Portion of a
colony of Hydractinia sp.

Access the text alternative for slide images.

© McGraw Hill ©William C. Ober/Medical Scientific Illustration 8

 Cnidarian Classes

Four traditional classes of Cnidaria.

• Hydrozoa: Hydroids, fire corals, and Portuguese man-of-
war.
• Scyphozoa: True jellyfishes.
• Cubozoa: Cube or box jellyfishes.
• Anthozoa: Largest class; includes stony corals, soft corals,
and sea anemones.
A fifth class, Staurozoa, has been proposed.
• No medusae in life cycle but has polyp topped by medusa-
like region.

© McGraw Hill 9
 Cnidarian Cladogram

Figure 13.2 Cladogram showing hypothetical relationships of cnidarian classes.

Access the text alternative for slide images.

© McGraw Hill 10
 Form and Function: Dimorphic Body Plan

Cnidaria generally have dimorphic body plans.

• Polyp (hydroid) form.
• Adapted to sedentary or sessile lifestyle.

• Medusa (jellyfish) form.

• Adapted to free-living and floating existence.

• Common name jellyfish is now being replaced by more accurate

term of sea jelly.

• Polyps and medusa forms both retain sac-like body plans.

• Medusa is essentially unattached polyp with tubular body widened
and flattened into bell shape.

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 Polyp Form

Tubular body with the mouth directed upward and surrounded

by tentacles.
Mouth leads into a blind gastrovascular cavity.
Aboral end attached to substratum by pedal disc.
Reproduce asexually by budding, fission, or pedal laceration.
• If a bud detaches from the polyp that made it, it forms a
clone.
• If a bud remains attached to the polyp that made it, it
becomes a colony with shared gastrovascular cavity.

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 Polymorphism

A shared gastrovascular cavity permits specialization of

polyps and results in many morphologically distinct types of
polyps.

• In colonial forms, polyps may be specialized for feeding,

reproduction, or defense.
• Individual polyps are called zooids.

• In class Hydrozoa feeding polyps (hydranths)

distinguished from reproductive polyps (gonangia) by
absence of tentacles in gonangia.

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 Medusa Form

Bell- or umbrella-shaped and exhibit tetramerous symmetry

(arranged in fours).
Usually free-swimming or free-floating.
Mouth centered on the concave side (subumbrellar region)
directed downward into frilly lobes that may extend beneath
the bell.
Tentacles may extend down from the rim of the umbrella.
Medusae equipped with statocysts (orientation) and ocelli
(light sensors).
Integration of sensory information into motor response by a
nerve ring located at base of the bell with hydrozoans having
two rings.
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 Hydromedusae and Scyphomedusae

Hydromedusae (medusa of hydroids) differ from

scyphomedusae (medusa of schyphozoa) by the presence of
velum
• Shelf-like fold of tissue from the bottom of the bell that
extends into the bell
• Velum reduces the cross-sectional area at the bottom of
the bell
• Increases the exit velocity of water from the bell when the
velum pulsates to make swimming more efficient.
This is an important distinguishing feature between hydroids
and true jellyfishes.

© McGraw Hill 15
 Medusa and Polyp Body Plans

Figure 13.3
Comparison
between polyp and
medusa types of
individuals.

Access the text alternative for slide images.

© McGraw Hill 16
 Life Cycles

Polyps and medusa play different roles in the cnidarian life

cycle.
Typically, zygote develops into a motile planula larva.
Planula settles on hard substrate and metamorphoses into a
polyp.
• Produce other polyps asexually.
Polyps eventually produce a free-swimming medusa by
asexual reproduction (budding or strobilation).
Medusae are generally dioecious and reproduce sexually.

© McGraw Hill 17
 Life Cycle Variations

True jellyfish (class Scyphozoa) have typical life cycle.

• Have both attached and free-swimming stages that
occupy both pelagic (open water) and benthic (bottom)
environments.
• Medusa is large and conspicuous.
• Polyps are typically very small.
Most colonial hydroids.
• Feature a polyp stage that form colonies and a pelagic
medusa stage.
• Some hydrozoans (Physalia sp.) form floating colonies—
the Portuguese man-of-war.
• Hydra only have small freshwater polyps.
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 Body Wall 1

Cnidarian body has.

• Outer epidermis derived from ectoderm.
• Inner gastrodermis derived from endoderm.
• Mesoglea in between.
Gastrodermis lines the gut cavity and plays a role in
digestion.
In Hydra sp., the epidermis contains epitheliomuscular,
interstitial, gland, sensory, cnidocytes, and nerve cells.

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 Anatomy of Body Wall and Cnidocytes

Figure 13.4 Left: A hydra polyp showing the gastrovascular lining. Center:
Portion of the body wall of a hydra. Right: Structure of a stinging cell.

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 Epitheliomuscular Cells

Cnidarian bodies extend, contract, bend, and pulse with no

mesodermally derived muscle cells.
• Epitheliomuscular cells, which form most of epidermis, both cover
organism and function in muscle contraction.

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 Epitheliomuscular and Nerve Cells

Figure 13.5 Epitheliomuscular and nerve cells in hydra.

© McGraw Hill 22
 Mesoglea

Mesoglea is located in-between and attached to the

epidermis and gastrodermis.
• Gelatinous (at least 95% water).
• Epidermal and gastrodermal cells send processes into it.
• Continuous layer over body and tentacles in polyps.
• Thick mesogleal layer containing amoeboid cells in
anthozoans and scyphozoan medusae; scyphozoans also
have fibers in mesoglea.
• Hydromedusa mesoglea lacks amoeboid cells and fibers.

© McGraw Hill 23
 Cnidocytes

Many cnidarians are effective predators due to cnidocytes.

Cnidocytes are located in the invaginations of ectodermal
cells and some endodermal cells.
• Produce one of over 20 types of cnidae that can be
discharged, reabsorbed, and replaced.
One type of cnida is the nematocyst, which can inject toxin
for prey capture or defense.
• Nematocysts are tiny capsules made of chitin-like material
and containing a coiled filament.
• End of capsule is covered by operculum.
• Filament may have tiny barbs or spines.

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 Nematocyst Discharge

Cnidocytes have trigger-like cnidocil (modified cilium).

• Except Anthozoa, which have modified ciliary
mechanoreceptor, tactile stimulation causes nematocyst to
discharge.
Discharge due to the combination of tensional forces and
high osmotic pressure in the nematocyst.
When stimulated to discharge, high internal osmotic pressure
causes water to rush in, increasing hydrostatic pressure.
The operculum opens and rapidly launches the filament from
the capsule with spiny barbs that inject poison into prey.
Only a few box jellyfishes and the Portuguese man-of-war
can seriously harm humans.
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 Various Types of Cnidae

Figure 13.6 (A) Several types of cnidae shown after discharge. At bottom are two
of a type that does not impale prey. (B) Fired and unfired cnidae from Corynactis
californica.
© McGraw Hill ©Thien T. Mai 26
 Feeding and Digestion in Polyps

Polyps are typically carnivorous.

Catch prey with tentacles, then pass it to gastrovascular cavity.
Gland cells discharge enzymes to begin extracellular digestion.
Intracellular digestion continues in cells of the gastrodermis.
• In colonial hydrozoans, polyps capture prey and begin digestion
in the mouth, then pass food into a common gastrovascular
cavity where intracellular digestion occurs.
Anthozoan polyps are mostly carnivorous.
• Expand and stretch tentacles in search of prey of any suitable
size.
• Some feed on small forms by ciliary currents.
• Others have symbiotic algae within body.

© McGraw Hill 27
 Feeding and Digestion in Medusae

Hydromedusae are similar to polyps.

• Their body is oriented with mouth facing down at the
center of the bell.
• Mouth is at end of tube, the manubrium.
Scyphomedusae are larger than hydromedusae.
• Have extended mouth edge (manubrium) with four frilly
oral arms or lobes that are used in capturing and ingesting
prey.

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 Nerve Net

Cnidarians have a diffuse nervous system.

• Plexus of nerve cells at base of epidermis and plexus at
base of gastrodermis are interconnected into a nerve net.
• Nerve processes (axons) end on the synapses of sensory
or effector organs.
• Nerve action potentials transmitted across synapses by
neurotransmitters.
• Unlike higher animals, cnidarian nerve nets have neurotransmitters
on both sides.
• Allow faster transmission in either direction.

• No myelin sheath that insulates axons.

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 Cnidarian Neuromuscular System

Neuromuscular system is formed by nerve cell synapses with

slender sensory cells, epitheliomuscular cells, nematocysts,
and contractile fibers.
This nerve net pattern is also found in annelid and human
(nerve plexus) digestive systems that coordinates rhythmic
peristalsis of the stomach and intestines.
Cnidarians do not have local concentration of nerve cells to
form a central nervous system as in higher animals.
• Nerve net and ring system allows processing and
response to stimuli from all directions.

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 Medusan Nervous Systems

Scyphomedusae and cubozoan medusae.

• Have sense organs along the margins of the medusa
called rhopalia that house chemoreceptors, statocysts,
and ocelli.
• Have two nerve systems, a fast-conducting system to
coordinate swimming and a slower one that controls
tentacle movement.
In hydromedusae, epidermal nerve net is condensed into two
nerve rings at bell margin.
• Process information from sense organs and respond by
changing swimming direction, pulsation rate, and position
of tentacles.
© McGraw Hill 31
 Class Hydrozoa

Mostly marine and colonial with asexual polyp and sexual

medusa stages, as in Obelia sp.
Typical hydroid has a base, a stalk, and one or more terminal
zooids (individual polyp animals).
• Base is a rootlike stolon, or hydrorhiza.
• Stolon gives rise to stalks called hydrocauli.
• Living part of the hydrocaulus is a tubular coenosarc.
• Hydrocaulus covered by a nonliving chitinous sheath, the
perisarc.

© McGraw Hill 32
 Hydroid Life Cycle

Figure 13.7 Life cycle of Obelia sp., showing alternation of polyp

(asexual) and medusa (sexual) stages.

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 Hydranths

Attached to the hydrocaulus are individual zooids or polyp cells.

• Hydranths (gastrozooids) are feeding polyps which are tubular,
bottle-shaped or vase-like and form a circle of tentacles
surrounding mouth.
• Hydranths may be thecate, where the perisarc forms a
protective cup around the polyp, or athecate, which is a naked
polyp.
• Hydranths capture and ingest prey ranging from worms, tiny
crustaceans, and various larvae so as to provide nutrition for
the whole colony.
• Food is partially digested in the hydranth (extracellular
digestion), then moves to common gastrovascular cavity where
ciliated gastrodermis helps circulate it, moving food into
gastrodermal cells for digestion.
© McGraw Hill 34
 Athecate Hydroid
Figure 13.8 Athecate
hydroids. (A)
Ectopleura integra,
with naked hydranths
and gonophores. (B)
Corymorpha sp. has
free-swimming
medusa with a single
trailing tentacle.

© McGraw Hill ©William C. Ober/Medical Scientific Illustration 35

 Hydroid Reproduction

Colonial hydroids bud off new individuals to increase size of

colony.
• Individuals may be new feeding hydranths or medusae buds.
• In Obelia sp., the medusae bud from reproductive polyp called
a gonangium.
• Young medusae leave the colony as free-swimming individuals
that mature and form gametes.
• Some medusae remain attached to the colonies and shed their
gametes.
• Some species do not have medusae but shed male and female
gonophores.
• Ciliated embryo called planulae are formed.

© McGraw Hill 36
 Hydroid Gonophores

Figure 13.9 In some hydroids like Tubularia crocea, medusa are reduced
to gonadal tissue, gonophores, and do not detach.
Access the text alternative for slide images.

© McGraw Hill ©William C. Ober/Medical Scientific Illustration 37

 Hydroid Medusae

Hydroid medusae are usually smaller than schyphozoan

medusae.
Margin of the bell projects inward as a shelf-like velum that
partly closes the bell and aids in swimming.
Tentacles around bell have many nematocysts.
Mouth opens at the end of a suspended manubrium and
connects to a stomach and four radial canals.
Radial canals connect to a ring canal that runs around the
margin of the bell and connects with the hollow tentacles.
Gastrovascular cavity is then continuous from mouth to
tentacles.
© McGraw Hill 38
 Hydroid Bell Medusa

Figure 13.10 Bell medusa, Polyorchis penicillatus

© McGraw Hill ©Daniel W. Gotshall 39

 Hydroid Nervous and Muscular Systems

Nerve net forms two rings around the base of the velum.

Bell margin has many sensory cells.

• Typically bears statocysts, specialized sense organs that

function in equilibrium, and light-sensitive ocelli.

Most cnidarians do not have true mesodermal muscles, but

hydromedusae have smooth and striated muscles derived
from ectoderm called the entocodon.

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 Structure of Hydroid Medusa

Figure 13.11 Structures of Gonionemus sp.

© McGraw Hill 41
 Freshwater Hydroid Medusae

Craspedacusta sowberii.
• Most likely evolved from marine ancestors in the Yangtze
River of China.
• Now found in Europe, the United States, and parts of
Canada.
• Medusae may reach a diameter of 20 mm.
• Polyps are tiny (2 mm), simple body with no perisarc and
no tentacles.
• Polyp employs three methods of asexual reproduction:
Budding off new individuals, constriction of planula, and
medusa buds.
© McGraw Hill 42
 Life Cycle of Freshwater Hydrozoan

Figure 13.12 Life cycle of Craspedacusta sp., a freshwater hydrozoan.

© McGraw Hill 43
 Hydra: Solitary Hydrozoan

Hydra is found on the underside of aquatic leaves and lily

pads in cool, clean freshwater.

Worldwide distribution (16 species in North America).

Body is a cylindrical tube that can extend or contract to a tiny

gelatinous mass.

Aboral end forms a slender stalk and end in a basal or pedal

disc for attachment.

The mouth (oral end) on a conical elevation, the hypostome

with 6 to 10 hollow tentacles around the mouth.

© McGraw Hill 44
 Feeding in Hydras

The mouth opens to a gastrovascular cavity that connects to

the cavities of the tentacles.

Hydras feed on a variety of small crustaceans, insect larvae,

and worms.

Food organisms that brush against the tentacles are

captured by nematocysts.

Helpless organisms moved by tentacles, and engulfed by the

mouth.

Inside gastrovascular cavity, gland cells discharge enzymes

to digest food.

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 Digestion in Hydras

Digestion is extracellular but some food particles are moved

by pseudopodia into nutritive-muscular cells of the
gastrodermis where intracellular digestion occurs.

Nutritive-muscular cells are tall columnar cells with lateral

extended bases containing myofibrils.

• Myofibrils run at right angles to the body axis, forms a

weak circular muscle layer.

Body and tentacle extension mostly due to water that enters

the gastrovascular cavity due to beating cilia.

• Water serves as a hydrostatic skeleton.

© McGraw Hill 46
 Hydra Catching Prey

Figure 13.13 Hydra catches an unwary water flea with the nematocysts of its
tentacles. This hydra already contains one water flea, eaten previously.

© McGraw Hill ©Tom Branch/Science Source 47

 Hydra Epidermis

Epidermal layer contains many types of cells with a variety of

functions.
• Epitheliomuscular cells.
• Form most of the epidermal covering and for muscular contraction.

• Interstitial cells.
• Undifferentiated stem cells that can develop into cnidoblasts, sex
cells, buds, or nerve cells (but not epitheliomuscular cells).

• Gland cells.
• Tall cells around the basal disc and mouth which secrete the
adhesive substances and sometimes a gas bubble for floating.

© McGraw Hill 48
 More Epidermal Cell Types

Cnidocytes (three types).

• Penetrants—Penetrate prey and inject poison.
• Volvents—Recoil and entangle prey.
• Glutinants—Secrete adhesive for locomotion and
attachment.
Sensory cells.
• Scattered among epidermal cells near mouth, tentacles,
and basal discs.
• Have a flagellum for chemical and tactile stimuli and as
synapse with nerve cells.
Nerve cells.
• Generally multipolar with both one- and two-way synapses.

© McGraw Hill 49
 Reproduction in Hydras

Asexual reproduction.
• Budding via outpockets of body wall that form young
hydras.
Sexual reproduction.
• Most hydra species are dioecious with temporary gonads
that appear in autumn as stimulated by lower temperatures
or stagnation of water.
• Eggs and sperm shed externally into water.
• Zygotes undergo holoblastic cleavage to form hollow
blastula.
• Cyst forms around embryo to survive the winter and young
hydras hatch in the spring.
© McGraw Hill 50
 Budding Hydra

Figure 13.14 Hydra with developing bud and ovary.

© McGraw Hill ©M. I. Walker/NHPA 51

 Other Hydrozoans

Orders Siphonophora and Chondrophora form polymorphic

swimming or floating colonies.
• Contain modified medusa.
• Contain several types of polyp individuals.
• Gastozooids are feeding polyps with single long tentacle.
• Dactylozooids are fishing tentacles that sting prey and lift them to
feeding polyps.
• Gonophores are sacs containing ovaries or testes.

Other hydrozoans secrete calcareous skeletons resembling

true corals and are called the hydrocorals.

© McGraw Hill 52
 Physalia

Physalia sp., commonly known as the Portuguese man-of-

war, is a colony of hydroids.
• Has a float called a pneumatophore that allows it to move
along the surface waters of tropical seas.
• The float is thought to have expanded from the original
larval polyp and act as nurse-carrier for future generations
of buds.
• Has long, graceful tentacles called zooids that contain
many nematocysts.

© McGraw Hill 53
 A Portuguese Man-of-War Colony

© McGraw Hill ©Wild Horizon/Getty Images 54

 Hydrocorals

Figure 13.16 These

hydrozoans form
calcareous skeletons that
resemble true coral.

Stylaster roseus

Species of Millepora
with nematocyts

© McGraw Hill A:©Larry Roberts/McGraw -Hill Education; B:©William C. Ober/Medical Scientific Illustration; C:©William C. Ober/Medical Scientific Illustration 55
 Class Scyphozoa

Most of the larger jellyfishes belong to this group and are

also called “cup animals.”
Nearly all float and swim in open seas and can reach up to
depths of 3000 meters.
Bells vary in shape and size but Scyphozoans lack shelf-like
velum found in hydrozoan medusae.
Tentacles around the bell may be numerous or few and can
be short or long.
Margin of the umbrella has indentations, each bearing a pair
of lappets.

© McGraw Hill 56
 The Giant Jelly

Figure 13.17 Giant jelly, Cyanea capillata. It is called the “sea blubber” by
fishermen.

© McGraw Hill ©Rick Harbo 57

 Scyphozoan Structures

Between lappets is a equilibrium sense organ, a rhopalium.

• Club shaped, it contains a hollow statocyst that is used for
equilibrium.
The nervous system consists of a nerve net with one set
controlling bell pulsations and the other more diffused one
controlling feeding.
Tentacles, manubrium, and often entire body may have
nematocysts.
Mouth located beneath the umbrella and leads to the
stomach.
Manubrium forms four oral arms to capture and ingest prey.
© McGraw Hill 58
 Scyphozoan Digestion

Scyphozoans have four gastric pouches that have small

tentacle projections called gastric filaments.
• Filaments (lacking in hydromedusae) are covered with
nematocysts.
Radial canals branch out from gastric pouches and connect
to ring canals around the margin and forms part of the
gastrovascular cavity.
Food items are caught in mucus on the umbrella surface,
and are carried to “food pockets” on the umbrella margin by
cilia.
Cilia in the gastrodermis layer keep a current of water
moving to bring food and oxygen into the stomach and to
expel wastes.
© McGraw Hill 59
 Cosmopolitan Moon Jelly

Figure 13.18 Moon jellyfish Aurelia aurita.

© McGraw Hill ©Larry Roberts/McGraw -Hill Education 60
 Scyphozoan Reproduction

Scyphozoans have separate sexes with gonads in the gastric

pouches.

Fertilization is internal with sperm carried by ciliary currents

into the gastric pouch of the female.

Zygote develops into a ciliated planula larva and later

attaches to the substrate and develops into a scyphistoma
which has a hydra-like form.

Scyphistoma undergoes strobilation and form saucer-like

buds called ephyrae that break loose to form jellyfish
medusae.

© McGraw Hill 61
 Moon Jelly Life Cycle

Figure 13.19 Life cycle of Aurelia sp., a marine scyphozoan medusa.

© McGraw Hill 62
 Class Staurozoa

Commonly called stauromedusans, these have no medusa

stage.

Solitary polyp body that is stalked and uses adhesive disk to

attach to seaweeds and objects on the sea bottom.

Polyp top resembles a medusa with eight extensions (“arms”)

ending in tentacle clusters surrounding mouth.

Polyps reproduce sexually and the nonswimming planula

develops directly into new polyp.

© McGraw Hill 63
 Staurozoan Polyp

Figure 13.20 Thaumatoscyphus hexaradiatus are an example of class

Staurozoa.
© McGraw Hill ©Rick Harbo 64
 Class Cubozoa

Medusa form is the predominant stage with polyp form being

inconspicuous or unknown.

Umbrella is square and one or more tentacles extend from

each corner.

At base of each tentacle is a flat blade called a pedalium.

Rhopalia house three sets of unique eyes—two types of

ocelli and one set of camera type eyes with cornea and lens.

© McGraw Hill 65
 Cubozoan Features

Umbrella edge turns inward to form a velarium that increases

the swimming efficiency.

Feed mostly on fish in nearshore areas.

Polyp stage is tiny, solitary, and sessile.

New polyps bud laterally, detach, and float away.

Do not produce ephyrae, but directly change into medusae.

The sea wasp (Chironex fleckeri) has caused several

fatalities in Australia.

© McGraw Hill 66
 Structure of a Cubozoan

Figure 13.21 Carybdea sp., a cubozoan medusa.

© McGraw Hill 67
 Class Myxozoa

Obligate parasites that typically have two hosts.

• Fish.
• Have damaged salmon, whitefish, and trout populations.

• Freshwater annelid.
Unique cell division.
• New cells grow inside older cells.
Have polar capsules that are homologous to nematocysts.
One of the smallest genomes in the animal kingdom.

© McGraw Hill 68
 Class Anthozoa

Called “flower animals” as most are polyp form and lack a

medusa stage.
All marine, in both deep and shallow water, and vary in size.
Two general groups based on body plan:
• Hexamerous plan: Multiples of six—also called
polymerous symmetry; tentacles in one of more circles
around the oral disc.
• Octomerous plan: Built on a plan of eight with tentacles
having eight featherlike tentacles around the mouth.

© McGraw Hill 69
 “Flower Animals”

Figure 13.22 Sea anemones, Tealia piscivora, are the familiar and
colorful “flower animals.”

© McGraw Hill ©Daniel W. Gotshall 70

 Anthozoan Subclasses

Three subclasses in the group:

• Hexacorallia (Zoantharia): Sea anemones and hard corals.
• Ceriantipatharia: Tube anemones and thorny corals.
• Octocorallia (Alcyonarias): Soft corals, sea fans, and sea
pens.
Solitary or colonial with some forms having hard skeleton.
Large gastrovascular cavity divided by septa (mesenteries).
• Inward extensions of body wall that are coupled.
• Hexacorallian septa are also paired.

© McGraw Hill 71
 Sea Pen and Gorgonian

Figure 13.23 (A) White sea pen Pteroeides sp. from the Solomon
Islands. (B) Close-up of a gorgonian with pinnate tentacles characteristic
of subclass Octocorallia.
© McGraw Hill a:©Diane Nelson; b:©Larry Roberts/McGraw -Hill Education 72
 Body Wall 2

Usually circular muscles in the body wall.

Usually longitudinal and transverse muscles in the septa.

Mesoglea is a mesenchyme containing ameboid cells.

© McGraw Hill 73
 Sea Anemones

Polyps are larger and heavier than hydrozoan polyps and

can be very colorful.

Found in warm water coastal areas attached to shells, rocks,

timber, etc. by pedal discs.

Some burrow in mud or sand.

Have cylindrical body form with crown of tentacles in one or

more circles around the mouth of a flat oral disc.

Slit-shaped mouth leads into a pharynx.

© McGraw Hill 74
 Sea Anemone Structure

Figure 13.24 Structure of a sea anemone.

© McGraw Hill 75
 Anemone Digestive Structures

Siphonoglyph (ciliated groove) creates a water current

directed into the pharynx.
Ciliary motion transports oxygen, removes wastes, and
maintains fluid pressure for a hydrostatic skeleton.
Pharynx leads to gastrovascular cavity.
• Divided by six pairs of primary septa (mesenteries)
extending vertically from body wall to pharynx.
• Smaller incomplete septa partially subdivide the large
chambers and increase surface area of the gastrovascular
cavity.

© McGraw Hill 76
 Septal Filaments

Free edge of each incomplete septum forms a sinuous cord

called septal filament.

• Contains nematocysts and gland cells used for digestion.

In some, lower end of filaments extend into acontia threads.

• Also equipped with nematocysts and gland cells.

• May protrude through mouth or body pores to help capture

prey.

• When in danger, water is rapidly expelled through pores

as the anemone contracts to a smaller size.

© McGraw Hill 77
 Feeding and Musculature

Sea anemones are carnivorous.

• Zoantharian feeding behavior often under chemical
control, responding to asparagine and glutathione levels.
Longitudinal muscles of the epidermis occur in the tentacles
and oral disc.
Longitudinal muscles of the column are gastrodermal and
located in the septa.
Most anemones can glide slowly on pedal discs and some
can swim with limited ability by rhythmic bending.

© McGraw Hill 78
 Swimming Anemone

Figure 13.25 (A) A sea anemone that swims. (B) When attacked by a
predatory sea star Dermasterias sp., the anemone Stomphia didemon
detaches from the bottom and rolls or swims to a safer location.

© McGraw Hill (a,b): ©Rick Harbo 79

 Anemone Mutualism

Most harbor symbiotic dinoflagellates called zooxanthellae

within their tissues and benefit from products of algal
photosynthesis.

Some anemones attach to the shells of hermit crabs and

provide protection for the crab while getting free
transportation and food scraps from the crab.

Damselfishes (anemone fishes) form associations with large

anemones whereby the fish provide ventilation and clean-up
for the anemone while getting shelter and protection.

© McGraw Hill 80
 Fish-Anemone Mutualism

Figure 13.26 Orangefin anemone fish (Amphiprion chrysopterus)

nestles in the tentacles of its sea anemone host.

© McGraw Hill ©Larry Roberts/McGraw -Hill Education 81

 Anemone Reproduction

Some anemones have separate sexes (dioecious).

Some are hermaphroditic or monoecious.
• Monoecious species are protandrous so produce sperm
first and eggs later.
Gonads on margins of septa and fertilization is usually
external or may occur in the gastrovascular cavity and forms
a ciliated larva.
Asexual reproduction occur by pedal laceration where small
pieces of pedal disc break off and regenerate a small
anemone.
Longitudinal and transverse fission and budding may also
occur.
© McGraw Hill 82
 Hexacorallian Corals

Also called true or stony corals of the order Scleractinia.

• Look like miniature sea anemones that live in calcareous
cups they have secreted.
Gastrovascular cavity is hexamerous.
Hollow tentacles surround mouth.
No siphonoglyph.
No pedal disc.
Secrete a limestone skeletal cup with sclerosepta projecting
up into the polyp between true septa.

© McGraw Hill 83
 Stony Corals

Figure 13.27 (A) Cup coral Tubastrea sp. is not a reef-building coral
(ahermatypic) and has no symbiotic zooxanthellae. (B) Polyps of
Montastrea cavernosa are tightly withdrawn during daytime but open to
feed at night, as seen in (C).

© McGraw Hill (all): ©William C. Ober/Medical Scientific Illustration 84

 Hexacorallian Exoskeleton

Sheet of living tissue forms over the exoskeleton and allows

polyps to retract into it when not feeding.

All gastrovascular cavities of the polyps are interconnected

by this tissue sheet.

In many colonial stony corals, the exoskeleton can grow to

massive sizes as the living tissue continues to build up over
many years resulting in extensive coral reef formations.

© McGraw Hill 85
 Hexacorallian Structure

Figure 13.28 Polyp of a hexacorallian coral.

© McGraw Hill 86
 Boulder Star Coral

Figure 13.29 Boulder star

coral, Montastrea annularis.

© McGraw Hill ©Larry Roberts/McGraw -Hill Education 87

 Tube Anemones and Thorny Corals

Members of subclass Ceriantipatharia have coupled but

unpaired septa.
Tube anemones (order Ceriantharia) are solitary and live
buried to the level of oral discs in soft sediments.
• They occupy tubes made with secreted mucus that they
can retract into.
Thorny or black corals (order Antipatharia) form colonies
attached to firm substrata.
• Skeletons are thorny and have few species limited to
warmer seas.

© McGraw Hill 88
 Tube Anemone

Figure 13.30 A tube anemone of subclass Ceriantipatharia, order

Ceriantharia.
© McGraw Hill ©William C. Ober/Medical Scientific Illustration 89
 Thorny Coral

Figure 13.31 (A) Colony of Antipathes sp., a black or thorny coral most
abundant in deep waters in the tropics. (B) The polyps of Antipatharia
have six simple, nonretractile tentacles that have spiny processes.

© McGraw Hill a: ©William C. Ober/Medical Scientific Illustration 90

 Octocorallian Corals

Octomerous symmetry with eight pinnate tentacles and eight

unpaired complete septa.
All colonial with gastrovascular cavities that connect through
tubes called solenia.
• Solenia pass through an extensive mesoglea
(coenenchyme).
Colony surface is covered by epidermis.
Skeleton is secreted in coenenchyme.
• Combination of limy spicules, fused spicules, and protein
forming an endoskeleton.

© McGraw Hill 91
 Octocorallian Structure

Figure 13.32 Polyps of an octocorallian coral.

© McGraw Hill 92
 Octocorallian Soft Coral

Figure 13.33 A soft coral, Dendronephthya sp., an octocorallian on a

Pacific Ocean coral reef.
© McGraw Hill ©Larry Roberts/McGraw -Hill Education 93
 Colonial Gorgonian Corals

Figure 13.34 Colonial gorgonian, or horny, corals from the western

Pacific. (A) Red gorgonian Melithaea sp. (B) A sea fan, Subergorgia
mollis. (C) Red whip coral, Ellisella sp.

© McGraw Hill (a-c): ©Larry Roberts/McGraw-Hill Education 94

 Coral Reefs

Great diversity of organisms.

One of the most productive ecosystems rivaled only by
tropical rainforests.
Seaweeds and animals are limited to top layer of large
calcium carbonate (limestone) deposits, normally in shallow
seas that were formed over thousands of years.
Scleractinian hermatypic corals (reef-building) and coralline
algae form most coral reefs around the world.

© McGraw Hill 95
 Coral Reef Requirements

Hermatypic corals require warm waters, abundant light, and

salinity of undiluted sea water.
Reefs limited to shallow waters found between 30 degrees
north and 30 degrees south latitude that excludes areas of
cold water upwelling and major river outflows with high
turbidity.
Photosynthetic zooxanthellae live in their tissues and provide
food for corals while enhancing deposition of calcium
skeleton.
Also tightly recycle phosphorus and nitrogenous wastes
within reefs.

© McGraw Hill 96
 Coral Symbiosis

Symbiosis between zooxanthellae and corals is threatened

by ocean warming.
• Warmer water damages the photosynthetic mechanism in
zooxanthellae.
• Harmful oxidants build up and diffuse into coral tissues.
• Zooxanthellae die or are expelled and coral tissues turn
white and brittle, resulting in coral bleaching.
• White color of bleached corals worsens problem due to
reflective color bringing even more light into the affected
pathway.
• Bleaching has occurred prior to recent global warming
measurements.
© McGraw Hill 97
 Healthy and Bleached Polyps

Figure 13.35 A comparison of healthy and bleached polyps within a colony of the
zoanthid Palythoa caribbaeorum in La Parguera, Puerto Rico.

Photo courtesy of Ernesto Well, Dept. of M arine Sciences, University of Puerto Rico.

© McGraw Hill ©Ernesto Weil 98

 Types of Coral Reefs

Fringing reef.
• Near the land with no lagoon or a very narrow lagoon.
Barrier reef.
• Parallel to shore with a wide and deep lagoon.
Atolls.
• Encircle a lagoon but not an island.
• Have a steep bank on the seaward slope.
Patch or bank reefs.
• Occur at some distance back from any steep seaward
slope of lagoons and reefs.
© McGraw Hill 99
 Zones of Coral Reefs

Reef front or fore reef slope.

• Facing the sea which is parallel to shore.
• Can gently slope toward deeper water or may drop
suddenly.
Reef crest.
• Usually shallow water or emerging at the top of the reef
front.
• Upper front and crest get most wave action and breaks
pieces off.
Reef flat.
• Slopes toward the shore and receives debris and coralline
sand.
© McGraw Hill 100
 Structure of a Coral Reef

Figure 13.36 (A) Profile of a barrier reef. (B) Portion of an atoll from the air.

© McGraw Hill ©Cleveland P. Hickman, Jr. 101

 Coral Reef Significance

Coral reefs form highly irregular walls facing the sea with
numerous grooves, caves, crevices, and channels that can
support a diversity of organisms ranging from invertebrates
to vertebrates.

Few nutrients enter or leave the coral reef system and little is
lost due to efficiency in recycling among interacting
organisms.

The loss of coral reefs due to pollution or catastrophic events

can result in major economic and ecological impacts.

© McGraw Hill 102

 Phylum Ctenophora

Ctenophora is composed of about 150 species, commonly

called “sea walnuts” or “comb jellies.”
All marine forms and most prefer warm waters.
Have eight rows of comb-like plates of cilia used for
locomotion and normally follow tides and ocean currents.
Nearly all free-swimming, few creep or are sessile, and range
from shallow water to deep ocean.

© McGraw Hill 103

 Pleurobrachia

Pleurobrachia sp. is a representative ctenophore.

• Two tentacles on body produces biradial symmetry; are
normally long, solid, and extensible or can be retracted
into tentacle sheaths.
• Has no definite head but has oral–aboral axis with mouth
leading from pharynx into a branched digestive tract
ending in anal pore.
• Have a sensory organ called statocyst on the aboral pole.
• Has eight equally spaced bands called comb rows that
extend as meridians from the aboral to the oral region.

© McGraw Hill 104

 Comb Jellies

Figure 13.37 (A) Comb jelly Pleurobrachia sp. Its fragile beauty is evident
at night when it luminesces from its comb rows. (B) Mnemiopsis sp.

© McGraw Hill a: ©William C. Ober/Medical Scientific Illustration; b:©Gregory G. Dimijian/Science Source 105
 Ctenophoran Features

Comb plates are long, fused cilia that form transverse plates
across the body.
• Movement by cilia beating on the comb plates.
Body is transparent with gelatinous layer called collenchyme
derived from ectoderm and endoderm.
Gelatinous layer has extensive muscle fibers forming radial,
meridional, and latitudinal banding patterns.
Muscle cells are distinct and are not contractile portions of
epitheliomuscular cells as in cnidarians.

© McGraw Hill 106

 Ctenophoran Tentacles

Ctenophore tentacles capture planktonic prey from

surrounding waters using epidermal glue cells called
colloblasts.
• Then food-laden tentacles are passed across mouth.
Some ctenophores have short tentacles that collect food on
ciliated body surface.
Other ctenophores without tentacles may feed on gelatinous
creatures like medusae, salps, and other ctenophores.
Those that feed on cnidarians can collect undischarged
cnidocytes for incorporation into their own epidermal tissue
and use as a defense mechanism.

© McGraw Hill 107

 Ctenophoran Structure

Figure 13.38 Comb jelly Pleurobrachia sp. (A) External view. (B) Hemisection.
(C) Colloblast, an adhesive cell. (D) Portion of comb rows showing comb plates.
Access the text alternative for slide images.

© McGraw Hill 108

 Ctenophoran Digestion and Respiration

Gastrovascular system comprises a mouth, pharynx,

stomach, and canals that run to the comb plates, tentacular
sheaths, and other regions of the body.

Two blind canals terminate near mouth.

Aboral canal divides into two small anal canals that expel
wastes.

Digestion by both extracellular and intracellular.

Respiration and excretion occur by diffusion across the body

surface.

© McGraw Hill 109

 Nervous and Sensory Systems

Resembles cnidarian nerve network with subepidermal

plexus concentrated under each comb plate with no central
control.
Statocyst is a bell-like chamber with tufts of cilia that sense
changes in pressure from statolith as animal changes
position.
Epidermis has sensory cells that detect chemical and other
stimuli so that when unfavorable conditions occur, cilia
reverse their beat and moves organisms backward.
Comb plates are also sensitive to touch and can withdraw
into the animal when touched.

© McGraw Hill 110

 Reproduction and Development

Mostly monoecious (hermaphroditic) with gonads on the

lining of the gastrovascular canals under the comb plates.

Fertilized eggs discharged through epidermis into water.

Cleavage varies from determinate to indeterminate.

Free-swimming cydippid larva gradually develops into an

adult without metamorphosis.

© McGraw Hill 111

 Cydippid Larva

Figure 13.39 A cydippid larva.

© McGraw Hill ©Ronald L. Shimek, 2004 112
 Other Ctenophores

Beroe sp.
• Large conical or thimble-shaped ctenophore that has large
mouth but lacks tentacles.
Cestum sp., or Venus’ girdle.
• Band-like ctenophore over 1 meter long and use sinuous
body movements along with comb plates for locomotion.
Ctenoplana sp.
• Disc shaped flattened bodies that creep rather than swim.
Most ctenophores are bioluminescent at night like
Mnemiopsis sp.

© McGraw Hill 113

 Ctenophoran Diversity

Figure 13.40 Diversity among

phylum Ctenophora. (A) Beroe sp.
(B) Cestum sp. (C) Coeloplana sp.

© McGraw Hill 114

 Phylogeny of the Diploblasts

Ctenophores and cnidarians have typical diploblastic

characteristics.
• Gelatinous middle layer surrounded by outer epidermal
layer derived from ectoderm and inner gut lining derived
from endoderm.
• However, some cells of the gelatinous layer cause
problems as they form ectomesoderm.
• Most cnidarians have few cells within the mesoglea, so
little debate that they are diploblastic.
• However, during hydrozoan medusae stage, the
development of entocodon layer has been judged to be
triploblastic.
© McGraw Hill 115
 Diploblastic or Triploblastic?

In triploblasts, true muscles are produced by mesodermal

cells, yet the hydrozoan entocodon is ectodermal in origin.
Studies of gene expression show that genes homologous to
triploblast mesoderm were expressed as diploblast
endoderm.
Recent re-examination of the development of ctenophores
has led to the observation that muscle cells in the middle
layer originate from endodermal cells and not ectodermal
cells.
Since mesoderm derives from endoderm, if further studies
confirm the endodermal origins of ctenophore muscles, then
they are triploblastic.

© McGraw Hill 116

 Radial or Bilateral Symmetry?

Body symmetry also debated.

• Adult cnidarian is radially symmetrical and adult
ctenophore is biradially symmetrical.
• But, cnidarian planula larva swims with one end moving
forward.
• This end could be designated as anterior end giving the
planula a distinct anterior–posterior axis.
The questions remain.
• Did the radially symmetrical cnidarians have a bilaterally
symmetrical ancestor?
• Does the genetic potential for bilateral symmetry predate
the bilateral body plan?
© McGraw Hill 117
 Cnidarian Phylogeny

Relationships among cnidarian classes are still controversial

with a major issue being the life cycle.

• Which came first, the polyp or the medusa?

• One hypothesis postulates that the ancestral cnidarian

was a trachyline-like hydrozoan with a medusa stage.

• Another hypothesis suggests that the ancestral cnidarian

was an anthozoan polyp without a medusa in the life
cycle.

© McGraw Hill 118

 Adaptive Diversification

Cnidarians achieved large numbers of individuals and species with

high diversity considering the simplicity of body plan.
• Polyp and medusae have similar construction but medusa have
more sensory and locomotory capacities.
• Efficient predators ranging from feeding on prey larger than
themselves to feeding on small particles.
• Some have colonial forms that grow to great sizes, as in coral
reefs.
• Some show polymorphism and specialization of individuals in a
colony.
Ctenophores.
• Have comb plates and biradial symmetry.
• Vary in body shape and presence or absence of tentacles.
• Some have sessile or creeping lifestyle.
© McGraw Hill 119
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