Sponges (Phylum Porifera)

Sponges are
sessile, spending their lives anchored
to a solid surface underwater.
Most are marine although some live
in fresh water.
diploblastic; that is, the body wall is
made of two layers of cells with a
jellylike mesoglea between them;
The body wall is perforated with
pores (hence the name Porifera)
through which water containing food
particles is filtered. The water is
drawn in through the pores by collar cells like those found in choanoflagellates.
(Some sponges can process a volume of water more than 100,000 times their
own volume in the course of a day!)
dispersed by small, free-swimming larvae;
about 10,000 species known;
probably the most ancient of today's invertebrates, their fossils appear in the
geological record as far back as 635 million years.
Despite their simple body plan, sequencing shows that their genome (> 18,000
genes) contains many genes homologous to those found in much more complex
animals.

Cnidarians (Phylum Cnidaria)

Characteristics:

 diploblastic; two layers of cells

ectoderm and endoderm with a
jellylike mesoglea between them;
predominantly radial symmetry: body
parts (e.g., tentacles) arranged in whorls.
However, in some sea anemones, there is
only one plane through the tubular body
that divides it into two mirror-image
halves; thus revealing bilateral symmetry.
cnidoblasts: specialized cells that secrete a
stinging capsule called a nematocyst.
Food is taken through a mouth into
the gastrovascular cavity. The cavity is
also called a coelenteron and for many
years the name of this phylum
wasCoelenterata. There is no anus.
Sexual reproduction produces a free-swimming, ciliated larva called a planula.
The phylum contains about 10,000 species distributed in 3 classes:
o Hydrozoa Although the freshwater hydra is a much-studied
representative, it is not typical of the class.
Most members are
marine
colonial
produce two body forms: the sessile polyp (like the hydra) and the
free-floating medusa (which disperses the species)
o Scyphozoa Jellyfishes (the medusa stage is dominant). The jelly of the
medusa is a much-enlarged mesoglea.
o Anthozoa Sea anemones and corals. Have only the polyp stage.
External Link

Site devoted to the Cnidaria with illustrations

Please let me know by e-mail if you find a broken link in my pages.)

Bilaterians
All the remaining groups of animals belong in a clade whose members share:
bilateral symmetry (hence the name); that is, dorsal-ventral and leftright axes
triploblastic (three tissue layers: ectoderm, mesoderm, endoderm)
HOX genes in one or more clusters with the genes within a cluster arranged in
the same order as the body parts they affect.
The bilaterians contain two clades, the protostomia and the deuterostomia.

Protostomia vs. Deuterostomia

Long before the days of genome analysis, taxonomists were convinced of a
fundamental division in the animal kingdom between the
protostomes ("first mouth") and the
deuterostomes ("second mouth").
Protostomia

Deuterostomia

Blastopore forms future mouth (in most

groups).

Blastopore forms future anus. Mouth forms later.

Few HOX genes for the posterior

Multiple HOX genes for the posterior

Spiral cleavage of embryo

Perpendicular cleavage planes in embryo [View]

Early cleavage cells committed [Link]; no

identical twins

Early cleavage cells totipotent; identical twins

possible

Coelom arises by splitting of mesoderm

Coelom arises between invaginating mesoderm

during gastrulation

Lophotrochozoans and Ecdysozoans

Echinoderms, Acorn worms, and Chordates

Let's first examine the protostomes. The deuterostomes are discussed below.

Lophotrochozoans vs. Ecdysozoans

Genome analysis, especially the analysis of
18S rRNA genes and
HOX genes
supports a major division of the Protostomia into two superphyla:
Lophotrochozoans and
Ecdysozoans

Lophotrochozoans
Their name was created from the names of formerly-separated groups that have now
been joined in a single clade on the basis of the similarities of their genomes:
They all share a cluster of HOX genes quite different from those found in the
ecdysozoans (and deuterostomes).
They share similar sequences in their 18S rRNA genes.
The clade contains a number of phyla of which we shall examine only 3.
flatworms (Platyhelminthes),
annelids (Annelida), and
mollusks (Mollusca).

Flatworms (Phylum Platyhelminthes)

This phylum contains some 20,000 species distributed among three classes:

 Turbellaria, free-living forms of which

the planarian (right) is a commonly-studied example.
Planaria share with the other members of the phylum
o a flat, almost ribbonlike, shape
o bilateral symmetry.
The bilateral symmetry of planarians is associated with
o active locomotion
by secreting a layer of mucus underneath
them and propelling themselves forward with the many cilia on
their ventral surface.
and by swimming.
o a concentration of sense organs in the head (called cephalization).
Planarians feed through a mouth on their ventral surface. It leads to an
elaborate gastrovascular cavity. But there is no separate exit so undigested food
has to leave by the mouth.
Trematoda, a group of
parasitic
o lung flukes
o liver flukes and
o blood flukes
(e.g., Schistosoma)
All of these have at least two
different stages in their life
cycle, each parasitic in a
different host one of which
is usually a snail.

The diagram gives the life cycle of the blood fluke, Schistosoma mansoni.
Once within the alternate host, a snail, a single miracidium may produce as
many as 200,000 infectious cercariae. Both sexes must infect the human if the
cycle is to continue. With the increasing use of irrigation in tropical regions, the
incidence of human infection known as schistosomiasis or bilharzia is
rising alarmingly.
How schistosomes evade the immune defenses of their host.

Cestoda; the parasitic tapeworms. They, too, alternate between an

intermediate host (e.g., pig, fish) and a definitive host (e.g., us). The growing
popularity of sushi and sashimi made of raw Pacific salmon has caused
infections by the fish tapeworm to become more common in the U.S.
Link to page illustrating the life cycles of the fish and pig tapeworms.

Annelids (Phylum Annelida)

Characteristics:
segmented; that is, their body is made up of
repeating units. Although some structures, e.g., the
digestive tract, run straight through, others like the
excretory organs are repeated in each segment.
The major nerve trunk runs along the ventral side.
a large, fluid-filled coelom; It is lined with mesoderm and enables the internal
organs to slide easily against one another making for easy locomotion.
There are >15,000 species known. Some examples:
the common earthworm [View]
leeches [View]
marine forms such as the clam worm [View] These animals produce a freeswimming trochophore larva (figure), which partly accounts for the name
Lophotrochozoan.

Mollusks (Phylum Mollusca)

With over 100,000 living species identified so far, the mollusks must be counted as
among the most successful animals on earth today. Most belong to the first 3 of the 6
classes shown here:
1. Bivalvia. Two shells encase the body. Includes the clams, mussels, oysters, and
scallops.
2. Gastropoda. Snails and slugs. Snails have a single shell ("univalves') while
slugs have none.
3. Cephalopoda. This marine group includes the various species of octopus,
squid, as well as the chambered nautilus. A record 28-foot (8.5 m) octopus and
60-foot (18 m) squid make these the largest of all the invertebrates.
4. Scaphopoda. Marine, filter-feeding "tooth shells".
5. Monoplacophora. Until a live specimen was discovered in 1952, these animals
were thought to have been extinct for millions of years. It has a single shell
(hence the name) and, unlike the other mollusks, is segmented (as are its
relatives the annelids).
6. Polyplacophora. The animals in this group, called chitons, have their dorsal
surface protected by 8 overlapping plates or "valves".
The trochophore larvae of mollusks is also evidence that they belong in the
same clade with the annelids.
Link to drawings of representative mollusks (132K)

Ecdysozoans
All the members of this clade
grow by periodically molting shedding their skin or exoskeleton;
share a unique pattern of HOX genes, e.g. Ubx and Abd-B.
The clade includes a number of phyla of which we shall examine 2:
the nematodes and the

 arthropods.

Roundworms (Phylum Nematoda)

Features:
A one-way digestive tract running from mouth to anus.
A cavity between the digestive tract and the body wall. It develops from
the blastocoel and is called a pseudocoel.
Some 25,000 species have been identified but this may be less than 10% of the
true number.
Most are free-living; found in soil where they are important decomposers.
One of these is Caenorhabditis elegans, a model laboratory animal.
Some are parasitic, including
o hookworms (In 2003 the number of humans infected by hookworms was
estimated at 740 million worldwide.)
o pinworms and whipworms
o filarial worms threadlike worms that are transmitted to the definitive
host from an intermediate host causing such human ailments as
river blindness (Onchocerca volvulus) acquired from the bite
of infected black flies
elephantiasis (Wuchereria bancrofti) acquired from infected
mosquitoes
dracunculiasis (Guinea worm disease) (Dracunculus medinensis)
acquired from ingesting water containing infected "water fleas"
(Cyclops)
o many parasites of commercially important plants like strawberries and
oranges.

 Most are small although one that parasitizes whales reached 30 feet (9 m)!

Arthropods (Phylum Arthropoda)

Some characteristics:
Incredible diversity. Over a million living species have been identified so far
more than all the other species of living things put together and this is
probably only a fraction of them.
Live in every possible habitat: fresh water, salt water, soil, even in the most
forbidding regions of Antarctica and high mountains.
A jointed external skeleton made of chitin, a polymer of N-acetylglucosamine
(NAG).
Segmented.
Pairs of jointed appendages; one pair to a segment used for locomotion,
feeding, sensation, weaponry.
Bilateral symmetry.
Main nerve cord runs along the ventral side.
We shall look at four groups (subphyla):
Crustacea
Hexapoda (the insects)
Myriapoda
Chelicerata
Crustacea
Head and thorax fused into a cephalothorax.
At least 40,000 species.

 Most are aquatic, found

in both fresh water and in
the oceans.
Includes crayfish,
lobsters, barnacles, crabs,
shrimp (drawings at
right).
Hexapoda the insects
Body segments grouped
into head, thorax,
and abdomen.
Each of the 3 thoracic
segments carries a pair of
legs (hence the 6legged "hexapoda")
Many have wings,
usually 2 pairs (only one
pair in flies diptera).
Gas exchange through a tracheal system.
Nitrogenous waste is uric acid thus conserving water.
Some 950,000 species, and this may be only 10% of the number out there.
Dominate all habitats except for the oceans.
Most intensively-studied representative: Drosophila melanogaster.
Representative colonial insect: the honeybee, Apis mellifera
Table listing some of the insect orders.

Myriapoda
Some 13,000 species of

 centipedes and
millipedes
(Neither group has the number of legs their
name suggests, although one species of

Limulus

millipede does have 375 pairs.)

Chelicerata
Anterior segments fused into a cephalothorax.
The first pair of appendages the chelicerae are used for feeding.
There are no antennae.
Includes:
o Merostomata. The only member alive today is Limulus, the horseshoe
"crab". It has existed in the sea virtually unchanged for 200 million
years.
o Arachnids (some 75,000 species)
8-legged
scorpions, mites, ticks, spiders, daddy longlegs [ View].
Evolutionary relationships of the arthropods
An ever-increasing number of arthropod gene sequences appear to have answered
some long-standing questions about the evolutionary relationships of the various

arthropod groups. A recent study (Regier, J. C., et al., Nature, 463:1079, 25 February
2010) examined 63 nuclear genes from 75
species of arthropods and concluded that
the crustacea are paraphyletic; that is,
the single common ancestor from
which all the animals we call
crustaceans are descended was also
the ancestor of another group, the
insects (Hexapoda). So insects are
terrestrial crustaceans!
All these groups plus the millipedes and centipedes (Myriapoda) make up
a clade designated Mandibulata.
So millipedes and centipedes are more closely related to the crustaceans than
to, as once thought, the Chelicerata.

The Deuterostomes
The features of these animals are listed above. The most prominent members of this
group belong to the echinoderms and the chordates.

Echinoderms (Phylum Echinodermata)

Characteristics:
radial symmetry. HOWEVER, their
larvae have bilateral symmetry so the
echinoderms probably evolved from
bilaterally symmetrical ancestors and
properly belong in the Bilateria.
water vascular system. Seawater is
taken into a system of canals and is used
to extend the many tube feet. These have
suckers on their tips and aid the animal in
attaching itself to solid surfaces.
About 6,000 species all of them marine.

There are 5 classes of echinoderms:

Sea urchins and sand dollars (Echinoidea)
Sea lilies (Crinoidea)
Sea Stars (aka "Starfish") (Asteroidea) The photo (courtesy of Dr. Charles
Walcott) shows a sea star that lost an arm and is in the process of regenerating a
replacement.
Brittle stars (Ophiuroidea)
Sea cucumbers (Holothuroidea)
Link to drawings of representative echinoderms and the water vascular system (100K)

Chordates (Phylum Chordata)

During their embryonic development, all chordates pass through a stage called
the pharyngula [View] with these features:
a dorsal, tubular nerve cord ("1")
running from anterior to posterior. At its
anterior end, it becomes enlarged to form
the brain.
a flexible, rodlike notochord ("2") that
runs dorsal to the digestive tract and
provides internal support. In vertebrate
chordates, it is replaced by a vertebral column or backbone long before
maturity.
pairs of gill pouches. These lateral outpocketings of the pharynx are matched
on the exterior by paired grooves. In aquatic chordates, one or more pairs of gill
pouches break through to the exterior grooves, forming gill slits ("3"). These
provide an exit for water taken in through the mouth and passed over the gills.
a tail that extends behind the anus

The vast majority of chordates have a skull enclosing their brain (Craniata), and all
but one of these (the hagfish) convert their notochord into a vertebral column
orbackbone. These latter are the vertebrates.
Vertebrates also differ from all the other animals by having quadrupled their HOX gene
cluster; that is, vertebrates have 4 clusters of HOX genes located on 4 different
chromosomes.
The vertebrates are described in a separate page. Link to it.

Here we shall examine two groups of invertebrate chordates:

Urochordata and
Cephalochordata
Urochordata
This group (also called Tunicata) includes animals
known as ascidians (and commonly called sea
squirts). They are
marine
sessile animals that
feed by filtering food particles from seawater
taken in through one opening, or siphon, and
squirted out the other.
The one on the right is Halocynthia, the sea peach (photo courtesy of Ralph
Buchsbaum).
It is hard to see what makes these animals chordates. The adults have neither
notochord nor a dorsal tubular nervous system.
However, these animals disperse themselves with free-swimming larvae that have
a dorsal tubular nervous system
notochord and

 gill slits
(see the diagram above).
One of the most common species (Ciona intestinalis) has had its genome sequenced.
It has a very small genome: ~1.6 x 10 8 base pairs encoding ~16,000 genes.
(Some 20% of these are organized in operons.)
Link to table giving comparative genome sizes.

Its larva is small (with ~2,600 cells) including only

o 36 muscle cells
o 40 notochord cells
o 100 neurons
These cells (as well as the others) develop along rigid pathways which can be
easily observed because the larva is
transparent.
All these features are shared with C. elegans, but now we are talking about an animal
far closer to the evolutionary line that produced us. In fact, with 80% of Ciona's genes
having homologs in us, tunicates are probably our closest invertebrate relatives.
Cephalochordata
The representative member of this tiny subphylum of so-called lancelets is a small (5
cm), marine, fishlike creature called amphioxus (on the right). (For years its genus
name
was Amphioxus but
that has now been
replaced by the name
Branchiostoma.)
Amphioxus retains:
a dorsal nerve cord

 notochord and
gill slits
throughout its life.
There is a small cluster of neurons at the anterior tip of the nerve cord with certain
similarities of structure and gene expression to the vertebrate fore-, mid- and
hindbrain.
Although able to swim, the lancelet spends most of its time partially buried in the sand
while it filters microscopic food particles from the water.
Welcome&Next Search