BILATERAL ORGANISMS

How did the first bilateral organisms look like?

- Classical view (based on morphological and developmental data): they were
something like a flatworm.
- Today the above view is not at al clear. Most flatworms have been recognized as
secondarily simplified lofotrochozoans. The only flatworms whose phylogenetic
position has not been resolved are the acoelomorphs. In some molecular analyses they
appear as a basal group, but not in all. The position of acoelomorphs is relevant for
reconstructing the characteristics of the common ancestor of all bilaterians. If the
acoelomorphs are not basal, then there is no reason to assume that the first bilaterians
were small and acoelomate (lacking a coelom), but rather large, coelomate and
relatively complex animals, like many protostomes and deuterostomes.

Which are the first fossils that could be attributed without doubt to bilaterians?
- Treptichnus type trace fossils. They appear at the end of the Proterozoic. They are
relatively complex three-dimensional galleries attributed to relatively large bilateral
worms.

PROTOSTOMES:

lophotrochozoa + ecdysozoa form the monophyletic group PROTOSTOMIA. The

division of bilaterians into protostomes and deuterostomes is relatively old (prior to
molecular phylogenies, although both groups have been confirmed by molecular
phylogenies) and based mainly on embryological data. The division of prostostomes in
lophotrochozoa and ecdysozoa is a result of molecular phylogenies and was a big
surprise and a revolution in evolutionary biology.

LOPHOTROCHOZOA: the name of the group refers to the fact that most of its
members have a lophophore (filtering organ) or a trochophore-type larva. An alternative
name for the group (proposed by some scientists) is "Spiralia". This name refers to the
characteristic spiral cleavage of many members of the group (cleavage: an early
embryological process consisting of a series of cell divisions of the fertilized egg that
occur before gastrulation).

MOLLUSCA:

- High diversity and abundance of molluscs in marine and freshwater environments;

also some groups in terrestrial environments.
- Possible phylogenetic relationships among the member of the Mollusca: Aculifera
(Aplacophora + Polyplacophora) + Conchifera [Monoplacophora + (Gastropoda +
Scaphopoda + Bivalvia + Cephalopoda)]. However, the phylogenetic relationships
within the group Mollusca are controversial and have not yet been resolved.
- Although the name molluscs means “soft”, many have a mineralized skeleton.
- They are bilaterians and have a very characteristic body organization based on the
combination of a dorsal shell and very muscular soft parts, with the muscles
generally attached to the shell. The muscular organization of molluscs is very
particular and is based on the presence of three-dimensional nets of muscles. This
type of muscle organization is called a "muscular hydrostat" and allows complex

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 movements without joints. Our tongue or the elephant's trunk also has this type of
muscular organization.

The muscular networks of molluscs are "anchored" to the shell leaving an impression, a
scar. These muscle impressions are practically the only remaining evidence of the
organization of soft tissues in many fossil molluscs.
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MONOPLACOPHORA:

Univalve shell + seriate and symmetrical muscle scars

The class MONOPLACOPHORA was formally proposed in 1952, although the term
had been used informally since the 1940s. The class was created to accommodate
certain fossils that looked like limpets (limpets are gastropods) but that had a very
particular pattern of muscular insertions that indicated they were not gastropods: it was
a pattern that indicated seriation of muscles.
- In the 1950s, the Danish government organized an expedition to study the deep
seafloor, the Galatea Expedition. The expedition collected deep-sea samples from
various parts of the world and took them back to Denmark for further study. In 1956,
when inspecting a sample of the deep seafloor of western Central America collected 4
years earlier (in 1952), the first "living" monoplacophoran was recognized and given the
name Neopilina. A few specimens were collected with an average size of 3 x 3 cm. In
1957 the detailed description of Neopilina galatheae was published.

Today's monoplacophorans are a relatively uniform group (not very diverse) and live in
deep waters. Today's monoplacophorans have a shell similar to that of limpets (which
are gastropods) but the anatomy of their soft parts is quite different.

The morphology of today’s monoplacophorans:

The body has the shape of a "mushroom" (but elongate along the anteroposterior axis).
They have a dorsal calcareous shell, secreted by the dorsal epithelium, which is called
mantle. The shell grows by marginal accretion and the deposition of shell material takes
place mainly at the margin of the mantle. The ventral part of the animal is the muscular
foot. Between the shell and the foot, the body gets thinner forming a “waist. From the
margin of the shell to the "waist", the body forms a concavity, called “mantle cavity”,
where the gills are located.
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It is difficult to establish the beginning of this lineage because of the great diversity of
univalved shells that exist since the early Cambrian. The first monoplacophorans that
we can confirm, because of the presence of muscular impressions are middle Cambrian.
The last fossil monoplacophorans are Devonian. However, monoplacophorans persist to
this day so we have to assume that there is a lapse of many millions of years without
fossil record of this group. The interruption of the record could be due to the fact that
the monoplacophorans, after having lived in shelf waters (shallow waters on the
continental margins) during the early Paleozoic, "sought refuge" in deep waters (where
the paleontological record is not good) and where modern relict species have been
found.

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GASTROPODS:

Characterized by several apomorphies:

- trochospirally-coiled shell
- skeleton with two elements: shell and operculum
- torsion
The skeleton of gastropods consists of a trochospirally coiled shell and an operculum
(which may or may not be mineralized and which has been lost in most land snails).

Gastropods have a thin, elongated, cone-shaped shell which is trochospirally coiled. In

their soft parts there is a "strangulation" that divides them into a dorsal part that contains
the visceral mass and is always inside the shell and a ventral part consisting of the foot
and head. This ventral part is able to be completely outside the shell or hide inside it.
The operculum, located above the foot, closes the opening of the shell once the soft
parts have entered.

The torsion is a twist of the part of the body that is inside the shell with respect to the
ventral part (the foot and head). After torsion, the anus and gills are more or less
resituated above the head. The muscular impressions (the scars left by the muscles
attached to the shell) of the gastropods are asymmetrical, which is an indication of the
torsion of the soft parts. This asymmetry is the only direct indication of the existence of
torsion in fossils.

The first indisputable gastropods date from the upper Cambrian, have a trochospiral
shell and a pair of unequal muscles, which is an indication of torsion. Gastropods have
conquered marine, freshwater and terrestrial environments and have developed very
varied morphologies, e.g. several groups have lost the shell secondarily and have
become "detorted". Their diversity has grown over geological time, and has done so
extraordinarily in the post-paleozoic. Terrestrial gastropods appear in the Carboniferous

BIVALVES
- The vast majority of bivalves are marine, but there are also freshwater species.
- Most are filter feeders, some are detritivores.
- The filtering organs are the gills, which are highly modified and are used both for
respiration and to trap food particles from the water column.
- Bivalves are laterally compressed and have a shell with two valves, one left and one
right, which are specular images of each other. The plane of symmetry coincides with
the plane of the commissure (the plane where the two valves meet).
- The bivalves have lost the cephalization characteristic of molluscs and the foot has
been modified and narrowed laterally (like the whole body) to become a thin tongue.
- Bivalves can be "understood" as a monoplacophoran that has been laterally
compressed and whose shell has been demineralized along the plane of symmetry,
yielding two independent shells.
- Dorsally the shells are connected by the ligament: an elastic "tissue" (which represents
the "demineralized" shell band) that tends to keep the valves open. In addition, the
valves are interlocked by a system of teeth.

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- In the case of bivalves, the most powerful muscles, those that leave the most marked
impressions on the shell, are not the pedal muscles (those directed to the foot) as occurs
in monoplacophorans and gastropods, but the adductor muscles and the pallial muscles.
- adductor muscles: the relaxed condition of the bivalve shell is with the valves open.
The adductor muscles are in charge of closing the valves; there may be two (one
anterior and one posterior) or one that is situated occupying a more central position.
- The pallial muscles: they are retractor muscles of the mantle margin. When the valves
are open the mantle margin can protrude a little beyond the valves and, when the valves
are going to close, the mantle margin has to be retracted quickly. The mantle margin has
sensory functions among others. The pallial muscles are disposed along a fine line
parallel to the margin of the valves, the pallial line.
- Posteriorly, the mantle margin can fuse and be modified to form two tubes called
siphons: one inhalant and one exhalant. Siphons can be very long in species that live
deeply buried in sediment. The corresponding pallial muscles also specialize as siphon
retractors. The presence of siphons modifies the outline of the pallial line, which forms
an embayment (the pallial sinus) to leave room for the siphons to retract. There are
species in which the siphons are very long and cannot be completely retracted into the
shell; therefore the shell does not close completely (this happens "razor clams", for
example). The presence of a pallial sinus in extinct species allows us to infer the
presence of siphons and therefore a deep infaunal habit for the species.
- Bivalves can be regarded as a result of the adaptation of molluscs to a filter feeding
and relatively sedentary mode of life.
The paleontological record of bivalves is generally good. However there is an enigma
that raises doubts about the origin of bivalves. The enigma is that there are a few species
of what appear to be tiny bivalves in the early Cambrian and early middle Cambrian,
but then there is no record until the early Ordovician, period in which the bivalves
experience their first important radiation. This leads us to question whether the first
small bivalved fossils are in fact the ancestors of the Ordovician bivalves.
The vast majority of Paleozoic bivalves lived in soft sediments, generally buried
superficially or only partially buried. They lived in shallow waters. However, the
dominant filter feeders in this environment were the brachiopods, most of them
epifaunal organisms (they lived above the substrate). Brachiopods were decimated in
the extinction at the end of the Paleozoic, while the bivalves were not that much
affected. Bivalves experienced an important radiation during the Mesozoic. It is thought
that the expansion of epifaunal bivalves during the Mesozoic was related to the decline
of brachiopods, while the expansion of infaunal bivalves was related to the origin and
development of siphons. Siphons allow bivalves to bury themselves deeply. This
allowed them to burry themselves deeper in those environments already inhabited by
them, and also to occupy the intertidal zone, which they probably did not occupy during
the Paleozoic.
The first freshwater bivalves appeared in the Devonian. Their shells are characterized
by a very thick periostracum that protects them from dissolution in an environment not
saturated with calcium carbonate.
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CEPHALOPODS:
Most modern cephalopods have a derived (apomorphic) morphology, which is quite
different from that of other molluscs.
They are exclusively marine. Modern cephalopods are generally active predators and as
such have a well-developed nervous system and a marked cephalization. Many have a

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high metabolic rate compared to other invertebrates and some are good swimmers,
capable to compete with fish. Eyes are usually highly developed; in particular, the eyes
of squids are very similar to those of vertebrates and are a remarkable example of
convergent evolution. The largest invertebrates can be found among the cephalopods.

Today it is a relatively diverse group (400 modern species) that includes many species
with internal shell or without shell, and only a few with external shell. In the past there
was a high diversity among the species with external shell, this being the plesiomorphic
condition for the group.

Modern cephalopods can be divided in two groups:

1- Nautiloids → today there are only a few species. The best-known species is Nautilus
pompilius, exclusive to the South Pacific, where it lives around the coral reefs.
Nautiloids retain plesiomorphic features, particularly the possession of a chambered,
external shell.

2- Coleoids→ shell internal or absent. Derived group: cuttlefish, squid and octopus.
Characterized by the internalization of the shell, an evolutionary process that is difficult
to conceive (how can an external shell "get inside" the body?). We can visualize what
happened in general morphological terms if we understand that it was the edge of the
mantle (the epithelium that surrounds the margin of the shell and makes it grow) that
"overflowed" and bent backwards to engulf the entire shell.
Cuttlefish retain a relatively robust skeleton. The cuttlefish shell is homologous to the
skeleton of Nautilus and, although very modified, the same basic structure can still be
recognized. Squids also have a homologous internal "shell", but it is not mineralized.
Octopuses do not have shells. Cuttlefish still use their shells to control their buoyancy
like the cephalopods with external shells, squids do not.

Nautiloids: the example of Nautilus pompilius:

Shell:
Is a thin-walled planispirally coiled cone (this is the type of coiling characteristic of
cephalopods; although there existed a few species with irregular or trochospiral coiling,
this is unusual among cephalopods; on the other hand, species with straight shells were
common, especially in the lower and middle Paleozoic).
The soft parts of the animal occupy the most recent part of the shell, the part close to the
opening, which is called body chamber. As the animal grows, the soft parts move
forward, progressively "abandoning" the oldest part of the shell. The "abandoned" part
of the shell is the phragmocone and is chambered. The septa are formed successively,
one in each episode of displacement of the soft parts. In each displacement, the soft
parts move forward towards the opening, allowing water to enter into the shell. Then the
back of the soft parts secrete a thin wall, the septum, which seals the abandoned part of
the shell. Septa are hemispheric walls of calcium carbonate, perforated by a single
circular hole (the septal foramen). This perforation is usually funnel-shaped, with a neck
(the septal neck). The space between two adjacent septa is a chamber and is initially
filled with water (which the soft parts have let inside the shell when moving forward).

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The soft parts of Nautilus include a fleshy cord (a kind of "umbilical cord") located
posteriorly. This fleshy cord is called siphuncle and contains blood vessels. The
siphuncle is the only part of Nautilus that does not abandon the old chambers, but
extends from the oldest part of the shell to the body chamber. Therefore, when the
animal builds the septa, it leaves a hole for the siphuncle. The siphuncle gradually
extracts the water from the abandoned chambers and fills them with air.

The most recently formed chamber is filled with liquid, and the immediately preceding
ones partially so, but the older chambers are instead filled with a nitrogen-rich gaseous
mixture. Little by little Nautilus replaces the liquid in the chambers with gas by means
of the siphuncle. Using an osmosis-based mechanism, the liquid from the chambers
passes into the blood, and the gas diffuses from the blood into the chambers. Thus
Nautilus regulates its buoyancy, adding more gas to the abandoned chambers as its body
grows and its weight increases. The result is that the animal achieves neutral buoyancy
by equating the total density of its body (soft parts plus partially full of gas shell) to that
of seawater.
Once Nautilus reaches maturity it stops adding chambers to its shell.
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NAUTILOIDS
A paraphyletic and morphologically diverse group. Includes the first cephalopods,
which appear at the end of the Cambrian. They are abundant and diverse in the
Paleozoic, particularly during the Ordovician. Many early Paleozoic nautiloids had
straight shells (generally known as orthocones) and some reached up to 9 m in length
(the largest shells ever secreted by invertebrates). They almost became extinct at the end
of the Triassic (the straight-shelled nautiloids disappeared and only coiled forms
remained), but still survive today represented by a few species.

AMMONOIDS
Their soft parts are not known in detail but are assumed to be similar to those of
nautiloids. Ammonoids originated from nautiloids. The lineage of ammonoids separated
from the nautiloids in the Devonian and diversified greatly during the Mesozoic. They
became extinct at the end of the Cretaceous, like the dinosaur. There is evidence that at
least some ammonoids fed on plankton.

COLEOIDS
Today they are a diverse group. They also originated from nautiloids. Coleoids are a
fundamentally Mesozoic group and, although their origins are not well known, probably
originated at the end of the Paleozoic.

Belemnites were an important coleoid group during the Mesozoic and became extinct in
the same extinction that ended with ammonites and dinosaurs.
They were similar to today's squids but had a more robust skeleton than any of today's
coleoids.
The skeleton of belemnites consists of:
- phragmocone → homologous with the phragmocone of nautiloids
- pro-ostracum → homologous with the body chamber
- guard or rostrum → has no homologous structure in cephalopods with external shell. It
is a massive deposit of calcium carbonate that was deposited, layer by layer, at the end
of the phragmocone. It is believed to have served as a counterweight to the anterior
portion of the body, to help the animal maintain a horizontal position.

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--------------------------
POLYPLACOPHORANS:
-Multiple skeletal elements, seriate muscle scars
-The mode of life and adaptations of polyplacophorans are similar to those of limpets
and monoplacophorans. Most live on rocky shores, in sub- or intertidal energetic
environments.
- Dorsoventrally flattened and with a wide muscular foot. Dorsally they have a row of
eight articulated plates, and in the margins can have spicules.
- The geological record of polyplacophorans is not very good, perhaps partly because
the shells are easily disarticulated and are fragile and aragonitic. Also, they tend to
inhabit energetic environments that are not favourable to fossilization.
- The first polyplacophorans date from the upper Cambrian.

SCAPHOPODS
A very specialized group. They are infaunal and micropredators (often prey on
foraminifera). Shell and mantle are ventrally fused forming a tube open in both the front
and back. It is an enigmatic group and little is known about their evolution.
Probably the first scaphopods are Devonian. Possible specimens in the Ordovician are
uncertain.
Scaphopods are not common in the fossil record, except for sporadic concentrations in
Cenozoic sediments.

BRACHIOPODS

Brachiopods are exclusively marine filter feeders. Today they are relatively common
animals (about 300 living species), however they are not familiar animals. In general
they are numerically unimportant and do not usually occupy coastal environments,
which are the marine environments most familiar to us. Most species live in the Pacific
Ocean. They are usually found in "marginal" environments: deep water, cold latitudes
and environments poor in oxygen or brackish. They were much more abundant and
diverse in the past, and occupied platform and coastal environments.

Traits shared by all brachiopods:

1- External skeleton with two unequal valves. The valves grow by marginal accretion,
like those of molluscs.
2- They are filter feeders: the filtering organ is the lophophore.
3- They have setae along the mantle margin.
4- Unlike molluscs, they are not very muscular animals. Most of their body mass, the
viscera and muscles, occupies only a small space "at the back" of the shell and most of
the shell is actually occupied by the lophophore and seawater. The main muscles are
those that articulate the valves. There are no muscles comparable to the massive pedal
muscles of molluscs, responsible for the movements of the foot.
5- They have a pedicle.
6- Their plane of symmetry is perpendicular to the plane of the commissure and cuts
each valve in half.
7. Brachiopods are always sessile and benthic.

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The different species of brachiopods have different life positions, but the vast majority
throughout the history of the group have been epifaunal (epibenthic). This is an
important difference with bivalves, which have been mostly infaunal (endobenthic)
organisms. Many living and extinct species live attached to the substrate by the
peduncle; others are reclined on one of their valves on the soft substrate. Some species
are cemented to a hard substrate and others (few) live buried in the sediment.

RHYNCHONELLIFORM BRACHIOPODS (ARTICULATED):

- Strongly mineralized calcareous shell.

- Valves:
pedicle valve → usually larger, associated with the pedicle. Often have a foramen or
groove through which the pedicle comes out.
brachial valve → usually smaller, associated with the lophophore. Sometimes has a
mineralized internal structure that serves as a support for the lophophore.

- Articulation: valves are directly articulated through a system of teeth and sockets. The
opening and closing of the valves depends on diductor and adductor muscles
respectively.

LINGULIFORM BRACHIOPODS (INARTICULATES)

- organophosphatic shell
- do not have direct articulation of the valves. They open the shell by hydrostatic
pressure in its soft parts. Have powerful adductor muscles to close the valves.

Some species are infaunal (endobenthic), burrowers, and have muscles that allow them
to rotate the valves one with respect to the other to burrow. These species have a long
pedicle.

CRANIIFORM BRACHIOPODS (INARTICULATES)

- calcareous shell
- cement themselves to the substrate with the pedicle valve→ do not have a pedicle.

PALEONTOLOGICAL RECORD
The first brachiopods appear in the lower Cambrian and are linguliforms. Soon
rhynchonelliforms appear as well. From the Ordovician onwards rhynchonelliforms
undergo a great diversification. Brachiopod shells are among the most common fossils
in the Middle Paleozoic.
The group was decimated after the extinction of the end of the Paleozoic and never
recovered, although some rhynchonelliform groups remained important in the
Mesozoic.

BRYOZOA:
Aquatic metazoans (mostly marine, some in freshwater), exclusively colonial, sessile
and suspension feeders (the zooids have a lophophore). Zooids are small, and colonies
are usually also small. The colony is a clone. Zooids originate by asexual reproduction
from a first zooid and remain connected throughout life. Sometimes there is a marked

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division of labour between zooids in the same colony (zooids specialise in reproduction,
defence, cleansing).
Many bryozoans are encrusting and have the ability to adhere to hard surfaces and
algae. There are also non-encrusting bryozoans whose colonies can be lamellar, massive
or branched.
Bryozoans build external skeletons that can be organic or mineralized with calcium
carbonate.

They appear in Ordovician and reach to the present day. They are diverse and important
during the Paleozoic and are an important members of Paleozoic reef communities,
along with stromatoporoids and corals. They are decimated in the Permian extinction
and many groups of bryozoans become extinct. There are very few species in the
Triassic but during the Jurassic they experience a new expansion. They are abundant in
today's seas.

ANNELIDS
The name means "small rings" and refers to their segmentation. They are soft-bodied
animals with a hydrostatic skeleton, so their preservation potential in most sedimentary
environments is low. In spite of having a soft body, the fact that they possess a cuticle
and setae facilitates their preservation in exceptional deposits such as Burgess Shale.
The first fossil annelids are Cambrian, and very similar to recent polychaetes.
In some polychaetes the mouth leads to an evaginable pharynx provided with strongly
sclerotized "mandibles". These masticatory structures are often preserved and are called
scolecodonts. Scolecodonts appear in the Ordovician and their stratigraphic range
stretches to the present. It is assumed that "mandibulate" annelids originated in the
Ordovician.