GEO 405: Life through Time (Spring 2021)

Lab 7: Survey of Invertebrates

The Digital Atlas of Ancient Life has

great resources! Check them out!
www.digitalatlasofancientlife.org/vc

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Animalia

Porifera
Demospongea (C. - Rec.)
Hexactinellida (Hyalospongea) (C. - Rec.)
Calcispongea (C. - Rec.)

Coelenterata (also known as Cnidaria)

Hydrozoa (C. – Rec.)
Scyphozoa (C. – Rec.)
Anthozoa
Rugosa (Ord. - Per.) Extinct form
Tabulata (Ord. - Per.) Extinct form
Scleractinia (Tri. - Rec.) Modern form

Annelida

Bryozoa
Stenolaemata (Ord. - Rec.)
Trepostomata (Ord. - Tri.)
Cryptostomata (Ord.- Per.)
Gymnolaemata
Cheilostomata (Tri. - Rec.)

Brachiopoda
Inarticulata (C. - Rec.)
Articulata (C. - Rec.)
Strophomenida (Ord. - Jur.)
Rhynchonellida (Ord. - Rec.)
Spiriferida (Ord.- Jur.)
Terebratulida (Dev. - Rec.)

Arthropoda
Trilobita (C. - Per.)
Chelicerata
Merostomata
Eurypterida (mid-Ord.- Per.)
Xiphosura (C.- Rec.)
Uniramia
Myriapoda (Sil.- Rec.)
Insecta
Crustacea (C.- Rec.)
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Mollusca
Gastropoda (snails, slugs, limpets, etc.) (C – Rec.)
Bivalvia (Pelecypoda) (clams, oysters, scallops, etc.) (C – Rec.)
Cephalopoda
Nautiloidea (chambered nautilus) (Late C – Rec.)
Ammonoidea (ammonoids; extinct) (Silurian – Cretaceous)
Coleoidea (belemnoids, squids, octopus, cuttlefish) (Dev. – Rec)
(Plus many other classes uncommon or rare as fossils)

Echinodermata
Echinozoa
Echinoidea (Ord.- Rec.)
Crinozoa
Crinoidea (mid- C.- Rec.)
Blastozoa
Blastoidea (Sil.- Per.)
Asterozoa
Stelleroidea (Ord.- Rec.)

Hemichordata
Graptolithina (C. - Miss.)

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PORIFERA

Poriferans are known to most of us as sponges, and they are among the simplest of
multicellular animals. The basic body plan is a two-layered sack perforated by numerous pores.
The pores lead into a central cavity, the paragaster. The channels from the pore mouths to the
paragaster are lined with special, flagellum-bearing cells called choanocytes. The flagella beat
continuously creating a current that pulls water, and whatever food particles are in the water,
through the pores. Special amoeboid cells trap food particles and distribute them throughout the
other cells of the sponge. Water is then expelled through the opening at the top, called the
osculum.
Today sponges are mostly marine, though freshwater forms do exist. They can be quite
abundant and have probably been around since before the Cambrian (some potential sponge
sterols have been found in rocks that are 2.7Ga!). The fossil record of sponges is spotty because
of the construction of their skeletons; sponges can be either calcareous (skeletons made of
carbonate) or siliceous (skeletons made of silica/glass spicules).
Calcareous sponges, like stromatoporoids, have skeletons made of carbonate, these have
a good fossil record and formed some of the earliest reefs.
Siliceous sponge skeletons are composed of thousands of tiny, interlocking structures called
spicules. Spicules have one or more rays and the number of rays can aid in identifying the order
of sponge present. Because they are not cemented together, when a sponge dies the spicules tend
to come apart relatively quickly. As a result, intact skeletons are rare but isolated spicules can be
common in rocks. Another problem with the fossil record of sponges is that many demosponges
have spicules made of spongin, an organic protein you commonly see in the form of bath and art
sponges. Spongin decomposes and does not preserve. Because sponges have a relatively poor
fossil record and have not changed much over the course of their history, they are not very useful
for biostratigraphy.

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CNIDARIA (hydras, jellyfish, corals)

Anthozoa (corals)
Several Anthozoan coral groups produce
carbonate skeletons. At its simplest the skeleton is a thin
circular disk with radiating ridges (septa) secreted by
the coral. In most forms, however, the young coral adds
to the disc, forming a cup (calyx) in which it rests. Septa
are usually present at this stage, too, and form vertical
calcite walls radiating away from the center of the calyx.
The small coral and its skeleton continue to grow,
expanding in diameter and height. The animal remains
in the upper calyx portion and soon requires skeletal
supports to prevent its base from sagging. These skeletal supports may be either flat plates called
tabular or curved "blister-like" dissepiments. These are the principle taxonomic characteristics of
a coral skeleton. Not all corals build the complete skeleton, however, by recognizing which
skeletal parts are present or absent different coral groups can be identified.

Rugose Corals
Rugose corals have well developed septa distinguishing them from the tabulate corals,
which do not. Well developed tabulae and dissepiments are also present, but the fossils usually
must be broken to see these. Both solitary and colonial forms (multiple corallites in one solid
colony) are common. Solitary Rugosa (horn corals) are shaped like a horn. Colonial Rugosa
range from isolated tubes connected to each other by lateral tubes, to compact honeycomb-like
masses. Rugose corals commonly range from the Ordovician to Permian, and were extremely
diverse and abundant in that time. They preferred warm, clear, shallow water and many were
important reef builders. Many are also important index fossils.

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Tabulate Corals
Tabulate corals have well developed tabulae but no (or very small) septa, or
dissepiments. All are colonial. Four distinct skeletal types exist, represented by the following
genera.
Syringopora (organ pipe coral) has erect, rough, irregular tubes with funnel-shaped
tabulae (rarely seen). The individual tubes are connected by horizontal tubes.
Favosites (common name: honeycomb coral) is shaped like a compact honeycomb.
Colony may be flat or mound shaped. In cross section numerous well-developed horizontal
tabulae are readily seen.
Halysites (common name: chain corals) appear, on top, as small chains. The "chains" are
in fact vertical tubes that join and diverge in various ways leaving the areas between the chains
of tubes open. Well-developed horizontal tabulae are present but not often seen.
Aulopora are small (< 1 cm) trumpet shaped corals each growing outward from
somewhere along the side of the previous. Curved tabulae are present but usually not seen. This
coral encrusts shells and other foreign surfaces.
Tabulate corals commonly range from the Ordovician to the Permian, but are especially
abundant in the Silurian and Devonian. Some are important in reef formation, these were
restricted to warm, clear water, and some are useful as index fossils.

Scleractinian Corals (Mid-Triassic)

Like the Rugosa, Scleractinian corals always possess well-developed septae and at times
dissepiments and a columella, but never tabulae. Both colonial and solitary forms exist. Because
of their similarity Scleractinian and Rugose corals may be hard to separate, but they are
distinguished, technically, by two taxonomic characters. First, Rugosa add septa in multiples of
four, Scleractinians in multiples of six. Careful counting under a microscope may distinguish
between them on this basis (although not always well in more complex forms). Second, in the
Rugosa septae may sometimes be undeveloped leaving a gap, a fossula. In some Scleractinians
distinct calyces are not present; instead rows of septa meander across the coral surface. Finally,
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some Scleractinian fossils are original hard parts with a white, bleached look, unlike Rugose
corals that look like rock.
Scleractinian corals first appeared in the mid Triassic, and have rapidly expanded to

become abundant and diverse today. Because rugose corals became extinct at the end of the
Permian these two groups are never found in the rock record together.

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8
 (LOPHOPHORATES) (Brachiopods, Bryozoans, Phoronid worms)

BRACHIOPODA

Brachiopods are benthic (bottom dwelling) marine animals with an external, two-valved
shell. They look superficially like bivalve molluscs, but differ in important ways. First, the shells
of brachiopods are different sizes and are symmetrical along a median plane. Bivalve shells are
symmetrical in a plane passing through the hinge line (see below). Second, most of the space
inside a brachiopod shell is taken up by its ciliated, mucus-covered feeding structure, the
lophophore. The beating action of the cilia creates a current that pulls water and food particles
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into the shell. Food is trapped by the mucus and cilia, and carried by ciliary action to the mouth
and intestine at the back of the shell, near the hinge line. They lack any specialized nervous or
sensory system. The valves are opened and closed by a set of diductor and adductor muscles
respectively. Some brachiopods live in burrows, while others rest in bottom muds or cement
themselves to solid bottom surfaces or objects. Many types anchor themselves with a fleshy stalk
called a pedicle. The pedicle projects from an opening, the pedicle foramen, through the pedicle
valve. The other valve is known as the brachial valve. Brachiopods have no effective means of
locomotion, other than in some forms that can push themselves around with their pedicle. The
shell can be turned in any direction on the pedicle by the use of adjuster muscles. This way they
can angle themselves into or away from the water current. Today brachiopods are not as diverse

as they were in the past, and they live in cool to temperate marine waters. They are most
common in the western Pacific. Most modern forms live in shallow water (<200m), although a
few are found at depths of 5000m.

Inarticulata
The valves of inarticulate brachiopods are usually chitinophosphatic or sometimes
calcareous in composition. The hinge articulation lacks teeth and sockets. The valves are oval or
circular in outline, are gently convex, and look quite similar. Recent specimens are often black
and shiny as a result of the phosphatic composition. Some ancient fossil specimens retain this
shiny black or dark appearance. Faint growth lines are sometimes visible, but other
ornamentation on the shell is very rare. Inarticulates range from the lower Cambrian to the
Recent, but are rare as fossils. The well-known genus Lingula ranges from the Cambrian to the
present and is frequently found in the fossil record.

Articulata
Articulate brachiopod valves are calcareous and commonly remain articulated after death.
Most are 2-3 cm in length, but sizes range from a few millimeters to tens of centimeters. Many of
the skeletal structures used in classification are difficult to see. However, many external features
make identification relatively easy. Many forms have a fold and sulcus, and the relative size and
shape of the hingeline is also diagnostic. Some skeletons are globular in form, others are disc-
shaped. The valves may be biconvex, plano-convex, or concavo-convex:
Articulate brachiopods are extremely abundant in the fossil record, especially in the
lower and middle Paleozoic. Approximately 30,000 fossil species are known. Since the
Paleozoic, they have declined to about 200 living species. Nearly all the extinct forms lived in
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shallow water, and some were associated with reef communities. Because of their abundance and
diversity, they are good tools for Paleozoic biostratigraphy.
BRYOZOA

At first glance many people mistake bryozoans for small colonial corals. However,
bryozoans are more advanced in their body structure than any coelenterate, and are actually more
closely related to brachiopods. The small size of individual living chambers and the lack of
septae are good characters for telling them apart from colonial corals.
Individual zooids (the name given to each bryozoan animal) are tiny, less than 1 mm
across. They form colonies that may be stick or twig-like, massive, delicate fans, or encrusting.
The colonial skeletons are made of either chitin, a tough organic substance, or by calcite.
Individuals live in tiny box or tube-shaped
chambers called zooecia (sing. zooecium).
The opening into an individual zooecium is
called an aperture. They feed by the use of a
lophophore, a set of ciliated tentacles that
filter water and trap passing microscopic
organisms. The mouth is centered at the base
of the tentacles. The body structure is a
simple, with a “U”-shaped gut and no well
developed sense organs. When resting or
hiding from danger the individual pulls itself
into its living chamber and blocks off the
aperture with a tough operculum.
Fossil bryozoans are exclusively calcareous, because chitin does not usually stand up
well to fossilization. Most modern forms are chitinous. Fossils are very common and make good
index fossils. Some Paleozoic carbonates (especially Mississippian aged) are made up in large
part by bryozoan remains. They first appear in the fossil record during the Ordovician. Massive
and thick, stick-like forms dominated during the Silurian and Devonian. From the Mississippian
to the Permian the more delicate twig-like and fan forms were the most abundant. Bryozoans
declined in diversity at the Triassic-Permian boundary, but rebounded in the Jurassic and persist
to the present day. They were important reef-builders in the Paleozoic, with tightly packed
clumps of bryozoans often making “mud mounds”. These were formed when the bryozoans
acted as baffles to trap fine sediment, eventually accumulating large amounts in mounds.

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ECHINODERMATA (sea stars, urchins, crinoids, blastoids)

Echinoderms are exclusively marine invertebrates. They are deuterostomes, an

embryological indication that they are actually more closely related to chordates than to the other
invertebrate groups you have seen so far. They all have an internal skeleton (endoskeleton)
composed of large numbers of porous calcite plates. A layer of spiny skin covers the animal
externally (hence the name of the group). Skeletons usually have pentameral (5-sided)
symmetry although some forms have a bilateral body plan superimposed on the pentameral plan.
A feature unique to echnioderms is their water vascular system; this is a water filled system of
complex canals and sacs used for locomotion, respiration, and acquiring food. In more advanced
forms, water enters the system through a sieve-like opening, the madreporite. Tube feet,
extensions of the water vascular system, emerge through pores in areas called ambulacra (sing.
ambulacrum). Different groups of echinoderms have radically different body forms and
lifestyles, reflecting their diversity and long evolutionary history. Some are active predators
while others are detritus feeders. A few sessile forms are known.

Crinoidea (sea lilies, feather stars)

Crinoids have a disc-shaped or globular calyx (body) and a variable number of arms,
which often are much larger than the calyx. A long stem, the column, extends downward from
the base of the calyx and attaches to the substrate with a holdfast. The column is made up of
numerous columnals, small calcite plates with a central hole stacked one atop the other. Isolated
columnals are common fossils. Because the entire skeleton is made up of small plates, complete
fossil skeletons are rare. Some crinoids, including most living forms, lack a column and can
move about freely by crawling on their numerous arms. Crinoids range from the middle
Cambrian to the Recent, but were most diverse and abundant from the Silurian to the
Mississippian.

Blastoidea (blastoids)
Do not confuse blastoids and crinoids, for they do look superficially similar. Blastoids
also attached themselves to the substrate with a long stem made of columnals. The calyx is a
compact cup made up of 13 calcite plates, arranged in 3 highly symmetrical rings, with five
ambulacral areas in between them. The ambulacra are distinctive for their numerous, regularly
spaced transverse grooves. The mouth is located at the center of the top of the calyx, and is
surrounded by five smaller openings, including the anus. Fossil calyces are almost always found
whole, and isolated stems are effectively indistinguishable from those of crinoids. Blastoids
appeared in the Silurian and reached their greatest abundance during the Mississippian. By the
end of the Permian they were extinct.
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Echinoidea (sea urchins, sea biscuits, sea hearts, and sand dollars)
These are disc, heart, or biscuit-shaped creatures. They lack arms or appendages.
Regular echinoids (sea urchins) are pentamerally symmetrical, with the anus centrally located
on top of the animal. Irregular echinoids (sand dollars and sea biscuits) are bilaterally
symmetrical and have the anus located underneath or laterally on top of the skeleton. The outer
surface of the skeleton is covered with movable spines. Spines don’t always make it into the
fossil record, as they are easily scattered after the animal’s death. The spines attach to small,
rounded knobs (tubercles) on the plates making up the skeleton. They first appear in the
Ordovician but are uncommon in Paleozoic rocks. Mesozoic and Cenozoic echinoids are
common fossils, and some are useful in biostratigraphy.

Asterozoa (sea stars, brittle

stars)
The sea stars and brittle
stars are mobile echinoderms with
five radiating arms connected to a
central disc. The mouth is located
in the center of the disc’s
underside. The anus and
madreporite are found on the upper
surface of the disc. The ambulacra
are strongly developed and one is
located on the underside of each
arm. Large tube feet extend
through them, facilitating the
mobility of these creatures. Many
sea stars and a few brittle stars are
deadly efficient predators on other
marine life, able to pry open
bivalve mollusc shells, and
generally consume anything not
fast enough to keep clear of them.
Other stelleroids are suspension or
detritus feeders. The skeleton is
made up of many small, very
loosely attached plates, so when
stelleroids die, the skeletons are
quickly broken down and
scattered. Fossils are rare.

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ARTHROPODA

Arthropoda is an unusually large and diverse group of complex invertebrates

(e.g.trilobites, eurypterids, horseshoe crabs, arachnids, crustaceans, insects). The basic body plan
consists of a variable number of body segments and jointed appendages, encased within a tough,
chitinous exoskeleton. Primitively, each body segment has a single pair of legs, but these are
often reduced or modified for feeding or sensory capabilities. There is usually a distinct head and
abdominal region.
Arthropods have been successful in all environments. There are estimates that put the
number of identified arthropod species at over one million. Some species are extremely abundant
(e.g. flies and ants) while others are known only from one or two fossils. Many arthropod
skeletons become mineralized and wind up as important components in the fossil record.
Arthropods grow by shedding their exoskeletons, so those chitinous body fossils are numerous.

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Trilobita
The trilobite body is divided into three
sections from anterior to posterior: the
cephalon (head), thorax (body), and
pygidium (tail). It is also divided
longitudinally into a central axial lobe and
two pleural lobes. The axial lobe includes the
glabella on the cephalon. The glabella can be
bulbous and enlarged, or quite reduced; but
under the glabella sat the stomach of the
animal. The thorax is comprised of individual
jointed segments. There may be as few as two,
though the average is twelve. The pygidium is
triangular to semi-circular in shape. Its
segments are fused into a solid structure, but
grooves show where individual segments once
were.
Trilobites are commonly found whole,
and even broken fragments are easily identifiable. Some are found enrolled, a defensive
maneuver where the animal curled up and the cephalon and pygidium are in contact. The
appendages are rarely preserved.
Trilobites first appear early in the Cambrian and extend through the Paleozoic,
disappearing in the Permian. More than 1500 genera are identified, and many make excellent
index fossils for biostratigraphic purposes. They were widespread in marine environments and
most played the roles of herbivore and scavenger. They moved about by crawling along the
bottom, or by swimming.

Crustacea
Most crustaceans are free-living aquatic animals, but some are terrestrial, some are
parasitic, and some are sessile (e.g. barnacles). The group has an extensive fossil record,
reaching back to the Cambrian. The body of a crustacean is composed of body segments, which
are grouped into three regions: the cephalon or head, the thorax, and the pleon or abdomen. The
head and thorax may be fused together to form a cephalothorax, which may be covered by a
single large carapace. The crustacean body is protected by the hard exoskeleton, which must be
molted for the animal to grow.
Each somite, or body segment can bear a pair of appendages: on the segments of the
head, these include two pairs of antennae, the mandibles and maxillae; the thoracic segments
bear legs, which may be specialized as pereiopods (walking legs) and maxillipeds (feeding legs).
The abdomen (think about ordering “lobster tail”) bears pleopods, and ends in a telson, which
bears the anus, and is often flanked by uropods to form a tail fan. The number and variety of
appendages in different crustaceans may be partly responsible for the group's success.

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16
MOLLUSCA

Molluscs (or Mollusks) are one of the most successful of the invertebrate phyla, both in
diversity and in abundance. Their adaptations are so diverse, however, that few generalizations
can be made about them. What joins all these superficially different appearing animals is a basic
body plan. Most are soft, fleshy, unsegmented organisms with bilateral symmetry. Molluscs
have a distinct head and tail region and within the head are sensory structures and a ribbon-like
row of teeth called a radula. In most groups the posterior or ventral part of the body is modified
into a “foot” used for crawling, burrowing, and other locomotion. Nearly all molluscs are
covered by a fleshy mantle, which secretes a calcareous shell in many species.
Since their origin in the Cambrian the molluscs as a group have evolved into every
aquatic habitat. Some are floaters, some swimmers, and others live on the bottom either attached
in a burrow, or crawling about. The gastropods as well have been one of the very few animal
groups to successfully invade the terrestrial environments (arthropods and vertebrates are two
others). Feeding styles range from vegetarians to scavengers, to the most predacious of the
invertebrates. In addition, molluscs include very tiny members (snails less than 1 mm long) and
the largest of the invertebrates (the giant squid at 15 meters long). Squids and octopi are also
believed to be the most intelligent invertebrates. The vast fossil record of the molluscs indicates
they were as successful in the past as today.

Gastropoda
Gastropod soft anatomy consists of a well-developed head with sense organs, a foot for
crawling, and a mantle that secretes the shell, although many forms like the common garden slug
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have lost the shell. The most unique biological feature occurs during embryological development
when the body is torsioned so that the gills and anus lie on top of the head facing forward rather
than rearward. The torsioning of the soft anatomy should not be confused with shell coiling
which is an independent process.
The basic gastropod skeleton is a hollow, tapering calcite tube, which is coiled around a
central axis, like a pipe coiled down the length of a stick. Beyond this generalization the
gastropod shell comes in enormous variety from tall spires to squat forms. Some are relatively
plain, and others highly decorated. Despite this great variability most gastropod shells are easily
recognized by their distinctive coiling. There is little basic difference between fossil and recent
shells.
Gastropods are the most adaptable of the mollusks. They include vegetarians,
scavengers, and fierce predators. Ecologically they include marine, freshwater, and terrestrial
forms. And within each of these environments they have evolved into nearly every
subenvironment. At the present time gastropods exceed all other mollusks in variety of shell
form and numbers. There are 35,000 living species and many extinct groups.
Gastropods have the oldest fossil record among the living molluscs. They first appear in
the early Cambrian and are common fossils throughout geologic history. Despite their success
and abundance the paleoecology of fossil forms is difficult to interpret. Therefore, gastropods are
not valuable for correlation.

Bivalvia/Pelecypoda (clams, oysters, mussels, scallops)

Pelecypods are molluscs with two-valved shells that surround and protect the body. DO
NOT CONFUSE THEM WITH BRACHIOPODS! With a few exceptions, bivalve valves are
equal-sized and not symmetrical along the median plane. The valves are hinged at the top with
teeth and sockets, and the shells are held open by a specialized ligament. The shells can only be
held closed by the exertion of the adductor muscles. As a result, bivalve shells spring open upon
death, and are then often separated.
A flattened, muscular foot is used to creep along bottoms or burrow deeply into sediment.
A few clams are able to bore directly into hard substances such as rock, or wood. Oysters cement
their shells to hard substrates and never move again, while scallops “swim” by jetting water as
they rapidly flutter their valves. Some bivalves (e.g. scallops and mussels) attach themselves to
the substrate by secreted byssal threads.
On the internal surface of valves, adductor muscle scars are commonly preserved, even in
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fossils. Scallops and oysters have only a single, large scar. A pallial line around the edge of the
shell indicates the site of attachment for the mantle. A pallial sinus, indicated by a sharp
inturning of the pallial line, is present in forms that had large siphons. The siphons are retracted
into this pallial sinus area when threatened.
Most pelecypods are filter feeders (a few deposit feeders exist). Water and food particles
are brought into the bivalve through an incurrent siphon. Food particles are trapped in the gill
filaments and carried to the mouth by cilia-action. Oxygen exchange takes place in the gills and
the filtered waste-water is ejected through an excurrent siphon. Shells are variable and relatively
good indicators of the lifestyle of the species. Razor clams are elongate, smooth-shelled, and
thin, a good design for rapid movement through fine sediments. Strongly ridged or rough shelled
forms are able to resist the pounding of a high-energy environment, such as wave zones or rocky
shorelines. Some oysters are well designed for sitting on top of bottom mud (e.g. Gyphaea) while
others cement themselves onto hard substrates.
Scallops "swim" by rapidly fluttering their shells creating jets of water by which they can
propel themselves quite effectively. These shells are distinguished by: (1) non-mirror image
shells- one is convex the other almost flat, (2) nearly symmetrical anterior and posterior halves of
each shell, (3) one large internal muscle scar situated almost centrally in the shell, (4) well
developed plica radiating out from the hinge line center.
Oysters and relatives have become specially adapted for resting on soft bottom muds or
being cemented to hard substrates. Forms like Gryphaea and Exogyra lived in soft bottom muds
with a large, boat shaped shell. The second shell formed a much smaller protective cap.
Clams appear in the lower Ordovician and were common by the Silurian and Devonian.
Fresh-water forms are known by the Devonian. After the Paleozoic clams surpassed the
brachiopods in number and diversity. During the Mesozoic clams became increasingly abundant
and important. Today they are the second largest group of mollusks.

Rudists

Some of the most strangely aberrant kinds of bivalves known are the Rudists
(Rudistacea). In this group, which occurred during the Cretaceous, the lower valve takes the
form of an elongate thick-walled cone. Longitudinal ribs and grooves commonly mark the
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exterior surface. Superficial resemblance to a large horn coral is striking. The upper valve is
reduced to a nearly flat lid-like structure in which the ligament is shifted to a subcentral position
and the elongate teeth are modified for vertical motion of the valve, rather than movement along
a hinge. Rotational movement of the upper valve is prevented by the configuration of the teeth
and the manner in which they fit into sockets on the fixed valve. Most of them were gregarious
reef dwellers that lived in a fixed location from the larval stage onward. They seem to have been
confined to warm-water areas of the Cretaceous seas; and in deposits laid down in such
environments, they include many important index fossils.
Cephalopoda (Ammonites, Nautilus, Squid, Octopus)

There are only 400 species of known, living cephalopods. In the past they were an
important and diverse group of marine predators. There are over 10,000 fossil species. Nautiloids
were dominant predators during the early and mid Paleozoic, while ammonoids dominated
during the Mesozoic.
Cephalopods have large eyes that parallel the vertebrate eye in structure and function.
Part of the foot has been modified into grasping arms and tentacles equipped with suction cups
to capture prey and draw it to the mouth. A beak like set of jaws tears the prey to pieces to be
swallowed. Water is drawn in under the mantle to aerate the gills. The water is expelled through
a tube called the hyponome, which propels the animal.

Nautiloidea
These first appear in the Ordovician and persist, albeit very reduced in diversity, to the
present day. There are more than 2500 fossil species but only one genus with two or three
species, survives today (Nautilus).
Nautiloid skeletons are straight, curved, or planispiral coiled (visualize coiling a rope or
garden hose on the floor). The tapering aragonitic tubes are divided into chambers by simple
calcite plates called septa. Where the septa and external wall meet, sutures are formed.
Nautiloid sutures are simple, circular joints around the shell. The last and largest chamber is the
living chamber. All the other chambers were living chambers that the animal outgrew. A small
tube, the siphuncle (do not confuse with the bivalve siphon), connects all the chambers and
serves to regulate fluid and gas pressures in the chambers for buoyancy. The siphuncle is usually
centrally located within the chamber of nautiloids. Calcite is also deposited in chambers for
ballast. Often the actual skeleton is not preserved, and what we often get instead is a cast of the
chambers and siphuncle. Fossil nautiloids are easily distinguished from their ammonoid cousins
by the simple, circular grooves running around the cast, indicating where the now missing septa
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once were.

Ammonoidea
Ammonoids are generally similar to nautiloids, but differ significantly in their sutures.
Instead of simple rings, ammonoid sutures range from simple wavy patterns to extremely
complex crenulations. The sutures are classified as goniatitic, ceratitic, or ammonitic (see next
page), in increasing level of complexity, and also depend on the arrangement of lobes and
saddles. The zigzag pattern results from the fact that the septa are not simple walls, but are
themselves complexly folded to aid in the prevention of implosion at great depths.
Unlike nautiloid fossils, preservation of shells is relatively common, in both altered and
unaltered condition. The down side to this is that the presence of a shell makes it impossible to
see the sutures, which are on the inside of the shell. The siphuncle is often near the outer edge of
the chambers, not centered as in nautiloids. Shells are more often coiled than straight, and
external knobs, ridges, and other ornamentation are common.
Ammonoids first appear in the fossil record in the Devonian, and steadily increased in
diversity and abundance throughout the late Paleozoic on through the Mesozoic. They abruptly
go extinct at the K-T boundary. Because of their wide distribution and complexity, they make
outstanding index fossils and are often used in biostratigraphic correlations.

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HEMICHORDATA

Hemichordates possess an embryology similar to echinoderms. They possess gill slits and
a short dorsal tubular nerve chord. Examples include Acorn Worms, Pterobranchs, and
Graptolites

Graptolithina

Graptolites were colonial, marine organisms that constructed and lived in chitinous tubes.
They fed by catching passing microscopic organisms on mucus-covered tentacles. Some were
attached to the substrate, but most formed colonies that simply floated in the water column. The
individual animals lived in small chambers called autotheca that were strung together along a
connecting thread, the nema. The colony was a branch shaped structure, the stipe. Stipes may be
alone or form multi-branched colonies called rhabdosomes.
Graptolites appeared in the Cambrian and went extinct in the Mississippian. They
reached high levels of diversity and abundance during the Ordovician and Silurian. Their fossils
are most often carbonized in black shales. Black shales were deposited mainly in deep marine,
anoxic conditions that prohibited the rapid breakdown of organic material. After death, graptolite
colonies would sink into these deep waters and be preserved in the bottom sediments. They are
also found in limestones, sandstones, and cherts. Because of their wide distribution and
morphological diversity, they are exceptionally good index fossils.

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 A Simple Dichotomous Key for the Identification of Some Common Fossil Invertebrates

Key Step Description Go To/Fossil Group

1. A. Colonial (with many uniformly shaped openings). Go to 2

B. Non-colonial (without a large number of openings). Go to 3
2. A. Openings smaller than 1 mm in diameter (microscopic). Go to 13
B. Openings larger than 1 mm in diameter. Go to 14
3. A. Coiled (shell in the form of a spiral). Go to 4
B. Non-coiled (shell does not spiral). Go to 6
4. A. Shell chambered internally. Go to 5
B. Shell non-chambered (no internal chamber walls) Gastropod
5. A. Chamber walls are straight or gently curved Nautiloid
B. Chamber walls are highly curved and wrinkled Ammonoid
6. A. Bilateral (two-fold) symmetry. Go to 7
B. Pentameral (five-fold) symmetry. Go to 9
C. Radial symmetry. Go to 11
7. A. Shell composed of two paired valves. Go to 8
B. Segmented body, divided into 3 longitudinal lobes Trilobite
8. A. Paired valves of the shell are mirror images Pelecypod
B. Paired valves are not mirror images;
each valve has its own bilateral symmetry Brachiopod
9. A. Globular or vase-like shell composed of polygonal plates;
stem (or attachment mark for stem) at the base. Go to 10
B. Globular or disk-like; without stem (or attachment mark) Echinoid
10. A. Five, prominent, radiating grooves on the shell Blastoid
B. No prominent grooves on shell; may have many long arms Crinoid
11. A. Conical or tapering shell. Go to 12
B. Cylindrical (non-tapering), composed of many flat, disk-like
“segments” that are circular in cross-section Crinoid (stem only)
12. A. Curved walls separating internal chambers in hollow shell. Go to 5
B. Radiating, vertical partitions inside the shell Rugose Coral
C. Shell solid (not hollow) with tiny, radiating, fibrous crystals in
cross-section Belemnite
13. A. Massive, rounded and bumpy; densely layered internally Stromatoporoid
B. Twig-like, button-like or sheet-like; not densely layered Bryozoan
14. A. Radiating, vertical partitions within each opening Scleractinian Coral
B. Horizontal partitions forming vertical stacks of tiny chambers Tabulate Coral

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Ecology
One of the neat things about paleontology is that we can use Uniformitarianism and
functional morphology to determine how a fossil organism used to live back when it was alive
(what niche it filled). For example, by looking at the fossils and knowing a little bit about their
modern relatives, we can determine whether something burrowed in the sediment, swam in the
water column, collected its food from the seawater (filter feeding) or actively hunted for food. In
addition, an increase in ecological complexity (more niches filled) often corresponds to increases
in diversity, while mass extinctions are often associated with lower ecological complexity.
The method of evaluating ecospace that we will use for GEO 405 was developed by
Andrew Bush and Richard Bambach as a way to compare ecological structure (niches filled)
within habitats and through time in a rigorous and standardized way.

Excerpt from Bambach et al. 2007 (paper in Canvas)

AUTECOLOGY, MODES OF LIFE AND ECOSPACE
Autecology, the relationship of animals as individuals to their biotic and abiotic
environment, focuses on modes of life. We have developed a classification of general modes
of life for marine organisms based on tiering position in relation to the substratum ⁄ water
interface, motility level and feeding strategy, which we regard as fundamental ecological
parameters (Bush et al. 2007). Tiering is important because each location with respect to the
sea bottom has particular physical characteristics that require adaptations to function
efficiently. Motility level reflects the spectrum of activities an organism can achieve in
response to disturbance and ⁄ or other external stimuli. Feeding strategy encompasses the
means of acquiring the energy necessary to maintain life. In effect, these axes describe the
basic autecology of an organism. The categories into which each axis was divided are listed
in Table 1 [see next page]. The categories into which each axis was divided are listed in
Table 1 and described fully by Bush et al. (2007).
This classification permits assignment of all marine animals to particular places in a
three-axis grid. Those locations define general modes of life and the three-dimensional
construct based on the three axes represents ecospace (ecological space) (Text-fig. 1A). The
three axes form a complete theoretical ecospace. We have tried to define the six subdivisions
of each axis so that all possible combinations of tiering, motility level, and feeding
mechanism are accounted for. The six subdivisions for each of the three axes designate 216
combinations, which we term modes of life (Text-fig. 1B).

ECOSPACE, NICHES AND GUILDS

Ecospace (location in a topographic setting, potential for motion or the maintenance
of a fixed position, and method of acquiring food) is always present, whether animals are
living in each tier or functioning in each possible way or not. The challenge for organisms is
to construct the adaptations necessary to exploit ecospace. Thus, complex ecosystems
develop as organisms acquire the behaviours and adaptations necessary to construct the
ecological interactions that enable them to exploit un-utilized opportunities. In effect, the
filling of ecospace over time conceptually parallels Valentine’s ‘tesserae’ model of evolution,
in which new taxa fill pre-existing empty ‘niches’, with small morphological changes
(speciation) making small steps and large changes (the origin of higher taxa) making large,
but rare, jumps (Valentine 1980, 1981; Walker and Valentine 1984).
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25
26
What to do in this lab:

This lab serves as a broad introduction to the different invertebrate groups (and their
preservation styles). In the lectures prior to this lab, we discussed fossilization and taphonomic
processes as well as biases in the fossil record, but we haven’t talked much aboutdifferent fossil
taxa. In this lab you will synthesize what you have learned about fossils, preservational
styles, and depositional environments.

Go to the Digital Atlas of Ancient Life (www.digitalatlasofancientlife.org/) and look at

the fossils of different invertebrates in the Virtual Collection. Explore the invertebrate
collections (taxa listed below). Create five “Cheat Sheets” for yourself (see template in Canvas
or on the next page) for the BODY FOSSILS of taxa from at least 3 different Phyla (pleural of
phylum) or bolded group below.

• Stromatolites • Phylum Mollusca

• Class Gastropoda
• Foraminifera • Class Cephalopoda
• Class Bivalvia
• Phylum Porifera (Sponges)
• Class Archaeocyatha • Phylum Arthropoda
• Class Calcarea • Chelicerata
• Class Demospongiae • Crustacea
• Class Hexactinellida • Trilobita
• Class Stromatoporoidea
• Phylum Echinodermata
• Class Anthozoa (Corals) • Class Asteroidea
• Rugose corals (Rugosa) • Class Crinoidea
• Tabulate corals (Tabulata) • Class Ophiuroidea
• Hexacorals (Scleractinia) • Class Edrioasteroidea
• Class Echinoidea
• Phylum Brachiopoda • Class Cystoidea
• Class Blastoidea
• Class Lingulata
• Class Strophomenata
• Class Rhynchonellata • Order Graptolithina (Graptolites)

• Phylum Bryozoa • Trace Fossils

• Feeding traces
• Phylum Annelida • Resting traces
• Locomotion traces
• Dwelling traces
• Predation traces
• Coprolites

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GEO 405: Survey of Invertebrates – Personal Cheat Sheet

Taxon Identification:

Age Range (If known):

Physical traits used to I.D. the specimen:

How would you tell this apart from a similar fossil?

Include photos of the specimens you looked at here.

Hint: use Dichotomous Key

Style(s) of Preservation (If you can tell):

Part(s) of the organism (What’s missing?):

Depositional Environment(s) (If you can tell):

Ecology of Organism:
(Shade in the appropriate boxes)

Comments:

Use the Digital Atlas of Ancient Life: www.digitalatlasofancientlife.org/