Lab #6: Molluscs: Cephalopods
At the end of this lab, you should be able to:
- Identify a fossil as a nautiloid, ammonoid, belemnoid, scaphopod or
Tentaculites.
- Recognize the age of an ammonoid from its sutures.
- Know the skeletal structure and material of each
of these animals.
- Know the ecological characteristics of each of
these animals.
- Know the geologic range of each of these groups.
- Know a few important genera (mentioned by name
in this handout) for each group.
- Be able to describe the mathematical variables
controlling molluscan growth.
Display: Cephalopods
I. Soft part morphology - For the tiny squids in the resin blocks,
use fig. 13.14 in your book to identify the tentacles, the hood, the hyponome (the tube it shoots water out through), and the
eyes (closed in these samples).
II. Hard part morphology - Nautilus shell and ammonites - be able to identify the septa
- A. Nautilus: (#832) find the septa, chambers,
septal necks
(where the siphuncle ran. The organism uses the long soft rope of
the siphuncle to regulate the amount of fluid and gas in the chambers).
- Structure of the chambers: compare the Nautilus (#832) with the three ammonite specimens, and #11 Baculites.
- A. In Nautilus, the chamber walls (septa) are
geometrically very simple - just a domed curved shape. Where the
septa meets the wall of the shell, it creates a simple curved
line. If the chambers were filled with sediment or minerals, the
septa would show up as that curved line - the suture.
- B. In these three fantastically beautifully preserved ammonite specimens,
the shells were preserved with nothing filling up the chambers.
You can see how complicated the septa are - not the simple curved dome
of the Nautilus, but a corrugated surface that creates a very
complicated suture. Look carefully at the septa to see the
complicated structure.
- C. #11 Baculites: This is a single chamber of a straight ammonite called Baculites - actually a mold
of that chamber (the chamber is an empty space, while this is the mud
that filled up the chamber). You can see the shape of the septa on each end. This specimen shows the complicated line where the septa meet the chamber wall - the sutures.
- Siphuncle: nautiloids and ammonoids use a long strand of
soft tissue the runs back through the septa to regulate the amount of
fluid and gas in the shell. The siphuncle is supported by small
projections of the chamber wall, called a septal neck.
- A. Find the septal necks on the Nautilus. In this species and in many nautiloids, the siphuncle runs near the center of the septa.
- B. If you look carefully at the large straight ammonite, you can see a tiny circle at the narrowest part of the shell. That's where the siphuncle passed through the shell.
- C. #11 Baculites: At the narrow end of the shell you can see a small circle of shell - that's the septal neck for the siphuncle.
III. Kinds of Cephalopods
A. Primitive straight-shelled cephalopods: These organisms were
once
all classified as nautiloids, but are now separated in several different
subclasses. ALl of these groups are Paleozoic in age, most Early to
Middle Paleozoic. Our samples are mostly too poorly preserved to
recognize, but you can see these things:
- 1. Straight sutures (#1034, 982).
- 2. On some specimens (#991, 1617, 1621) you can see the dome shape for the chambers.
- 3. #1100 & 1506: you can see a very small siphuncle
- 4. #994 Endoceratid with siphuncular deposits - look for the
big crystals in a very wide siphuncle. The organism used these
deposits as counterweights to keep the shell horizontal.
B. Nautiloids: Organisms like Nautilus have existed since the Paleozoic.
- #2014 Eutrephoceras dekayi: Notice the gently curved
sutures. The outermost chamber of the nautiloid is filled with
mud, and the inner coils are full of calcite. The hole in the
septa for the siphuncle was too samll to allow mud to get through, so
the inner chambers were not filled with sediment when the sediment was
deposited. This often happens, but usually the shell crushed as
more sediment accumulated. In this case, the mud was micrite -
calcite mud - so the water running through the sediment contained a lot
of dissolved calcite that deposited calcite crystals inside the shell.
C. Ammonoids:
- Nautiloids v ammonoids: You already saw that ammonoids have
much more complex sutures than nautiloids. The convulted
structure of the septa allowed the ammoinoids to adapt in two other
ways: 1) thinner shells and 2) more streamlined shapes. This meant
that ammonoids could grow larger than nautiloids without having to
produce huge amounts of shell. They could also become more
effective predators, because they could become fast, streamlined
swimmers. Compare the shapes of the ammonoids you see in this part
of the display with the very round nautiloid (#2014).
- 1. Sutures: The first ammonoids had septa only slightly
more complicated than nautiloids. Over time, the septa became more
convoluted, with more complex sutures. Use fig. 13.17 in your book to recognize the kind of suture.
- a. Goniatites (#1712 Gonioloboceras, #98, #281):
Goniatitic sutures are simple wavy lines. These ammonoids are
typically plump and more spherical, like nautiloids. Goniatites
are usually Late Paleozoic in age.
- b. Ceriatites (# 279 Phaneroceras, #49): In the
Permian and Triassic, ammonoid septa became more complex. This
produced ceriatitic sutures, where the sutures forms a rounded wiggle,
and then a spiky wiggle.
- c. Ammonites (unnumbered Scaphites, unnumbered large pachydiscid, #993 Baculites, unnumbered Baculties):
Jurassic and Cretaceous ammonoids usually have much more complex
septa, and so complicated sutures. Ammonitic sutures look a
bit like very fancy mittens, with many wiggles that represent the
corrugations in the septa.
- 2. Ornamentation.
- a. #416 Epenogonoceras: Very smooth, flat streamlined
ammonites were probably fast swimmers. This ammonite coiled
around one point, so the outer whorls completely covered the inner
whorls, leaving no indentation to create turbulence.
- b. #834 Haugia, #432, unnumbered specimen:
Many fine ribs actually enhance the hydrodynamics of the shell by
holding a small boundary layer of water around the shell. This
reduces drag on the water as the shell moved. These forms were
probably also fast swimmers.
- c. #398 & 21 Pervinquieria: Very large ribs suggest that
the ammonite was not very streamlined. The large ribs may have
provided some protection from predators (fish or marine reptiles) by
making the shell wider and harder to bite. However, the knobs on
the shell are not thickened parts of the shell, as they are on highly
ornamented gastropods. The shell would still have been thin and
vulnerable, so the ribs may not have a protective function after all.
- 3. Heteromorphs: Some families of ammonite evolved uncoiled shapes.
- a. #833 Scaphites coiled for part of its life and then grew straight.
- b. #1551 and unnumbered Baculites coiled only as an embryo in the egg, then grew straight for the
rest of its life. #1551 Baculites is a section from an organism
probably about a meter long.
- c. See the pictures for some other rather spectacular and bizarre heteromorph ammonites.
- d. Because most heteromorphs are not hydrodynamic, they were probably not active predators like the other
ammonites. They may have lived high in the water column,
filtering water, or lived near the bottom as scavengers. However, Baculites may have been able to swim.
D. Belemnoids: Squids have a terrible fossil record except for one
group of squids which grew internal skeletons: the belemnoids.
- 1. Rostrums - unnumbered specimens: The skeleton included an
aragonite phragmacone and a calcite rostrum (see diagram and the picture
of the entire fossilized animal). Usually you find just the
calcite rostrum, but one of these specimens still has the aragonite that
originally covered the calcite.
- 2. Phragmacone - unnumbered specimens. Some of our specimens show
the conical hole where the chambers of the phragmacone were.
- 3. Crystalline structure of the rostrum - unnumbered broken specimens.
Note the radiating calcite crystals that make up the rostrum.
Questions
1. #605 - This sample contains both nautiloid and ammonoid fossils. Which
is which? How do you know?
2. unnumbered - What kind of sutures does this organism have? Estimate the
age of the fossil.
3. #427 - There are two different classes of mollusc in this sample,
both with long straight tubular shells. What are they? How can you tell
them apart?
4. #606, three unnumbered specimens: What kind of preservation is this?
5. #874 - What kind of cephalopod is this? What is your evidence?
6. #832 - What group of fossil cephalopods does this belong to? What
is your evidence? How is it preserved?
7. #432 - What does the ornamentation of this cephalopods tell you about how it lived? Is it a nautiloid or an ammonite?
8. Unnumbered - Is this an ammonite or a nautiloid? Use the sutures to estimate the age of the fossil.
9. Unnumbered - Was this ammonite a fast or slow swimmer? How do you know?
10. Unnumbered - Notice that these ammonites look a little odd. How did these ammonites come to look like this?
11. two unnumbered specimens - No question for
these. They are here so you can see what happens to ammonites when
they weather. The suture lines get weathered out, leaving the
clay or calcite that filled the shell in a very complicated
pattern. Ranchers on the west side of the Valley sometimes think
they have discovered fossil brains when they find a large weathered
ammonite.
12. unnumbered - The large ammonite labeled 12a was
sexually mature when it died. In modern Nautilus, when the animal
reaches sexual maturity, it stops growing. The last chamber is
much smaller than the one before it, and the last two sutures are close
together (see picture). Look carefully at the sutures on 12a and
notice how the last two sutures overlap, while the next suture is
farther back.
The ammonite fossil labeled 12b includes the body chamber of
the organism - the space on the outside of the last septa where the
animal's body was. You can recognize the body chamber because it's
filled with mud (the rest of the shell is filled with calcite) and it
has no sutures (because there were no more chamber walls).
Your job is to look carefully at a minimum of five of the Scaphites
fossils in the boxes labeled 12c. For each one, figure out 1) is
the body chamber here? and 2) if so, was this animal sexually mature
when it died?
13. unnumbered belemnoids. No question here - just
look carefully. Both of these belemnoid rostrums were attacked by
boring organisms after the squid died. One was attacked by
clionid sponges - look for the tiny holes where the sponge dissolved a
hole in the belemnoid, and then moved into the belemnoid and continued
to dissolve tunnels through the belemnoid. The other belemnoid was
bored by some kind of crawling organism, leaving trackways in the
surface.
14. unnumbered - This is a pretty cool specimen of a primitive
straight cephalopod that was overgrown by another organism. The
cephalopod is the circular shell (on the polished ends) with the knobby
shell (on the weathered side). You can see a crescent of mud that
entered the empty shell and the crystals of calcite that grew in the
part of the shell that did not fill with mud. On one of the
polished ends you can see the hole of the siphuncle. Your job is
to:
- identify the organism that overgrew the cephalopod shell
- draw a little cartoon that shows what happened to this cephalopod
from the time it was alive, swimming just above the ocean bottom, until
it became fossilized.
15. Just a pile of pretty ammonites to look at. No question.
Constructional Morphology
Mollusc shells lend themselves well to mathematical modeling because
of the simplicity of their construction. All mollusc shells are essentially
some 2-dimensional shape coiled around an axis. What that 2-dimensional
whorl shape is depends upon the organism. For example, in some snails, the
whorl shape is almost a circle, as seen in the round aperature of the animal.
In Conus, the whorl shape is essentially a triangle. That shape starts
out very tiny and expands as the animal grows and the whorl shape is rotated
about an axis. Note: this whorl shape is not necessarily the same shape
as the aperture. In Conus, the shell overlaps itself tightly, so
the aperture is reduced to only a portion of the triangle that makes the
shell.
The rotation of the whorl about the axis can also vary. The whorl can
move out from the axis either quickly or slowly, producing whorls that overlap
completely, or whorls that do not even touch. The whorls may spiral in a
single dimension, producing flat shells like ammonoids, or may coil down
an axis, producing spired shells like gastropods.
We can describe mollusc growth, then, in four variables (see figs. 7.2
&7.3):
- Whorl shape
- Expansion rate (W): rate at which the cross-sectional area of the shell
increases
- Distance of aperture from the axis (D): a measure of how rapidly the
coil spirals outward
- Translation rate (T): rate at which the spiral moves along the axis
Why do we care? A couple reasons. First, if mollusc growth is controlled
by only four variables, it is probably controlled by a small number of genes.
It becomes easy to imagine how large changes in molluscan morphology are
possible through very small genetic change, making molluscs an evolutionarily
adaptable group.
Second, analyzing molluscs in this way gives us an idea of what is structurally
and biologically important in constructing shells. There are many possible
molluscan forms, but only some are found in nature. This analysis gives
us an idea why.
I. Examine the mollusc shells on the back tables. For each, describe
the shape of the whorl and the general value (low, medium, high) or each
of the other variables (W, D, T). Find the shell with the highest and lowest
values of each variable. Use figs. 7.2 and 7.3 to help you.
There are two sets of the same shells - CM1 and CM2. You just need to look at one of them.
II. Look carefully at Fig. 7.3. Notice that there no living or fossil
organisms fall in either the back right quadrant of the box (high D, high
W) or in the lower left quadrant (high T, high W). Why not? What would such
creatures look like?