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Brachiopods |
Ancient Shells at the Falls of the Ohio
Part I: Brachiopods
What would
it have been like to be a shell collector during the Devonian
period, between 408 and 360 million years ago? Today collector's
find shells in a variety of shapes, sizes and colors washed
up on a beach. Those are shells of mollusks, usually gastropods
(snails) and pelecypods (clams).
The fossil-bearing rocks at the Falls of the Ohio indicate
that a Devonian beachcomber would have found mollusks and
other types of shells washed up. The most common shell-bearing
creature in the Devonian at the Falls is virtually never found
washed up on a beach today - that is the brachiopod (pronounced
brak - e - o -pod). They are often called lampshells because
some varieties resemble a Roman oil lamp.
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| Figure
1: Shell Symmetry |
What
is a brachiopod?
Brachiopods are invertebrates -- animals without a back bone
-- in the phylum Brachiopoda. They have two shells that may
be composed of calcium carbonate (the minerals calcite and
aragonite) or a phosphatic mineral (like your teeth). Most
brachiopods live on the ocean floor, though some burrow. Fossil
records indicate that they have never lived in a freshwater
ecosystem (like the Ohio River).
Perhaps the easiest method to differentiate a brachiopod from
a clam is by shell symmetry (see figure 1). Each clam shell
or valve is a mirror image of one another. Brachiopods are
symmetrical from side to side, so that the top and bottom
shells look different.
The internal anatomy of brachiopod is very different from
a clam. Brachiopods have a coiled feeding organ called a lophophore.
The lophophore may be occasionally preserved in some Devonian
brachiopods found at the Falls of the Ohio and elsewhere in
the county (see figure 2).
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| Figure
2: Orthospirifer fornacula (Hall) showing lophophores |
There
are two major divisions or classes of brachiopods: inarticulate
and articulate. Inarticulate brachiopods were the first to
develop, and can be found in rocks dating back to the earliest
Cambrian period, more than 570 million years ago. These brachiopods
are not abundant in the local Devonian rocks. Petrocrania
hamiltonae (Hall) is perhaps the most common inarticulate
brachiopod (see figure 3).
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| Figure
3: Petrocrania hamiltonae (Hall), an articulate brachiopod |
Inarticulate brachiopods open and close their
valves with muscles and do not rely on sockets and teeth that
are a characteristic of articulate brachiopods. Devonian species
of this class are often found attached to other organisms
-- such as another brachiopod shell or horn coral. The bottom
ventral valve may be cemented to a firm surface, while the
upper dorsal valve is opened for feeding.
Articulate brachiopods have two valves that are different
size. The larger shell is called the pedicle valve. It contains
a hole through which a fleshy stalk called a pedicle attaches
to a substrate (rocks or sediment on the sea floor). The pedicle
acts as an anchor which firmly holds the brachiopod in place.
(Unlike clams which can move through sediment, the brachiopod
is fixed throughout its adult life.) The pedicle valve contains
projections called teeth (see figures 4a & b) that fit
into sockets on the opposite brachial valve.

The
inside of some brachiopod shells contain muscle scar patterns
(see figures 4a & b). The placement of these scars help
paleontologists determine the placement of important muscles.
Articulate brachiopods come in a variety of shapes. Valves
may be convex (bow out), concave (bow inward) or flat. There
can be variation in the shape of the brachial and pedical
valve. One might be concave, the other flat or convex. The
hinge line is where the two valves of articulate brachiopods
come together. It can be straight (as in figures 4a, 5, 6a)
or curved (as in figures 6a, 7a & b, 8a & b). It can
be short, relative to the width of the shell (figure 8b),
or long (figure 4a).
Brachiopod shells often show interesting external ornamentation.
Brevispirifer gregarius (Clapp) often shows pronounced growth
lines and is strongly ribbed (figure 5). Protoleptostrophia
perplana (Conrad) and Rhipidomella penelope (Hall) are examples
of brachiopods that have costae, which are very fine ridges
on the outer surface of the shell (see figures 6). Invertrypa
spinosa (Hall) and Productella spinulicosta Hall are two brachiopods
with spines on the external surface of the shell (figures
7). Invertrypa spines are usually not preserved. Spines helped
stabilize brachiopods in muddy sediment. Some brachiopods,
like Athyris and Cryptonella (figures 8) are very smooth.
Evidence of color patterns on valves are very rare from brachiopods
of this age.
Types of brachiopods at the Falls
There are six types or Orders of articulate and two Orders
of inarticulate brachiopods that may be found at the Falls
(see table 1). Most brachiopods are found on the "upper
fossil beds," rather than the coral-rich "lower
fossil beds." As a result, this group of fossils is visible
for much of the year. Although brachiopods are common in rocks
at the Falls, please remember that fossil collecting here
is prohibited.
| Table
1 Brachiopods of the Falls area |
| Inarticulate
(Class Inarticulata) |
P.
pavilionensis Hall |
| Acrotretids
(Order Acrotretida) |
P.
thusnelda (Nettelroth) |
| Acanthocrania
granosa (Hall and Clarke) |
Rhynchonellid
(Order Rhynchonllida) |
| Craniella
hamiltoniae (Hall) |
Attribonium
gainesi (Nettleroth) |
| Craniops
sp. |
Cupularostrum
carolina (Hall) |
| Orbiculidea
doria (Hall) |
C.
depressa? (Kindle) |
| *O.
lodiensis (Vanuxem)? |
C.
louisvillensis (Kindle) |
| Petrocrania
greenei (Miller |
C.
nitida (Kindle) |
| P.
hamiltoniae (Hall) (figure 3) |
C.
sappho (Hall) |
| P.
sheldoni (White) |
C.
tethys (Billings) |
| Philhedra
crenistria (Hall) |
Cyclorhina
nobilis Hall |
| Roemerella
grandis (Vanuxem) |
Spiriferids
(Order Spiriferida) |
| *Schizobolus
concentrica (Vanuxem) |
Acrospirifer
duodenaris (Hall) |
| Lingulids
(Order Lingulida |
Ambocoelia
umbonata (Conrad) |
| Glossina
triangulae (Nettleroth) |
Athyris
fultonensis (Swallow) (figure 8a) |
| *Lingula
spatula Vanuxem |
Atryparia
devoniana (Webster) |
| Articulate
(Class Articulata) |
A.
ellipsoida (Nettelroth) |
| Orthids
(Order Orthida) |
Brevispirifer
(?) davisi (Nettelroth) |
| Rhipidomella
penelope (Hall) (figure 4b, 6b) |
Brevisprifer
gregarius (Clapp) (figure 5) |
| Tropidoleptus
carinatus (Conrad) |
Cyrtina
crassa (Hall) |
| Schizophoria
striatula (Schlotheim) |
C.
hamiltonensis (Hall) |
| Pentamerids
(Order Pentamerida) |
C.
hamiltonensis var. recta (Hall) |
| Pentamerella
arata (Conrad) |
Elyta
fimbrata (Hall) |
| P.
indianaensis (Kindle) |
E.
wabashensis (Kindle) |
| Emanuella
subumbona (Hall) |
Fimbrispirifer
divaricatus (Hall) |
| F.
grieri (Hall) |
Invertrypa
spinosa (Hall) (figure 7a) |
| Mediospirifer
audaculus (Conrad) |
M.
(?) manni (Hall) |
| M.
(?) segmentum (Hall) |
Megakoglowskiella
raricosta (Conrad) |
| Meristella
barrisi (Hall) |
M.
nasuta (Conrad) |
| Nucleospira
concinna (Hall) |
Orthostrophia
fornacula (Hall) (figure 2) |
| Paraspirifer
acuminatus (Conrad) (figure 1) |
Parazyga
hirsuta (Hall) |
| Pentagonia
unisulcata (Conrad) |
Spirifer(?)
arctisegmentum (Hall) |
| Spirifer(?)
varicosus (Hall) |
Strophomenids
(Order Strophomenida) |
| "Chonetes"
acutiradiatus (Hall) |
*Chonetes
lepidus Hall |
| "C."
subquadratus (Nettelroth) |
Devonochonetes
coronatus (Conrad) |
| Eodevonaria
arcuata (Hall) |
Floweria
chemungensis var. arctostriatus (Hall) |
| Leptaena
rhomboidalis (Wilckens) |
Longirostra
mucronatus (Hall) |
| Megastrophia
concava (Hall) (figure 4a) |
Protodouvillina
inequistrata (Conrad) |
| Protoleptostrophia
perplana (Conrad) (figure 6a) |
Productella
semiglobosa (Nettelroth) |
| P.
spinulicosta Hall (figure 7b) |
Stropheodonta
inequistriata (Conrad) |
| S.
demissa (Conrad) |
S.
plicata Hall |
| Terebratulids
(Order Terebratulida) |
Camarospira
eucharis (Hall) |
| Centronella
glansfagea (Hall) |
Cranaena
(?) jucunda (Hall) |
| C.
harmonia (Hall) |
C.
lincklaeni (Hall) |
| C.
romingeri (Hall) |
Cryptonella(?)
lens (Hall) (figure 8b) |
| C.
ovalis (Miller) |
C.
(?) sullivanti (Hall) |
| * From the New Albany Shale
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Suggested Reading
The Audubon Society Field Guide to North American Fossils,
I. Thompson. Published by Alfred A. Knopf, 1982. Good general
fossil book.
The Devonian Fossils and Stratigraphy of Indiana, E. M. Kindle,
Indiana Dept. of Natural Resources, 1901. Out of print. Identifies
most Devonian brachiopods found in Indiana. May be found in
college, university and large public libraries.
Fossil Invertebrates, edited by R.S. Boardman, A.H. Cheetham,
and A.J. Rowell. Published by Blackwell Scientific Publications,
1987. Chapter 16. College level text.
Kentucky Fossil Shells, Henry Nettelroth. Kentucky Geological
Survey, 1889. Out of print. The first comprehensive, well
illustrated book of fossil brachiopods and mollusks from Kentucky.
This is a rare book, but may be found in large public or university
libraries in the region.
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