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A SEA WITHOUT FISH
LIFE IN THE ORDOVICIAN SEA
OF THE CINCINNATI REGION
David L. Meyer and Richard Arnold Davis
With a chapter by Steven M. Holland
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This book is a publication of
Indiana University Press
601 North Morton Street
Bloomington, IN 47404-3797 USA
http. //iupress. indiana. edu
Telephone orders: 800-842-6796
Fax orders: 812-855-7931
Orders by e-mail: iuporder@indiana. edu
© 2009 by Richard Arnold Davis and David Lachlan Meyer
Except chapter 15 © 2008 by Steven M. Holland
ll h d
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The wor ldwi de fame of the fossi ls and rocks of the Ci nc in na ti , Oh io , re
gion gr ew out of the labors of myria d am at eur fossil col l ector s. T h e curre nt
embodiment of those folk is the "Dry Dredgers, " a group founded in Cin
cinnat i in 1942 and, to this day, ded icat ed to col lec ti ng and un der st and ing
those fossils.
W e d e d i ca te this v o l u m e to the " D r y D r e d g e r s " an d to th e host o f fo ss il
col lectors they represent. Vos salukimus!
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CONTENTS
ix PREFACE
x i i l ACKNOWLEDGMENTS
xv REPOSITORIES OF FOSSILS ILLUSTRATED IN THIS BOOK
1 Introduction
1 <
2 Science in th e Hinterland
15 THE CINCIN NATI SC HO OL OF PALEO NTOL OGY
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9 Molluscs
11 7 HARD, BUT WITH A SOFT CENTER
10 Ann elids and Worm -Lik e Fossils
143 <
11 Arthropods
147 TRILOBITES AND OTHER LEGGED CREATURES
12 Echinoderms
167 A WORLD UNTO THEMSELVES
13 Graptolites and Con odo nts
19 5 OUR CLOSEST RELATIVES?
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PREFACE
Two princi pal goals moti vate d us to write this bo ok . First, kn ow le dg e of the
Earth's ancient history from geology provides a powerful lesson about the
eve r-c ha ng ing nature of the planet, and the ancien t history of one's ho m e
region can be particularly mean in gf ul . The present natu re of the lands cap e
in the Cincinnati region (southwestern Ohio, northern Kentucky, and south
eastern Indiana) is the pro du ct of its most recen t ge ol og ic history, the Pleist o
cene ice Age, when continental ice sheets repeatedly forced their way as far
south as the Ohio River. As recently as 20, 000 years ago, much of southwest
ern O h i o was cov ere d with an ice shee t m u ch as Green land is today. As the
glaciers receded, melt waters carved the present valleys and left a mantle of
debris that det erm ine d the t opograp hy, dr ain age , soils, and veget atio n of the
region. A magnif icent lee Age exhibit at the Cincinnati Museum Center
enhances public awareness of the profound environmental changes that took
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Many local residents who have been fascinated by the fossils und
col lec ted and studied t he m alm ost since the earliest settle ments of the
teenth and ninet eent h centuries. G ener ati ons of geologists and paleo
gists from abro ad ha ve visited the regi on and writt en of the abun da nt
and the strata, including the pioneering British geologist Charles Lye
1842. Because the Cincinnati region has been a focus for geological res
by so man y sci en tis ts over so m a i n ye ar s, there ex is ts toda y a vast a m o u
inf or mat io n ab ou t the fossils and rocks of the region. This info rma ti
scatt ered in m an y sou rce s, i nc lu di ng the latest issues of so me of the w
lead ing international geolog ical journals, Internet websites, and num
type s of publi cati ons, so me wid ely availabl e, some obs cure. Mu ch of the
work d e sc r i b i n g n e w spe ci es of C i n c i n n a t i fo ss il s dates to the secon d h
the nineteenth century, and is found in periodicals no longer published,
as the Cincinnati Quarterly journal of Science, The Paleontologist, and
Journal of the Cincinnati Society of Natural History. No sin gle library h
all of the geologi cal informa tion publi shed about the Cin cin nat i re
Mo re ov er , most stud ies deal with on ly a small fra ction of the total fossilness of the reg ion , and, mos t impo rtan tly for us, there has never b
synt hesi s of the vast ran ge of fossil diversity and its geol og ica l con text . I
b o o k we present a synthesis that wil l r econ stru ct the li fe of the Or d o v
sea in order to show not only what organisms inhabited this sea but also
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not have to sea rch at the bo tt om of the p ag e or the en d of the c ha pte r, or,
even, volume, for the pertinent reference.
Thus, when we refer you to a publication, the literature citation will
be in the f o l lo w i n g format: "(S. A. M i l l e r 1875 ). " T h i s m e a n s that y o u are
b ei n g re fe rre d to a p ub l i c at i o n aut hor e d by S. A. M i l l e r an d p u b l i s h e d in
1875; he nc e, you know w ho said what is bei ng cited and whe n. If you n eed
the complete bibliographic information about that publication, i t is pro
v i ded in the bi bl i ogr ap hy toward the en d of the v o l u m e . In c ases in w h i c h
it is important for you to know the page number within that publication
wh er e the i nfor mat ion or q u o tat i on is f o u n d , t h e l iterature c i ta tion wil l bein the form "(S. A. Miller 1875, 87). "
N a m e s o f O r g a n i s m s a n d G r o u p s o f O r g a n i s m s
By international ag ree men t of zoolo gists , the International Code of Zoo
logical Nomenclature i s the document that speci f ies how the names of
specie s, gene ra, and othe r gro ups of an im al s are stated an d used in scie n
tif ic works (International Co mm is si on on Zoo lo gi cal No me nc la tu re 1999).
Gen era l reco mm end ati on B io of the C o d e enc our age s that the author and
date of every taxon in the speci es gr ou p, genu s gr ou p, or fam ily gro up
mentioned in a publication be cited at least once in that publication and
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Technical Terms Sc ie nc e is replete wit h tec hni cal terms that do not appea r co mm on
and the Glossary non-sc ienti f i c conte xts. To ma ke matters worse, sc ientists often use
mon, everyday terms in ways that are not their common, everyday u
Thus, we felt it important to include a glossary; this is found near th
of the vol um e. I n the interests of spa ce, ho wev er, we hav e not inc
every technical term in this book in the glossary. For its first use,
tec hni cal term is def ine d and is in bo ld fa ce ty pe . Those techn ical
that are used in more than one chapter are listed in the glossary. A t
cal ter m that is used in onl y on e cha pte r, such as the na m e of an ana
cal feature that occur s in onl y on e major grou p of org anis ms, is def ine
first ti me it is used i n the vo lu me ; howe ver , we ha ve not listed such
in the glo ss ar y— ag ai n, in the interests of spa ce. S uc h words are list
the index to the volume.
So wh at do yo u do if yo u find a tec hn ic al te rm that is unf am il
y o u a n d t h e de f in i t io n is not right the re w h e r e y o u e n c o u n t e r the
Firs t, go to the glossary . If th e tec hn ic al te rm is not in the glos sary,
G o d forbid!, the cov era ge of that te rm in the glos sary is insuf ficien t,
go to the i nde x and t he n to the text of the bo ok to wh ic h you are ref
(C ol le ge professors, l ike us, so me ti me s are ac cu se d of stating the ob
Gen era ll y, this is do ne in an atte mpt to ans wer the ques tion s of so m
dent s in a given class befo re they are asked. Ther e is, of cou rse , a d
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ACKNOWLEDGMENTS
W e are very grateful t o the f o l l o w i n g c o l l e a g u e s w h o read p r e l i m i n a r y
drafts of various chapters : Lor en E. Ba bc oc k ( T h e O h i o State Univer sity) ,
Richard Bambach (Virginia Polytechnic Insti tute and State Universi ty),
Steven H. Felton (Cinc inna ti) , Ro bert J . Elia s (Universi ty of Man ito ba) , J .
Mark Erickson (St. Lawrence Universi ty), Steven M. Holland (Universi ty
of Geo rgia ), Steve n Lesli e (Universi ty of Ark ans as, Litt le Ro ck), James
Sprinkle (Universi ty of Texas), and C ol in Sum ral l (Universi ty of Te nn es
see). In particular, we thank Professor Holland for contributing the chapter
on the Ci n c i n n at i an p a le oe n v i r on me n t .
We thank the fo l lowi ng c o l l e a g u e s w h o kindly provided il lustrations for
our use or permitted us to reproduce their i llustrations: Loren E. Babcock
(The Ohio State University), Stig Bergstrom (The Ohio State University), Jon
W Bra ns tra tor ( E ar l h a m C o l l e g e ) D e v i n Bu i c k (University o f C i n c i n n a t i )
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We arc particular ly gra teful to John A g n e w of C i n c i n n a t i w h o pa
"The Cincinnatian" for the cover and color plate, and who also did
draw ing s of a spon ge, a stromat opor oid, a crin oid, and an edrioasteroid
illustrations could not have been completed without the technical and t ic skil ls of Ti mo th y Phi l l ips (D epa rt men t of Geo log y, Universi ty of Ci n
nati) , Evelyn Moh alsk i (formerly of the D epa rt me nt of Geo log y, Unive
of Cin cin nat i) , and Jay Yocis ( Photo graphi c Service s, Universi tv of Ci n
nati). Professor Kevi na Vu li ne c (D epa rtm ent of Agri cult ure and Na
Res our ces, De la wa re State University, Dover) kindly perm itted us to r
duce her drawings that were originally made for an exhibit at the Cincin
Mu se um o f Natural History.
Many colleagues and fr iends al lowed us to photograph specimen
their collections: Steve Brown (Zanesville, Ohio), Fred Collier (former
the M u s e u m of Com p ar at i v e Z ool ogy , Har v ar d Un i ve r si ty) , D an Co
(Cincinnati) , Steven H. Felton (Cincinnati) , Ron Fine (Cincinnati) , B
an d Char lo t te G i b s on (Ci n c i n n at i ) , Br e n d a Hu n d a (Ci n c i n n at i M u s
Ce n te r ) , K e n d al l Hau c r (L i m p e r M u se u m . M i am i Uni v er s ity) , W i l
Heim broc k (Cinc innat i) , Mark Peter (C ol um bu s, Ohi o), and Janice Th
son (Nat ion al Mu s eu m of Nat ural History, Sm ith so nia n Institution).
W i l l i a m B u t c h e r an d D e n n i s Kytasaari o f the N o r t h A m e r i c a n
V e r n e S o c i e t y p r o vi d e d the a c c u r a t e trans lation o f and i nfor mat ion a
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REPOSITORIES OF FOSSILS
ILLUSTRATED IN THIS BOOK
B M N H N atu r a l Hi story M u se u m , L on d o n
C M C I P C i n c i n n a t i M u s e u m C e n t e r , I n ve r te b ra t e P a l e o n t o l o g y C o l l e c t i o n s
F M N H Fi e ld M u se u m of N atu r a l Hi story , Ch i c ag o, I l l i noi s
M C Z H arv ar d Un i ve r s ity , M u se u m of Co mp a r at i v e Z o ol og y
M U G M M i a m i U n iv er s it y , C a r l F . L i m p e r G e o l o g i c a l M u s e u m , O x f o r d , O h i o
O S U O r i o n G e o l o g i c a l M u s e u m , T h e O h i o S t at e U n iv e rs i ty , C o l u m b u s , O h i o
U S N M N at i on al M u se u m of N atu r a l Hi stor y, S mi t hso n i an In st i tu t i on ,
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A SEA WITHOUT FISH
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INTRODUCTION
The vicinity of Cincinnati, in the Ohio River Valley of southwestern Ohio,
includ ing adjacent northern Kent ucky and southeastern Indiana, is am o ng
the most fossil-rich regions in N ort h Am er ic a, if not the entire world. T h e
profusion of fossils in the local li mes to ne and shale attr acted m an y pion eer
ing geologists and paleontolo gists of the nine tee nth cent ury, and mu ch fun
d ame n ta l w or k i n Am e r i c a n p a l e o n t ol og y an d s tr a t i gr ap hy w as ac c o m
plished here. Hun dre ds of fossil spec ies were first dis cov ere d and nam ed fro m
these rocks. Early geolog ists gave th e entire series of strata ex po se d here the
name "Cincinnatian, " and this name was applied to strata of s imilar age
throughout North America. Cincinnatian fossi ls are displayed in museums
all over the world. Researchers, students, and amateur fossil collectors regu
Of the many prolific col-lecting grounds in the
continental interior, none
excels the Ohio river
bluffs at Cincinnati, Ohio.
Here the Upper Ordovi-
cian rocks are almost
literally made of fossils;many are as perfectly
preserved as fossils can
be. The river banks,
road cuts, and even the
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Figu re 1. 2. Diversity of
marine fossil metazoan
families through the
Phanerozoic. The heavy
uppermost curve depicts
the sum of the three
"evolutionary faunas, "
each shaded differently,
while the stippled por
tion below the total
curve represents residual
diversity not accounted for by the three compo
nent faunas. Taxa listed
for each evolutionary
fauna are those taxa that
contribute most heavily
to the diversity of that
fauna. I = Cambrian Fauna, II = Paleozoic
Fauna, and III = Modern
Fauna. From Sepkoski
(1981) and reprinted by
i i f Th P l
ters (820 feet) of inte rbed ded l im est on e and sh ale was deposited du rin
L ate Or d ov i c i an , c on st i tu t i n g the Ci n c i n n at i an an d c on ta i n i n g fo
thro ugho ut Furt her discussion of the nature and subdivisions of Ci nci
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The env iro nme nt of Late Ordovici an t ime in the Ci nc in na ti region contri b
uted to the ab un da nc e and ri chne ss of fossils in several fu nd am en ta l ways.
Cincinnatian foss i ls and rocks bear profound test imony to the existence of
widespread shal low seas (cal led e p i c o n t i n e n t a l o r e p e i r i c se as) over m os t of
the North Ame ri ca n cont in en t at this tim e (Plate 1). Usin g ma ny sour ces of
evi den ce, geol ogist s have co mp il ed a reco rd of the rise an d fall of sea level
during the past half bill ion years of Earth history (f igure 1. 3). The Late Or
dovi cia n was one of the t imes of m a x i m u m rise of sea level over the entire
globe, r ivaled only by the Late Cretaceous (according to the reconstruct ion
by H a l l a m [1984]). The cau se of thi s f lo o d ing has b e e n attributed to h ig h ra te s
of sea f loor spr ea din g whi ch swel led the mid -oc ea n ridges , disp laci ng im
mense volumes of seawater f rom the deep ocean basins onto the continental
plates. The Atlantic O ce a n as we know it did not exist, but instea d, a narro wer
o cean ca l led t h e I ap et u s Oc e a n sep arated N o rt h A m e ric a f rom co nt inent al
plates later to const itute Eu ro pe and Africa (P late 1). T h e nearest la ndm ass es
to the Cin cin nat i region were the r is ing App al ach ia n moun ta in ch ai n, about
300 miles to the east, and the low-lying Canadian Shield to the north. Just
before and d u r i n g the La te Ordovician, a ph ase of m ajor t ec t o nic ( m o u n t a i n -
bui ld ing) acti vity, the T a c o n i c O r o g e n y , resulted in severe crustal de f o r m a
tion and uplift al on g the region bord eri ng Ne w York and N e w En gl an d. Is
lands were raised high above sea level as lofty and jagged mountain chains
Environment
We are accustomed to
thinking of North Amer
ica as terra firma, one
of the large high and dry
segments of the earth's
crust, and it is difficult for
us to imagine a time in
the past when our conti
nent was so submerged
beneath the sea that fish
could have swum directly
from the Atlantic Ocean
to the Pacific Ocean,
from Hudson Bay tothe Gulf of Mexico. Yet
such a time did exist 450
million years ago when
the epeiric sea spread
f A i G lf f
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Figure 1. 3. Global sea
level curves for the
Phanerozoic. A. Hallam
curve, B. Vall et al. curve
(1977). From Hallam
(1984) and reprinted by
permission of Annual
Reviews. According to
more recent studies
(Miller et al. 2006), maxi
mum rise of sea level in
the Cretaceous waslower than these esti
mates, reaching 100 m ±
50 m above present sea
level, but this does not
contradict the evidence
that Ordovician sea level
was also very high andextensive over North
America.
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Figur e 1. 4. Thickness of
Upper Ordovician strata
in relation to the ances
tral Appalachian Moun
tains (tectonic land) that
was uplifted during the
Late Ordovician Taconic
Orogeny. Contours are
lines of equal rock thick
ness (isopachs). From Kay
(1951, figure 4) and re
printed by permission of
the Geological Society of
America.
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preservation of hard parts is the Cam br ia n Burges s Shale of Bri t ish C
bi a, w i th its a m a z i n g w e a l t h of soft -bodied w o r m s , a r t h r o p o d s , an d
invertebrates, a l on g with shell -bea ring forms (C ou ld 1989). In the Ci
nat ian , there is virt ual ly no pres erva tion of soft- bodied spec ies or soft
of shell- or skel eton -bea ring species . T h e only records kn ow n to us o
b o d y pr ese r vat i on in t he C i n c i n n a t i a n are a w o r m de scr ib ed by U
(1878) and the re cen t dis cov ery of fossi lized "t ub e feet " in a brittl
(Glass 2006). Ou r kn ow le dg e of the Ci nc in na ti an biota is thus heav
ased i n favor of spe cie s wit h hard parts, th e shells and skeleto ns, com
or partial, known as body fossils. Fortunately, this is offset to some d
by e v i d e n c e of the activ i ty o f soft-bodied s pe c i es f rom t r a c e fossils
rows, tracks, and tra il s— th e subject of cha pter 14). H owev er, it mus t b
in m in d that pot enti all y great nu mb er s of spe cies in the biota will ne
known because they left no fossil record whatsoever.
Shells and skeletons preserved in Cincinnatian strata are pre
n an t ly c o mp os e d of c a l c i u m c ar b on a te ( C a C O 3 ) i n the mi n e r a l for m
cite . Some shells of brachiopods (see chapter 8) and the microfossi ls k
as conodonts (see chapter 13) are preserved as calc ium phosphate . D
the ab un da nc e of ca lc iu m carb ona te in Ci nc in na ti an fossi ls , not al l
having this chemical composition are equally well preserved. The r
for this is that so me org anis ms form ca lc iu m car bon ate shells or ske
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unequal . Species having shel ls formed of one or two valves (snails , c lams,
or brach iopods) h ave a high er prese rvati on potentia l than specie s with
multi-parted skeletons such as crinoids or trilobites. Multi-parted skeletons
are held together with connective t issue, which is susceptible to scavengingand decay, causing the skeleton to become disart iculated and scattered by
currents . The co ns eq ue nc e of al l these variable factors of shel l compo sit ion
and structure is that al l organisms producing a calcit ic shel l capable of
prese rvati on do not hav e an equal pot enti al for ac tu al prese rvat ion . Pres
erv at io n is h ig h ly s e lec t iv e ev en am o n g s h e l ls ch e m ic al l y and m ine ralo g i-
cally stable enough to survive post mortem.
The mode of l i fe of organisms determines preservation potential even
before a n i m a l s die. Fo r a qu at i c s p ec i es , b o t t o m - d w e l l e r s (bent ho s) h a v e a
highe r l ikel iho od of preservat ion than sw i mm in g (ne kto nic ) or f loat ing
(pl ank ton ic) species . A m o n g the ben tho s, spec ies that burrow into the
sediment for a l iv ing ( infauna) obviously have a much higher potential for
p res erv at io n t h an do s ur f ace dw el lers ( ep i f auna) . A m o ng t h e ep i f auna,
species l iv ing permanently attached to the bottom often have a higher
potential for preservation than free- l iv ing, mobile species , s imply because
they are una ble to esca pe sudden burial by se dim ent .
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that we can assess what species made u p the l i fe as se mb la ge . Th e de
as se mb la ge of rema ins already dead at the t ime of burial is a lso infor
tive, because, like a graveyard, it can record multiple generations and
cu rr en ce of rare specie s. Tab le 1 lists so me of the most useful charact eri
to look for in distinguishing fossils buried while living from those accu
lated gradually as dead remains.
Table 1. Characteristics of
Life Assemblages and
Death Assemblages
Life ass emb lag e Death asse mbla ge
Ar ti cu la ti on good disarticulated
Breakage rare common
Ab ra si on rare common
Preserved in life position maybe not often
Size-sorting uncommon possible
History
. . . our search for a
mechanism forces us to
f b d th
If, in light of the fo reg oin g dis cus sio n, the reader is not fully co nv in ce
the extr em e rarity of fossili zation and the uni qu ene ss of the fossil rich
of the Cincinnatian strata, the fol lowing section should provide additio
food for thought. Subsequent to l i fe and death during the Late Ordovi
P i d 0 il li th i f i i l b
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Figure 1. 5. Axis of the
Cincinnati Arch and its
branches, the Findlay
Arch (through Ohio) and
the Kankakee Arch
(through Indiana).
Shaded areas depict out
crop of Ordovician bed
rock; heavy lines indicate
Silurian-Devonian contac
that defines the Findlay
and Kankakee branches.
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SCIENCE IN THE HINTERLAND:
THE CINCINNATI SCHOOL
OF PALEONTOLOGY
The rocks beneath and around Cincinnati were deposited in an interval oftime universally called the Or do vi ci an Period . This tim e unit was prop osed
formal ly in 1879. In the sec ond h alf of the nine te ent h cen tur y, beg in ni ng
even before the Ordovician Period was named, there was in the region of
Cincinnati , Ohio, a group of paleontologists who have been cal led the "Cin
cinnati School of Paleontology. " There is no single, definitive list of the mem
ber s of t he C i n c i n n a t i S c h o o l , and diffe rent authors have i nc luded diffe rent
people as memb ers , dep en di ng on the purposes of their compi lat io ns. Nor is
there a definitive list of iron-clad criteria as to who should be considered a
me mb er and who should not. None thel ess, the indi vidu als inclu ded in the
bod ) of th is chapter have a n u m b e r of characterist ics in c o m m o n .
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ral Sciences that preceded the society). In present-day "buzz-word" te
n ology , the y c omp r i se d a " le ar n i n g c ommu n i ty . " T he y w or ke d toge t
the y shar e d r e sou r c e s ; the y c o mm u n i c at e d w i th on e an othe r ; the y e n c
aged one another; they competed against one another . Above al l ,
st imulated one another to perfor in at a higher level than they other
mi gh t have don e. The wh ol e was mor e than the su m of its parts. Th er e
tr u e syn e r g i sm i n the Ci n c i n n a t i S c ho ol o f Pa le on tology .
A l t h o u g h c a l l e d a s c h o o l , th e C i n c i n n a t i S c h o o l was not o n e , nor
it have any formal re lationship with any college or universi ty. (The Un
sity of Ci nc in na ti , as suc h, was not foun ded un ti l 1870, and there wa
De pa rt me nt of Ge o lo g y there unti l th e first de ca de of the twentie th
tu ry , w he n the D e p a r t me n t of G e ol og y an d G e og r a p h y w as i n i t ia te d.
But we need to put the Cin ci nn at i Sc ho ol into mo re of an histo
perspective . In the second de cad e of the nineteen th century, Cinc inn ati
the largest city west of the Al le gh en ie s, and a local physi cian, D ani el D r
f igured that the c i ty nee ded a f irst-c lass mu se um . H en ce , he spearh e
the esta bli shm ent of the Wes ter n M us e um . As part of the preparation
the ope ni ng of the ne w m us e u m , a taxiderm ist and artist na me d John J
A u d u b o n w as hired an d wo rke d for the or gan i zat io n for ab ou t a yea r, be
moving on eventually to become the most famous bird artist the Un
States has produced. In any case, the Western Museum opened in 1820
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Figur e 2. 2 A. Cov er of
an 7849 publication of
the Western Academy of
Natural Sciences published by U. P. James, a
member of the Cincinnati
School of Paleontology,
and his brother. B.
Cover of the Cincinnati
Quarterly Journal of
Science, volume 1, num
ber 7, published in Janu
ary, 1874, by S. A. Miller,
a member of the Cincin
nati School of Paleontol
ogy. C. Cover of the
Journal of the Cincin
nati Society of Natural
History, volume 1, num
ber 7. D. Cover of The
Paleontologist, Number
4, published in July 1879
by U. P. James, a member
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Figure 2 .6 . Urban out
crops in Cincinnati,
where members of the
Cincinnati School found
their inspiration. A. The
Bellevue House, on the
site of the present Belle
vue Hill Park, ca. 1895.
The stratigraphic section
exposed below begins in
the Kope Formation,
spans the entire Fairview
Formation and Miam-
itown Shale (a small
"step" below crest), and
is topped by the Bellevue
Limestone. Clifton Av
enue runs below the ex
posure, which was designated as the type section
of the Fairview Formation
by Ford (1967). (Image
courtesy of the Cincinnati
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sidered to be corals . Ma ny of the typ e-s pec ime ns in his col l ectio n en
up at the United States National Museum; other material went to the U
ve rsity o f C h i c a g o an d to th e Univer si ty of C i n c i n n a t i .
No t onl y was James th e autho r of ma ny paper s about lo cal fossils,
he was the publisher of many others. Indeed, James was the publishe
the journal The Paleontologist (Fi gu re 2.2), wh ic h ran for seven n um b
b e g i n n i n g in 18 79 . He also p u b l i s h e d a c a t a l o g u e of C i n c i n n a t i freshw
mussel s and ano the r of local p lants.
U. P. James retired from the bookstore business in 1886. He died
1889 and was buried in Cincinnati ' s beauti ful Spring Grove Cemete
(Becker 1938; Bradshaw, pers. comm.; Caster 1951,1981,1982; Croneis 19
Cuffey, Davis, and Utgaard 2002; Hendrickson 1947; Howe, Fisher ,
Ke ek el er 1889; J. F. Jame s 1889; Shi de le r [1952] 2002; A n o n . 1849,1878.)
Joseph Fra ncis James al mo st cer tai nly was indu cte d into the won der
fossil co ll ec ti ng by his father, U. P. Jam es (above) . However, Joseph's inests in natural history were broader than were his father's; the son p
lished not only about fossils, but about physical geology, botany, and ot
subjects.
Joseph F James began as a clerk in his father's bookstore but he
Joseph F. James
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Yo rk and L o n d o n d o i n g hospital work and ba cter io log ica l st udy, afte r w h i c h
he set up in medic al pract ice in Hi ng ha m, Massa chus etts .
Joseph K James was a prolif ic author , not onl y in pale ont olo gy, but also
in geol ogy and botany. Not co un ti ng many items in newsp aper s and mag a
zine s , h is outpu t am ou nt ed to wel l over on e hu nd re d scientif ic pape rs
about equal l y spread am on g those three areas , alo ng with a nu mb er of
others on mis cel lan eou s subjects . S om e of his pale onto logi c papers were
co-authored with his lather, U.P. James. The yo un ge r James was the aut hor
or co-au thor of a n um be r of taxa in the ty pe -C in ci nn at ia n, and at least on e
was n a m e d aft er h i m .
Joseph F. James died on March 29,1897, in Hingham, Massachusetts ,
and his ashes were buried in Cinc in na ti ' s Spr ing Gr ov e Ce me te ry . (Beck er
1938; Caster 19S2; Croneis 1963; Cuffey, Davis, and Utgaard 2002; Gilbert
1898; Shideler [1952] 2002; Anon. 1879,1882,1885b, 1886a.)
Charles Brian Dyer (Figure 2.1D) was born on April 1 , 1806, near Dudley C. B. Dyer
Cas tle , Worce stersh ire, England . Ha vi ng had to support himse lf and his
mother , he had l itt le formal edu ca tio n, i f any. He ca me to Cin ci nn at i in
1828 and set up as a ma nu fa ct ur er of soap and ca nd les . Ar ou nd 1850, ha vi ng
C. B. Dyer
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T h r o u g h his work with S. A. Mill er, C. B. Dye r was involved in
na m in g of many taxa of local fossi ls , inc lud ing annel id worm s, bryozo
snails , spo ng es , starfish and othe r ec hi no de rm s, trace fossils, and ot
Moreover, at least one genus and twelve species of fossils were named
hi m, inc lud ing the wel l-k now n species of cr inoids or iginally design
Glyptocrinus dyeri Me ek , 1872, now assigne d to Pycnocrinus.
A c c o r d i n g to records a t C i n c i n n a t i ' s S p r i n g G r o v e C e m e t e r y , C
Dyer died on July 11, 1883, in Harrison, Ohio, near Cincinnati. (Be
1938; Byrnes et al. 1883; Caster 1982; Croneis 1963; S. A. Miller and
1878a, 1878b; Raymond 1936; Sherborn 1940; Shideler [1952] 2002.)
Sa mu el A lm on d M ill er (F igu re 2.1B) is certai nly the most import ant o
"am ate urs " of the Ci nc in na ti Sc ho ol . He was born near Ath ens , Oh io
1837. By profession he was a lawyer; he had studied at the Cincinnati
Co ll eg e and was ad mi tt ed to the bar in 1860.
S. A. Mi ll er was also invo lved in pub lis hin g. In 1861-1862 he publ ithe M ar i e t ta , Ohi o , Republican, w hi c h, i n te r e st i n gly e n o u gh , w as a D e
crat ic new sp ap er. In 1874 and 1875, he was the propr ieto r of the Cinci
Quarterly journal of Science; many important papers on Cincinnatian fossils
bli h d i th t j l Aft t Mill ( d L
S. A. Miller
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tions of kno wl ed ge: The American Palaeozoic Fossils (1877), North Ameri
can Mesozoic and Cenozoic Geology and Palaeontology (1881), and North
American Geology and Palaeontology for the Use of Amateurs, Students, and
Scientists (1889, with supplements in 1892 a n d 1897). 'The last volume listed,
according to Kenneth Caster, is probably the most used volume about
A m e r i c a n p a le o n t o lo g y ever c o m p i l e d a n d ce r t a i n l y was the mo st a m b i
tious private publication in paleontology ever.
Mil ler 's compilatory works were looked down upon by most profess ion
als, but were used by them nonetheless. Caster recounted a story about his
professor, G. D. Harris, to the effect that Harris 's own professor, Henry
Shaler Wil l iams, was disdainful of Mil ler 's works. In Caster 's words: " 'Yet , '
said Harris , 'Mil ler' s grea t North American Geology and Paleontology was
alway s on Wi ll ia ms ' desk, and on th e desk of eve ry othe r pal eon tol ogi st of
the land!'" (Caster 1982, 24).
Nor did S. A. Miller confine his work to fossils from the Cincinnati
region. H e also worked on those of I l l inois , Misso uri , and Wisc on si n. Mil l
er's fossil collection must have been fantastic: one newspaper account re
ported that it contained over a mil l ion specimens! According to Bradshaw,
he rose early and worked on fossils until 10 AM, then went to his law office
until supper; after supper he worked a couple more hours on fossils.
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By now it has b ec o m e obv iou s that a num be r of threads of our s
arc intr icate ly inte rtwi ned. Variou s me mb er s of the Ci nci nn at i Sch
ha ve tie-ins with the Ci nc in na ti Societ y of Nat ura l Hist ory, with the U
ve rs it y of C i n c i n n a t i , or with b o t h. A n o t h e r thread in the skei n is W o
ward H i g h S c h o o l , as we shall see. But le t us fo ll ow the University of C
cinnati thread for a bit.
Albert G al l a t i n Wetherby ( f i g u r e 2.3A) was born in Pittsburgh, Pen nsy
nia, in 1833, but his family later moved to the Cleveland, Ohio, area. A
graduating from college, he spent several years teaching in a country sch
with s u m m e r s sp en t far min g. In 1861 he m o v e d to C i n c i n n a t i and was
pointed principal of Woodburn School, one of the public schools in the c
and spent some nine years there. In a eulogy written by George W. Harp
ano the r me mb er of the Ci nc in na ti Sc ho ol , it is reported that Wetherby
app oin ted professor of nat ural history at th e the n new Univer sity of Ci nc
nati in 1870 and stayed there six years. However, ac co rd in g to the Univerof Ci nc in na ti Recor d of Min ut es No . 2, a vo lu me in the archives of the U
ve rs it y of C i n c i n n a t i , Wethe rby ' s t im e at the univ er sit y b egan in the a u t u
of 1877, an d he is listed as "Ass't. Pr of A. C. Wet her by o f Na tu ral His tor y."
January of 1878 he was appointed "Curator of the Museum in the Universi
A. G. W e t h e r b y
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John M. Nickles s tudied un der Wethe rb y at the Uni vers ity of Ci nc in na ti ,
and Ge or ge W. Har per wr ote a eu log y abo ut Weth erby . Wet he rb y was one
of a comm it tee of ten who wrote a report on the geolo gica l no me nc la tu re
of the ty pe- Cin ci nn at ia n (S. A. Mil ler et al . 1879); six of the i ndivi duals on
t hat co m m it t ee are g eneral ly rec o g ni zed as m e m b ers o f t h e C in c in na t i
Sc ho ol, and all of th em ha ve bee n listed in one pla ce or an ot he r as co lle c
tors of loc al fossils. (Br and t and D a v i s 2007; C a s t e r 1982; H a r p e r 1902;
Johnson 2002; S. A. Miller et al. 1879; M i c k l e b o r o u g h a n d W e t h e r b y 1878a,
B; N ick les 1956; W e t h e r b y 1879a, 1879b, 1880, 1881; A n o n . 1876, 1878,
1879.)
Jo hn M ic k leb o ro u g h , Ph . D . , w as t h e p r inc ip al o f t h e C in c i nna t i N o r m al
School f rom 1878 until 1885. T hi s school wa s a part of the Cin ci nn at i pub -
l ie school system that was dedi cate d to trainin g teachers . Th e Ci nc in na ti
Board of Educa tion su spe nde d the opera t ion of the No rm al Sc ho ol in
1900, but that was a de ca de and a half after Dr. Mi ck le bo ro ug h had de
parted Ci nc in na ti for New York, wher e he be c a me the pri nci pal of the
Boys High School in Brooklyn.
M ick lebo ro ug h h ad been no m in at e d f or m em ber s h i p in t h e C i nc in
John
Mickleborough
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and as a co-author with S. A. Mil ler . As such, he was involved in the n
ing of a n um be r of taxa of foss i ls f rom the t yp e- Ci nc in na ti an .
A c c o r d i n g to t h e s a m e p h o t o g r a p h c a p t i o n , Faber lived unti l 1930 ,
en o u g h t h at Sh id e ler w en t co l lec t in g w it h h im . A c co rd in g t o Sh id
([1952] 2002, 3), Faber was, ". . . like the typical old timer he was[,] very
cret ive and suspi cious . H e wasn' t te l l in g an ybo dy any thi ng. I t took me
years to get h i m so ft ene d up a n d e d u c a t e d so that h e was w i l l i n g t o c o
out with his info rma tion . So we started goin g aro und to a nu mb er of
old secret localities where S. A. Miller got his types."
Lik e mos t of the other me mb er s of the Cin ci nn at i S ch oo l , Faber
associa ted with the Ci nc in na ti Society of Natu ral History . In fact , he
pro pos ed for me mb er sh ip in the society- in 1885, at the sa me time as Ch a
Sc h u ch e rt and Erns t Va up e l , and h e w as duly e lec t ed .
Faber sold his original col lect ion to the Univers ity of Chicago
$5000, ac co rd in g to Shid eler , and it includ ed spec im en s descr ibed by S
Mil ler . Be in g an inveterate foss i l col lec tor, howeve r, he pro cee ded to am
a second col le ct i on. This one was be qu ea th ed to the Univers ity of Ci nc
nati. The co ll ec ti on c a m e wit h so me mone y t o prov ide for a curato rial p
t ion and for pal eon tolo gic al p ubli cat i ons. The f irst holder of the curato
posit ion was Car rol l L an e Fen ton , who went on to write, alo ng with
w i fe w h a t is arguably th e best b o o k of its t i m e for a m a t e u r fo ss il col lec
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After q u i t t i n g s c ho o l , he was a sur veyor fo r a c o u p l e of years an d
worked on the E d e n Pa rk Reservoir, w h i c h , to thi s day, suppl ies d r i n k i n g
wa te r to d o w n t o w n C i n c i n n a t i . He was a s tud en t at B a l d w i n - W a l l a c e C o l
lege for two years, hut he did not finish college. During the 1876—1877
school year , he was a student in the Medi cal C ol le ge of O h i o in Ci nc in
nati, an independent institution at that time, but absorbed into the Univer
sity of Ci nc in na ti in 1915 (Bro addu s, pers. co mm .) . Ag ai n, he did not finish
work for a d e gr e e . Formal e d u c a t i o n an d he did not get on too we l l , b e
cause "he insisted he was taught too much he didn't want and too little that
he did" (Bassler 1945, 333).
In 1876, Ulri ch was ele cte d to me mb er sh ip in the Ci nc in na ti So cie ty
of Natu ral Hist ory. The fol low ing year he was ele cte d curator of pal eon tol
ogy, an unpaid position. A bou t that time the societ y acqu ired its ow n buil d
ing , and , in the mi nu te s of th e society- for the firs t me et in g held in t he new
b u i ld i n g on N o v e m b e r 6, 1877 ( C i n c i n n a t i S o c i e t y of Nat u r a l History), it
is recorded:
" Th e matter of appo inti ng a janitor for the Bui ldi ng co mi ng up, proposi
tions were recei ved from Mess rs. F. O. Ulrich , Talbot, an d J. C. Shor ten .
"Professor W ether by mo ve d that the Society pr oc eed to ball ot for a jani
for, the person elected to be subject to such rules as the Society may adopt,
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tions of bryozoans, which he sold to buyers both in the United States
Eur ope . In order to col lec t suff ic ient spe cim ens and mak e thin-sec
from th em , Ulr i ch emp lo ye d other local af ic ionad os of fossi ls, inc lu
Bass le r , N i c k le s , an d S e hu c he r t . (K e n n e th Caste r [1982] has credit e
rich with t he trait of enl ist ing the assi stanc e of local yo uth s, therein ch
ing their lives. This cal ls up the im ag e of the kindly old ma n hel pin
local kids; i t happens, though, that two of Ulr ich's three best-known p
ges, Sehuchert and Nickles, were only one and two years younger
Ulr ich, respectively.)
In 1897, Ulr ich was hired pe rm an en tl y by the United States Ge ol o
Su rve y and stayed the re for the rest of his care er, event ual ly b ec o m in
hea d of the strat igrap hic se cti on, and in effect the arbiter of strati gr
decisions in the country. Although specimens and thin-sections prov
by Ulrich are presen t in many insti tutions al l across the l an d , his per
collection went mainly to the United States Geological Survey and th
the United States National Museum. Ulrich officially retired in 1932
co nt in ue d scho larly work as an honorary assoc iate in pal eon tol og y aS mi thson i an In st i tu t i on .
Ul ric h autho red or co- aut hor ed many taxa of an im al s of all kinds.
ol these was a specie s of ost raco d cru st ace an n am ed alt er the ma n
cloth who had introduced him to fossils Ulrich wrote: "I name it after
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In that last year, James Hal l, the clean of Am er ic an paleo ntol o
vi sit ed C i n c i n n a t i a n d w as so i mp r es se d by S e h u c h e r t ' s c o l l e c t i o n th
hired hi m to b ec o m e an assistant for the N e w York Ge ol og ic al S urve y
inci dent ally , ob ta in ed his co ll ec ti on of fossils). In 1893, Se hu ch er t j
the United States Geological Survey, and, a year later , he went to
Un i te d S tate s N at i on al M u se u m, a lso i n W ashi n gton , D .C. Ev e n tu a l l
came to occupy the most prestigious geological professorship in N
A m e r i c a , that at Y a l e Universi ty.
S e hu c he r t b e g an a t te n d i n g me e t i n gs of the Ci n c i n n at i S oc i e
Natural History- in 1878. However, it was not until 1885, after he had le
furniture business for good, that he formally was proposed as a mem
A s i t h a p p e n s , C h a r l e s Faber , E rn s t V a u p e l , a n d S e h u c h e r t al l w er e n
nated at the same time.
Sehuchert was born in Cincinnati, but in some respects it is not f
cal l hi m a me mb er of the Cinc inn at i Sch ool , becaus e he did not real ly
lish anything locally. He had 234 scientific publications, but none app
in the local journals. Although he did not invent the paleogcographic
he bro ug ht it to its ma tu re state. A nd , up until the ti me of his death , in
he was also the foremost authority on fossil brac hio pod s of Nor th Am e
L i ke s ome othe r me m b e r s of the Ci n c i n n at i S c hool , S e hu c h e r
h d d i hi l i f t i H d d h t ' d
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By the ti me he gra du at ed from hi gh scho ol i n 1878, he ha d starte d a
b ib l iographic work o n the local b r y o z o a n s , and he a l rea dy w a s a c q u a i n t e d
personal ly with U lri ch and S chu che rt . After st ints of tea chi ng in Arka nsa s
and then Illinois, where he was a high school principal, he returned to
Cin cin nat i , al t houg h for a nu mb er of su mme rs he had spent vacat ions with
Ulrich col lect ing bryozoans al l over central and eastern North America.
In 1899 Ni ck le s me t Ray S. Bassler at the resi de nc e of E. O. U lr ic h in
Newport , Kentucky. Nickles and Bassler col laborated to produce United
States Ge ol og ic al Survey Bul let in 173— Syno ps i s of American Fo ss il Bryo-
zoa ( Nic kles and Bassler 1900). Ab ou t the sam e ti me , Josua Lin da hl , the
director of the Cin ci nn ati Soc iet y of Natu ral History and former state ge
ologist of Ill inois, asked Ni ckl es to pre par e a pap er on the geo lo gy of C i n
cinnati; this was published in the society's journal in 1902 and is used to
this day. In the su mm er of 1909, Nick le s pre par ed a manu sc ri pt ge ol og ic
m ap o f t h e W es t C in c in nat i Qu ad ra ng le f or t he p ro p o s ed C inc i nn at i F o l io
of the Unite d States Ge ol og ic al Surv ey; this has yet to be publis hed.
In 1903 Nick les was appo inte d to the Unit ed States Ge ol og ic al Sur vey
in Was hin gto n, D.C . , apparen tly on the strength of the bryozoan bibl iog
raphy that had appe are d in 1900; of cou rs e, his fri end shi p with Ul ri ch did
not hurt. Unti l his dea th in 1945, he dev ote d him se lf to co mp il in g bib liog
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organizing classes and in putting the institution in order; in fact,
co un te d as an int erim presid ent of the universit y (Gr ace an d Ha nd
139). Mo re ov er , he served on the boa rd and as president of the Co ll
Me di ci ne and Surg ery of the universi t y for a n um be r of years.
Harper was e lected to mem ber sh ip of the Cinc inn ati Society of N
History in 1871 (C in cin na ti S oci ety of Nat ura l History, 20). H is assoc
with the society w as lo n g and extensive. At various t imes he ser ve d as cu
librarian, me mb er of the publi shing co mm it te e , v ice president, and
dent. Mo reove r , he served as a me mb er of the co mm it te e on the nom
ture of the rocks of the ty pe- Cin cin nat ian ch aired by another me mb er
Ci nc in na ti Sch oo l and i ncl udi ng four others (S. A. M ill er et al. 1879).
W o o d w a r d Hi gh S c h o o l , in C i n c i n n a t i , was he aded by " kindly pr i
Ge o rg e W. Har per, a geol ogis t in his ow n right, whos e partic ular des
life was to train students of geology" (Bassler 1947, iv). For example, he
tated the progr ess of John M. Ni ck le s an d Ray S. Bassler by al lo wi ng
to re-arrange their schedules at the school so as to be able to work with
in paleontologic endeavors. Moreover , in 1896, he co-authored a pal
logical paper with Bassler, then a high school senior (Harper and B
1896). George W. Harper died in 1918. (Bassler 1947; Caster 1965,1982; C
Davis, and Utgaard 2002; Dury 1910; Harper 1886, 1902; Johnson
Kramer 1918; Martin 1900; Anon 1876 1878 1885b 1886a b )
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by Bass ler th at t he young m a n was a l o n e in the b u i l d i n g o n e day w h e n a
gentleman with a brill iant white beard showed up and asked to be shown
around the place. Bassler did so, and, afterwards, the bearded visitor de
parted on his way ba ck to Al ba ny , Ne w York. Thu s, Ba ssler me t Jam es Hall(1811-1898), perhaps the foremost paleontologist in the country (Yochelson,
pers . comm.).
Ulr ich le ft t h e C in c in na t i area fo r W a s h i ng t o n, D . C , in 1900, and
Bassler fol low ed in Ma rc h of 1901, w it hd ra wi ng from the Uni vers ity of
Cincinnati , before complet ing his senior year.
Bassler worked privately for Ulrich and went to school part-time at
Co lu mb ia n Univers ity (now Ge or ge Wa shi ng to n Univers ity) ; he was able
to transfer credits back to the Un iver sity of Ci nc in na ti a nd was aw ar de d a
bachelor 's de g r ee in June of 1902. About that same t ime, Bass ler began
w o rk in g for the Unit ed States N a t i o na l M u s e u m (where C h a r l e s S e h u c h e r t
was hi s i m m e d i a t e supervisor) . T h i s ass o c iat io n w it h the N a t i o n a l M u
seum lasted for nearly six decades, as Bassler rose through the ranks to
b e c o m e he ad curator of geology in 1929. Af ter his ret irement, in 1948, hecon ti nu ed as an honorary resea rch assoc iate until his dea th in 1961. M e a n
w h i le , he e a rn e d hi s master's and doct oral d e g r e e s at G e o r g e W a s h i n g t o n
University, in 1903 and 1905, respectively; thereafter he was associated with
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b en efi t of h i g h e r m a g n i f i c a t i o n (Caster 1965,1981). I f that we r e true
time, Bassler must have seen the light, at least with respect to bryo
". . . we ca n not be sure of the pos iti on of any form in the sc he me o
sification un til we have lea rn ed its inter nal stru ctur e by mea ns o
sections examined microscopically" (Nickles and Bassler 1900, 9).
over , according to Ell is Yochelson, Bassler had a compound microsco
his desk, and i t appears in photographs Yochelson had seen (Yoch
pers. comm.).
Bassler was recognized for his great accomplishments during hi
tim e. He was elect ed secr etary of the Pa leo nto log ica l Soc iet y and serv
that posi tio n from 1910 to 1931, an d then he b ec a m e pres iden t of th e sIn 1933, he was president of the G eo lo gi ca l Soc iet y of Am eri ca .
W h e n Ray S . Bassler d ied i n 19 61, t he C i n c i n n a t i S c h o o l o f Pale
og y was no mo re —e xc ep t in their vast num be rs of fossi ls in mu
around the world and in their publications in libraries. He was th
survivor. (Bassler 1933; Becker 1938; Brandt and Davis 2007; Caster
1981; Croneis 1963; Harper and Bassler 1896; Nickles 1936; Nickle
Bassler 1900; Shideler [1952] 2002.)
A l t h o g h a n m b e of th e m e m b e s of the C i n c i n n a t i S c h o o l of P aleThe Cincinnati
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NAMING AND CLASSIFYING
ORGANISMS 3 W h e n p e o p l e from a n u m b e r o f dif ferent c o u n t r i e s e n d e a v o r t o c o m m u n i
cate with one another, eventual ly there is a problem, namely , language.
Dif ferent peoples have dif ferent names for the same animal; for example,
"chat," "felix," "gato," "gatto," and "Katze" all refer to the animal we call
"cat." Moreover, the same word may be used to designate more than one
kind of an im al ; lor in sta nc e, we use the word "ca t" whe n ta lki ng abo ut a
house eat, or a lion, or a tiger, or a bobcat, or a mountain lion, or. . . .
Beg inning w el l o v er t w o cent ur ies ag o , i t g radual ly w as reco g nized
that , i f scientists arou nd the world were to co mm un i ca te with on e anoth er
successful ly , each kind of plant and an im al mus t hav e its ow n un iq ue
na me , and that eac h n a me mus t refer to on e, and on ly on e, kind of plant
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In some scientific works, you may see a scientific name followed
person's name, a comma, and a date, for example, Felis catus L i nnaeus,
This me an s that it was Ca ro lu s (or Ca rl) Linn aeus wh o nam ed the spec
th e tenth edit ion of his bo ok Systema Naturae, published in 1758. Lin
invented the binomial system of naming organisms, and he did so in
b ook. B e c a u s e of t he great sc op e and i m p o r t an c e of that work, i t is c o m
to abbreviate "Linnaeus, 1758" to "L." Hence, you might see Felis catus
(There has been some confusion as to Linnaeus's real name; see
[1992]. As was the custom in Sweden at the time, Carl Linnaeus's father,
originally was called Nils Ingermarsson, after his father, Ingermar. As a y
man, Nils intended to become a pastor, and, when he registered as a stuhe was required to give a family name, rather than just the patronymi
chose "Linnaeus," a Latin word referring to a lime tree—there was one
ing in the family garden. After Carl was famous and ennobled by the
he adopted the honorific form "von Linne." It is for this reason that the
"Li nn aeu s" som etim es is written "Li nn e" | Moo re, pers. comm.].)
A c t u a l l y , the r e g u l a r i z a t i o n o f b i o lo gi c a l n o m e n c l a t u r e (the s c
of n a m in g the group s into whi ch org ani sm s arc classified) was onl y o
the con tr ib utio ns of Li nn ae us . He also was the in ventor of the syste
use for that classific atio n. It works like this: ea ch basi c kind of or gan i
called a species Related species are joined together in a larger unit a g
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example, "family" is the level in the Linnaean hierarchy between "order"
and "genus," and the "Felidae" is the taxon at the family-level to which Felis
catus belongs.
All of th is i s w i t h in t he realm of t a x o n o m y — t h e s c i e n ce of a s s i g ni ngorganis ms to their proper biol ogica l grou ps. The word "ta xo no my " co me s
from two Greek words, taxis, m ea ni ng "arrang e m ent , " and norms, m e a n i n g
"law" or "science of" (Brown 1956); thus "taxonomy" literally means the "law
of arr ang eme nt" or the "science of arra ngeme nt"; al ternative n am es are clas
sif ic ation and syst ema tics . The real ch al le ng e of ta xon omy is, of cou rse ,
f iguri ng out the biologic al relationships of the organ ism s bei ng studied. After
that has been done, and only after that has been done, can the groups be
na me d in a truly me an in gfu l fashi on, at least, from a biologic al poin t of view.
Al t h o u g h s o m e sc ie nt ist s m i g h t separate " n o m e n c l a t u r e , " the n a m i n g of the
groups, from figuring out the biological relationships, the two activities are
irrevocably intertwined.
One purpose for the Linnaean hierarchy is to simplify the describing of
kin ds of org ani sms. Im ag in e if yo u had to des cr ib e a ho us e ca t COMPLETELY; it would ta ke reams of paper and m a ny m o nt h s of t ime. B ecau se of t he Linnaean
hierarchy, by saying "Felis catus," you convey to your listener all the character
istics of the species, gen us, family, and s o on , with out ha vi ng to use up paper
W h a t Funny Names! If o u ha e tried to r ap our to ng ue ar oun d th e scientific n am es of
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W h a t Funny Names!
. . . many learned words,
half-Greek, half-Latin,
and always difficult to
pronounce, many unpol
ished terms that would
scorch a poet's lips.
Jules Ve rn e, [1864]1992, Journey to the
Centre of the Earth, 4
If yo u have tried to wr ap your to ng ue ar oun d th e scientific n am es of
y o u c a n identi fy w it h Jul es Ve r n e 's on e - l i n e r a b o u t scientif ic term
th ou gh he was referr ing speci f ical ly to mine ral og ica l ones). Part
pro blem is that the na me s of fossi ls do not see m to ma ke sen se —t h
pear to be ran do m co mb in at io ns of letters. Yes, they have a uti l i
s ignif icance in denoting taxa, but the names generally have "real"
ings, too. However , those meanings generally have their roots in a
Latin or Greek or both, which is unfortunate for the vast majority
w h o are n o t s c h o o l e d i n these c lass ical l a n g u a g e s .
Some names are eponymous, that is , they arc derived from the
of peop le . Th er e is a genus of co m m o n articula te-br achio pods found t y p e - C i n c i n n a t i a n c a l l ed Rafinesquina after C. S. Ra fi nes qu e (1783-
a naturalist wh o tau ght at Trans ylva nia Co ll eg e in Lexin gt on, Ken
T h e c r i n o i d Pycnocrinus dyeri was na me d i n h ono r of local fossil col
C. B. Dyer (1806-1883). And so on.
Ot he r na me s derive from places . For exa mpl e, the edrioasteroid
cinnatidiscus and Isorophus cincinnatiensis were named after a city
o nc e up on a ti me , was the capital of the old Nor th wes t Territory.
Latin suffix "-ensis" or "-ense" denotes place or locality.)
Each genus has what is known as a type-species; this was desig
to represent the genus In some cases the trivial name indicates this s
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If you hear so me on e tryin g to talk ab out a subjec t, but that pers on r outi nely
stumbles over technical terms or mispronounces them, you are bound to
suspect that he or she does not know the subject very well. So how should
one pron ou nc e the nam es of ta xa —s o as not on ese lf to be lab ele d an
ignoramus?
Proper pron unc iat ion of words that are Lat in or Gr ee k de pe nd s on a
kno wle dge of those classical lang ua ges . Alas ! Very few peo pl e today have the
requisite knowledge. To make matters worse, even among those who might
lay a claim to being well-versed in Latin or Greek, not everyone agrees as to
what const itutes correct p ro n unc ia t io n. Fo r e x a m p l e , dev ot ees o f " C h u r c h
Lat in" and those of "Cl assi cal Lat in" do not sing the sam e Ch ri st ma s carols
the same way. There arc, howev er, som e rules of th um b:
1 . Unless you kno w oth erw ise , put the empha sis on the ant epe nul t i
mate syllable (that is, the syllable before the syllable before the last).
T h u s , B r a c h i o p o d a i s B r a c h i O p o d a .
2. C's and G's are ordinarily hard (as in "cat" and "gun," respectively).
3 . V's are pronounced as W's .
4. J's should sound like the Y in "your."
5. A "long i" in Latin is pronounced like a "long e" in English.
6 . The dip hth ong "ae " in Latin is pr on ou nc ed l ike a " l on g i" in Eng
Pronouncing
Those Lip-
Blistering Names
te r m nomen dubium refers to the n a m e of a taxo n tha t wa s so poo r
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te r m nomen dubium refers to the n a m e of a taxo n tha t wa s so poo r
scr ibed and otherwise documented that i t is not certain just what c
tutes the taxon and how to recognize i t . A dubious name, indeed.
A n o t h e r s u c h p h r a se is nomen nudum. W it h a term that li
me an s "naked n am e, " a bi t of ba ck gr ou nd is necessary. W h e n a spe
named, the author is expected to fol low certain conventions that have
agre ed to by the co mm un it y of the world 's biologists . Th e author
indicate that he or she is naming the species for the first time (G
2:19-20, not wit hst and ing ). Th e specie s mus t have a diag no sis ; thi
spec ial ki nd of des crip tio n that tells how indi vid ual s of the "ne w" s
dif fer from all othe r me mb er s of the genu s. A nd on e or more type-
me ns must be indica ted.
Type-specimens serve as the material on which the species is
T h e y shoul d be deposited in a bona fide m us e um so that scientists of
generations can study the exact specimens on which a given spec
b a se d . It u s e d to be c o m m o n to b ase a " n e w " sp ec ie s on a s ingle s p e c
No w, given the int rasp ecif ic vari atio n that has be en found to exist
a l l speci es, i t is mo re co m m o n to desi gna te a suite of typ e-s pec ime ns
a species is na me d. If a l l the typ e-s pec im ens arc cons idere d to be of
v a l u e as r e p r e se n t i n g the s p e c i e s , they arc sa id t o b e c o t y p e s . On the
ha nd if there is single t yp e- sp ec im en or onl y on e of a suite is cons
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considered to be a separate genus, and be called it Pycnocrinus. In subse
quen t years, experts on fossil cr inoi ds deci de d that G. dyeri actual ly is more
closely related to the type-species of Pycnocrinus than to that of Glyptocri-
nus, so the species named by Meek was re-assigned—from the latter to theformer genus. He nc e, i ts na me was ch an ge d to Pycnocrinus dyeri ( M e e k ,
1872)— the parentheses are shorthand to tell us that the species named by
Meek in 1872, with the specific name "dyeri," was later transferred to the
g enus Pycnocrinus. (Al tho ugh par ent hese s used i n that way ma y not appe ar
in some guide books for amateur fossil collectors, they can provide a valu
able hint to the paleontologist try ing to track down the nomenclatorial
history of a pa rti cul ar spec ies.)
On the other hand, sometimes the situation is the other way round. In
1935, Saburo Sh im izu and Tadah iro Obat a reco g nized a "new " g enus o f
fossi l cep ha lop ods and na me d it Orthonybyoceras. In 1942, R o us s eau
Flower, an eminent expert on foss i l cephalopods, recognized a "new" ge
nus and called it Treptoceras. Later workers , for exa mp le, Cu rt Tei che rt ,
on e of the most fa mou s paleont ologi sts of his clay and , it ha pp en s, an exp erton foss i l cephalopods, concluded that animals formerly recognized as be
longing in the two separate genera comprised a single taxon. In such cases,
biologists ap ply a c o n v e nt i o n ca l led p rio ri t y , viz., when there is an older
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ROCKS, FOSSILS, AND TIME
Fossils in ma ny col l ect i ons and mu se u m exhibits are of ten impre ssive for
f ine ly preserved detail and even be auty , bec au se the y hav e un de rg on e
pain stak ing preparat i on by whi ch every trace of the stony matr ix has bee n
removed. However, a fossil so isolated from its embedding matrix also loses
mu ch of i ts s ignif ic ance as a mea ns by wh ic h to unders tan d whe n and how-
it l ived. Only th ro ug h inv esti gati on of the fossil in the rock can we attain a
clear und erst and ing of the s igni f ica nce of the ab un da nt Or do vi ci an fossi ls
of the Cin ci nn at i Ar ch reg ion , or any fossils for that matt er. In this chap ter w e will exp lo re the na tu re of th e rocks in w h i c h C i n c i n n a t i a n fo ss il s are
found, the means by which they are subdivided, and the applicat ions of
this s tudy to understanding the environments in which they were formed
Era , and the Pal eoz oic Era is the earliest era of the Ph an er oz oi c Eo n (
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in g "t im e of rev eal ed life," for the ab un da nc e of fossils in thos e strata
oth er set of term s, time-s tratigr aphic units, applies to the actual rock
geolo gists assign to parti cula r intervals of the geol og ic tim e scale, an
rou ghl y equ iva len t to the divisions of co nt in uo us time , except that thema ny gaps in th e record of tim e that result from inc om ple te preservat
A c c o r d i n g to a f u n d a m e n t a l p r i n c i p l e of g e o l o gy , s u p e r p os i t i o n , the
rock layers in an undisturbed vertical sequence were deposited befor
ers lying above them. Thus, for the Cincinnati " layer cake," we know
layers ex po se d in the bed of the Oh i o Riv er for med befo re layers ex
higher along the hi l ls ides, but we do not know exactly how old the
are, or how much older are low er layers tha n h ig he r layers. We kn ow
that lower layers are relatively older than higher layers. Geologists
l ished th e re lat ive ag e se qu en ce of the majo r sedi men tary layers
Ear th ' s c r u st lon g b e for e an a lyt i c a l me t hod s (c hi e fly r ad i om e tr i c d
w e r e d e v e l o p e d to d e t e r m i n e the a b s o l u t e a g e o f rocks .
Roc ks that form ed du rin g a particular interval of geo log ic time are
Relative Age
Time Stratigraphic
T h e earliest studi es of the rock s and fossils of the Ci nc in na ti regi on date Early Studies
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T h e earliest studi es of the rock s and fossils of the Ci nc in na ti regi on date
to the early 1800s (Drake 1825; Riddell 1837; Locke 1838; Lyell 1845) and
coin cid e with the very be gi nn in gs of geo log y as a scien ce. These early
wor kers m a d e no a t t em p t t o s u b d i v i d e t he st ra ta a r o u n d C i n c i n n a t i , a nd
referred to them simply as the "Blue Miami Limestone" (Riddel l 1837) ,
"Blue L i m es t o n e" ( Lo c k e 18 38 ), o r "G rea t L im es t o ne D ep o s i t e" ( Br ig g s
1838). F. B. M e e k an d A. H . Wo rt he n (1865), bot h pion eer s of Am er i ca n
paleontology, f irst used the term Cincinnati Group for the Ohio strata.
Edward Orton (Orton 1873), Ohio 's third state geologist , proposed a four
fold subdivis ion of what he termed the Cincinnati beds proper: River
Quarry Beds ( lo w erm o s t ) , M iddle ( Eden) Sh ales , Hi l l Quarry Beds , and
Lebanon Beds (uppermost) .
The abu nda nt and wel l-pr eserve d foss ils of the Cin ci nn at i reg ion attracted
more and more attention du ri ng the latter part of the nine tee nt h cent ury ,
and as a result, deta iled kn ow le dg e of the distr ibu tion of par ti cul ar fossil
species in the strata was ac cu mu la te d thr oug h the ef forts of the Cin ci nn at i
scho ol of early pal eon tol ogi sts (see ch apt er 2). In 1902 John M. Ni ck le s
p ubl is h ed a co m p reh ens i v e rep o rt o n t h e g eo lo g y o f C i nc in na t i , an ex ce l
Early Studies
Subdivision Based
on Fossils
group ". . . may be divided into three subdivisions, more easily recog
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faunally than l i th ologi cal ly, tho ug h c lose study shows l i tholog ical
ences, which soon come to be fe lt , but are not easi ly descr ibed" (N
1902, 69) These were named the Lower Utica or Aspidopora new
Be d s , the M i d d le Ut i c a or Batostoma jamesi Beds, and the Upper Ut Dekayella ulrichi Bed s. The Lorr ai ne was l ikewise subd ivide d in
"beds," each given a local geographic name as well as a character istic
spe cie s, and he indi cat ed tha t th e gr ou p ". . . is easily separa ble on
gr ou n d s , w i th c or r e sp on d i n g mor e or le ss w e l l -mar ke d l i thologi c a l c
ters" (Nickles 1902, 75) He div ided the Richmond into lower, middle
upper divisions based on their faunas, but indicated that study has
insuff ic ient to establish their boundaries or l i thological characters.
In 1903 August F. Foerste introduced for the first time the term C
natian Series for the entire section, and referred to the Utica, Lorraine
Richmond as stages (Foerste 1903). Foerste (1906) termed the same
groups formations, with their subdivisions as members. He also disc
the New Yo rk ter m Lo rr ai ne and repla ced it wit h Mays vil le. In the
year R a y S. Bassler (1 90 6) e levated the n a r r o w e r su b di vi s ion s to fo rm
an d d e f i n e d the Cov i n gton Cr ou p to i n c lu d e the Ut i c a , Ed e n , Fa i
an d M c M i l l an For mat i on s , ov e r la i n b y the Ri c h mo n d Cr ou p , i n c l
the Ar n h e i m, W ayn e sv i l le , L i b e r ty , W hi te w a te r , an d S a lu d a For ma
Figure 4 2 Average per
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Figure 4.2. Average per
cent composition of the
allochem fraction of fau
nal groups and algae in
the upper (Saluda andWhitewater Formations)
and lower (Kope Forma
tion to Saluda Formation)
parts of the Cincinnatian
Series. (Allochems are
fragments of fossils or
other discrete grains in
limestones.) Adapted
from Martin (1975), cour
tesy of Wayne D. Martin.
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f raught with dif f icult ies Eve n the international ly rec ogn iz ed Cin ci nn at ia n
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f raught with dif f icult ies . Eve n the international ly rec ogn iz ed Cin ci nn at ia n
s t ages ( Ed eni an, M ay s v i l l i an , and R i ch m o n di an ) co u ld h e ca l l ed int o
quest ion as val id t ime-strat igr aphic units bec au se they wer e equ iva len t to
t h e Ede n, M ay s v i l l e , and R ich m o n d "C ro up s " w h ic h co m p ris e d " f o rm a
t ions" that lacked t ime-strat igraphic s ignif icance. Consequently , geologic-
research on t h e C i n c i n n a t i a n d u r i n g t h e 1960s resulted in major revisions
of the strat igraphic c lass if icat ion and n ome nc la tu re.
Three major research p rog ra ms of the 1960s ma rk ed a ne w ph as e in the
dev e lo p m ent and unders t a nding o f C i nc in na t i an s t rat ig rap h y ; a ll w e re in i
t iated outs ide of Ci nc in na ti . Severa l geologi sts at The O h i o State Univer
s ity (notably St ig M. Ber gstro m, Walter C. Sw eet , Ma lc ol m P. Weis s , and
their s tudents) publ ished a series of papers a im ed at revi s ing the Ame ri ca n
Up p er Ordo v ic i an St andard in bo t h l i t h o s t rat ig rap h ic and b io s t rat i g rap h y
terms. In the 1960s the Kentucky Geological Survey and the United States
Geological Survey init iated a major project to provide new geologic maps
of the entire s tate of Ke nt uc ky at scale of the 7.5 m in ut e qu ad ra ng le
(1:24,000 scale). In order to accomplish this enormous task, it was necessary
to provide a un ifor m strat igraphic c lass if icat ion of map pa bl e l i thostrat i
hi i k i i h h S i hi C d l i
Lithostratigraphy
and "Stateline
Boundaries"
lishe d for the Ci nc in na ti an sect ion in ea ch state of the tri-state region
t b l t d
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to be correlated.
A l t h o u g h th e ear ly z o n a t i o n of the C i n c i n n a t i a n based o n s u c h g r ou b r y o z o a n s an d b r a c h i o p o d s had p r ov e n i n a d e q u a t e for c o r r e l a t i o n ve
outside the local Cincinnati area, beginning in the late 1950s, new res
i n to the b i os tr a t i g r ap hy of the Ci n c i n n at i an e n ab le d a s i n gle t i me -s
graphic framework to be applied across the entire region. In order t
derstand how this new advance came about, we must consider what
acteristics are required for a fossil to be a reliable tool for long dis
cor rel ati on of strata.
Fossils that are mos t usefu l for cor rela tio n of strata over great dist
ideally must have two characteristics: a short vertical (time) range a
w i d e lateral d is tr i bu t ion . Fossil s h a v i n g a short vertical range wi l l be g
w h o s e p re se rv ab le m o r p h o l o g y ev olv es re latively rapidly over t im e. F
w i t h a w i d e lateral d is tr ib ut i on w i l l be g r o u p s that ei ther tolerate a
rang e of en vir on me nt s or e lse are cap abl e of wid e dispersal th roug h a
s wi m m in g larval stage or adult mo de of l i fe. Th e short vertical ran
many Cincinnatian fossi ls produced a detai led biostratigraphic subdiv
of the strata. H owe ver , as already men ti on ed , ma ny of the fossil zo n
New Advances inBiostratigraphy
Figure 4.4. Cincinnatian
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shoaling-upward cycles.
From Tobin (1982, figure
74). The base of the Cin
cinnatian is at the base ofthe Kope Formation in
the left-hand column; the
Corryville Formation con
tinues above the Bellevue
Limestone, and the
Waynesville Formation
overlies the Oregonia
Formation and continues
to the top of the Cincin
natian (Saluda Forma
tion). Arrows indicate
thickness of each major
cycle in meters. Note that
in each cycle, shale is
replaced by limestone
toward the top, indicat
ing a shallowing upward
transition. Rock symbols
Figure 4.5. Geological
Society of America Field
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Society of America Field
Trip, 1981. A. Left to
right: leader Rick C.
Tobin, participants Tim
Carr, Thomas W. Ams-
den, and Robert F.
Dill. B. Professor
Wayne A. Pry or with
poster illustrating Cincin
natian shoaling-upward
cycles and fades
relationships.
Figure 4.6. A. Three
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scales of Cincinnatian
stratigraphic cyclicity.
From Tobin (1982, figure
30). In storm cycle, basalbed with wavy and
oblique lines indicates
cross-stratification pro
duced by storm-induced
currents and
waves. B. Comparison
of three concepts of Cin
cinnatian meter-scale cy
cles. From Holland et al.
(1997, figure 1), courtesy
of the Journal of Geol
ogy. In each column,
width of bed pattern in
dicates lithology as
shown in scale (pack =
packstone, grain = grain-
stone). Arrows indicate
thickness of cycle in me
Figure 4.7. A. Storm
deposition and erosion
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p
across the Cincinnatian
sea floor. From Tobin
(1982, figure 37). Note
that hurricane rotation is
correct for the Southern
Hemisphere. B. Varia
tions in Cincinnatian
storm cycles. From Tobin
(1982, figure 36). Arrows
indicate layers of sedi
ment deposited by single
storm events. Sh = shale,
L = laminated unit, W =
whole fossil limestone, S
= siltstone, F = fragmen-
tal limestone, FL = fine
grained limestone, SL =
storm layer, Ls = any
limestone. Chondrites is
a trace fossil. A = abun
dant, C = common, R =
Figure 4.8. Colonization
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MATURE COMMUNITY
of a soft mud substratum
by Dalmanella (low di
versity) is followed by the
development of a shell pavement and succession
of erect bryozoans.
Higher diversity is
achieved in the mature
community by crinoids
growing on a firm sub
stratum. From Harris and
Martin (1979) and re
printed by permission of
the Society for Sedimen
tary Geology.
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scale of cycl ic ity the megacycle, and described it as a "f ining-upward" Figure 4.9. Kope cyclicity
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packet havi ng a basal co mp on en t of coar se-gr ain ed l im esto ne (grainstone) ,
a middle component with thin interbedded l imestones (packstones) and
shales , and an upper shale-rich component (Figures 4 .6A, B) . Jennette and
Pryor (1993) felt that the cycle was "coarsening-upward," beginning with
shale, overlain by interbedded thin l imestone and shale, and capped by
coarse-gr ained l im eston e (Figu re 4 .6B). In their interpretat ion the cycl e
was of a regressive na t ur e , c o r r e s p o n d i n g to short-term fluctuations of sea
level. More recent studies by Holland and others (1997) have continued to
recognize storm-related features within these meter-scale cycles , but note
that no single pattern of co ar sen in g or f in ing u pwa rd is domi na nt . T h e high
variability in cy c le ch aracter probably reflec ts a c o m p l e x interplay of se a
level ch an ge and fluctuations in storm intensity an d freque ncy. A remark
able result of all these stud ies is that desp ite the lon g-h eld v ie w that ind i
vidual str ata within the C i n c i n n a t i a n are not la te rally persistent, meter-sca le
cycles with in the Kop e For mati on can i nde ed be corr elate d over virtu ally
the entire outcr op area for tens of kilo mete rs (Br ett and Al ge o 2001b; Hol
land, Miller, and Meyer 2001; Webber 2002). Variations in proportions of
limestone to shale within the Kope and Fairview Formations define even
larger-scale cycl es, on the order of 10-2 0 m thick ("de came ter- sca le cycle s")
h bl l l l i l di ( i d
and Kope-Fairview for-
mational contact trace
able over broad area of
Cincinnati Arch. A. Geological Society of
America Field Trip, 1981.
Participants examining
meter-scale cyclicity in
the Kope Formation
along Kentucky Route
445 near the Ohio River,
Campbell County, Ken
tucky. B. Contact of
Kope Formation with
overlying Fairview Forma
tion (arrow) along Rad-
cliff Drive, Cincinnati,
Ohio. Photo by Paul E.
Potter.
In recent years several paleoecologic and taphonomic studies bro
renewed attention to thin fossil-rich horizons that are traceable over a
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renewed attention to thin, fossil rich horizons that are traceable over a
area. Frey (1987b) traced the nautiloid-rich Treptoceras d useri shale w
the W ayn e sv i l le For mat i on i n W ar r e n an d Cl i n to n Co u n t i e s , Oh i o ,
corre late d it with a lithologically equ iva le nt trilobite-ric h shale to the in Indian a. A str oph om eni d bra chi opo d shell pavem ent encrust ed by
sam e three edrioaste roid species occ urs in the upper Corryvi l le fo rm a
at Florence, Kentucky, and again 22 km to the north near the I-275 bel
nort hwes t of Ci nc in na ti (M ey er 1990). In the uppe r Kope fo rm at io
persistent bed of ca lc ar eo us siltstone replete with the U-s hap ed trace
Diplocraterion provided a key marker horizon for correlation from n
ern Kentucky to southwestern Ohio (Jennette and Pryor 1993). Sev
e xam p le s of p a le on t ologi c a l e v e n t hor i z on s i n the Ci n c i n n at i an ar e
cus sed in Brett and Baird (1997). T h e s e inc lu de: a trac eabl e storm hor
(Miller 1997); a one-meter thick interval character ized by the brachio
Onniella meeki traceable 135 km from Ohio into Indiana (Frey 1997b);
Richmondian faunal " invasion" (Holland 1997);and Isote lus-bearing sh
in the Wa yn es vi ll e for ma ti on that can be traced for at least 40 km a
south wester n O h io (Sc hu ma ch er and S hrak e 1997). Datti lo (1996) used
r e stri c te d oc c u r r e n c e of the b r ac h i op o d Heterorthina fairmountensis
da tu m for correl ation of the Mi am it ow n Shal e from southwestern Oh
may have resulted from earthquakes caused by tectonic activity in the rising
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A p p alach ia n m o u n t a i n s far to the east. U s i n g all of these ev ent h o r i zo ns ,
Brett and Algeo (2001a) were able to correlate decameter and meter-scale
cycles with in the K ope and Fair view F orma tio ns for dist anc es up to 80 km
alo ng the AA Highway (Ke ntu cky Route 9) in nor the rn Kent ucky .
To a certain degr ee, the most recent studies of Ci nc in na ti an stratigraphy
confirm and vindicate the heavy emphasis that was placed on fossil content
by generat ions of earli er wor ke rs in the C i n c i n n a t i a n . Clear ly , no study of
Ci nc in na ti an stratigraphy can afford to ignore the pletho ra of info rmat ion
offered by the abundance and diversity of fossils throughout the region. At
the time of this writing, efforts to devel op a hig hly detai led regi onal stratigraph ic framewor k for the Cin ci nn at ia n rely on a synthesis of all evi de nc e
available, including l ithology, sedimentology, and paleontology. Studies in
corpo rati ng this "total ev id en ce " app roa ch i ncl ude those of Dat til o (1996),
Holland (1993 Holland (1997) Holland. Mi ll er , an d Me ye r (2001), Bre tt an d
Al geo (2001b), We bber (2002), and M c L a u g h l i n and Bret t (2007).
Th ro ug h these studies , the recogn it ion that the Ci nc in na ti an is con
structed of a series of stratal "pa ck ag es " de lim it ed by rel atively short-term sea
level changes that mark sequence boundaries also to some extent revives the
old conc ept of the stratigraphic " layer cak e" that typified mu c h of the origi
l k h Ci i i ( l d ) Cl l h Ci i
enc as in g rocks on the geo log ic tim e scale. H owe ver, the rate of evolut ion
ch an ge varies greatly am o ng living orga nism s, and som e groups show mar
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g g y g g g , g p
ch an ge th rou gh a vertical successi on of strata while others do not; co
quently, different groups have different clocks. Evolutionary change
can no t provide a unif orm mea sur e of tim e by wh ich to estimate the abso
age of fossils (despite the mo de rn use of "m ol ec ul ar clo ck s" to set the a ge
c o m m o n ancesto rs of different livi ng groups).
A b s o lu te ag e d a t i n g of a n c i e n t rocks rel ies on the c o n st an t rate o
dioac tive decay of unsta ble isotopes of certai n e lem ent s con tai ned in m
erals fou nd in volc ani c and oth er type s of ig neo us rocks. Be ca use th
min era ls do not usua lly form in t he shells and skelet ons of org ani sms o
the sedi me nt s, direct dati ng of fossils and most sed im ent ary rocks is
difficult. At best, we can hope to find datable layers such as volcanic
b e d s or lav a flows inter bedded w ith fossil iferous st ra ta . It m a y then be
sible to state that a fossil-bearing layer lies either above or below a da
horizon, making the fossil either relatively younger or older. In the mar
sed ime nta ry record the best oppo rtun itie s for this met ho d of age deter
nation come from unique potassium-rich c lay beds known as K-bentoni
form ed t hro ug h che mi ca l a lteration of vol can ic ash fal ls into the
K-bentonites contain bioti tes and zircons that are datable using urani
lead and potassium-argon dating techniques.
f rom a dri l lcore in Seneca County, Ohio (Bergstrom and Mitchel l 1992),
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but it , too, has not b e e n dated .
By using radiometric dates from other regions to calibrate the time-
stratigrap hic record base d on range s of co no do nt s and grapt olite s, it is pos
sible to place further constrai nts on the Ci nc in na ti an time interval. The
b e g i n n i n g of the O r d o v i c i a n is now dated at a bo ut 488.3 +/-1.7 mi l l io n years
ago and its end at 443.7 +/-1.5 million years ago (International Commission
on Stratigraphy 2004). The tim e scale pro duc ed by the U N E S C O - sp on s or e d
I nt ernat io nal Geo lo g ica l C o rre lat io n Pro g ram m e ( W ebby , C o o p er , Berg
strom, and Paris 2004) places the be gi nn in g of the Cin ci nn at ia n at abo ut
452.5 mil lion years ago. How eve r, the top of the sectio n i n the Cin ci nn at i A rc h region does not include the u p p er m o s t O r d o v i c i a n record b e c a u s e of
pre-Silurian erosion, and an entire stage, known as the Hirnantian, is miss
ing. Acc ord in g to the timesca le provi ded by Web by, Co op er , Bergstr om, and
Paris (2004) the base of the H ir na nt ia n is at ab ou t 445 mil li on year s ago .
These considerations yield an approximate duration for the Edenian,
Maysv illia n, and Ri ch mo nd ia n of abo ut 7.5 mil lio n years (452.5 my—445 my).
The fact that dates for neithe r the lower or upper bo und ari es of the Ci nc in
natian come from samples in the local region should alert the reader for
future refinements! However, the dates and 7.5 million year estimate seem
b h b i f i l il bl
b e d d i n g c on tac t it se lf potent ial ly rep res ents a bre ak in sed ime ntat ion or
tus of indeterm inate dur ati on— day s, mon ths , years, or more. Man y Ci n
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natian limestones have irregular, pitted upper surfaces, sometimes bea
enc rus ti ng org anis ms, which strongly suggest formation of a hardg roun
the sea floor over a long time interval during which very little sedimentcumulated. In general , given the abundance of ev idence for storm-rela
deposition in the Cincinnatian (Tobin 1982; Jennette and Pryor 1993). m
limestones probably represent more time than the interbedded shales,
det erm ina tio n of absolute duration s is very uncert ain .
W h e r e v e r sedi mentary strata display a strongly cycl ic pa tt er n of rep e
sets of beds ha vin g unifo rm t hickne ss or variation in lithol ogies, geol og
have sought a possible link to cyclic or episodic causes reflecting seaso
an nu al , or lon ger timesc ales of periodicity. M os t intri guin g is the possib
that variations in the Earth's orbital parameters could exert influence
climate changes that in turn cause cyclic or periodic sedimentation
cesses (Gr otz ing er et al. 2007). If sedi men tat ion co ul d be sho wn to resp
to this kind of astrono mical met ron om e, interpolation bet ween calibra
points on the tim esca le coul d be ma de accurately, and the am ou nt of t
represented by pa rticu lar sets of strata cou ld be det erm ine d. However, cy
of this type, kno wn as Mi la nk ov it ch cycl es, have not as yet been dem
strated to exist wit hi n the cyclic C in ci nn at ia n strata
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ALGAE: THE BASE OF
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THE FOOD CHAIN
5 A l g a e are u n c o m m o n fo ss il s in the C i n c i n n a t i a n , bu t are p o t en ti a l ly si g
nif icant as prima ry prod ucer s of the Or do vi ci an eco sy st em and as indic a
tors of impo rtan t env ir on men ta l con dit ion s suc h as water dep th. Alg ae
inclu de s ingle-cel led as wel l as mult i -cel led plants that are con f in ed to
aquatic or moist habitats because they lack internal canals for water storage
and transport . They arc therefore termed nonvascular. Because they obtain
their energy through photosynthesis , algae require adequate exposure to
sunligh t . This essential requ ire men t general ly restricts th em to very shal
low water. Alg ae inc lude the blue -gre en algae (divis ion Cy an op hy ta ) , green
algae (divis ion Ch lo ro ph yt a, s ing le- and mult i -ce l le d) , red alg ae (divis ion
Rhodophyta) , and the brown algae (divis ion Phaeophyta) .
Figure 5.1. A - C . Cincin
natian acritarchs, all Ede-nian, Kope Formation,
Wayne County, Indi
ana. A. Veryhachium
edenense Colbath, x
569 (from Colbath [1979,
plate 13, figure 1]).
B. Ordovicium gracileColbath, x 507 (from Col
bath [1979, plate 8, fig
ure 4]). C. Multipli-
clocrinites darwinii
(Miller), Stephen Felton
ll ti M illi
group , the Dasyc lad acea e, is represented in the Ci nci nna tia n by several
cies (Cross et al. 1996). Modern dasyclads occur no deeper than about 30
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collection, Maysvillian,
Mt. Auburn Formation,
Butler County, Ohio, x
1.3. I. C. darwinii, sur
face detail of H, polygo
nal facet diameter ~0.4
mm.
and frequently less than 5 m (Wray 1977). Cyclocrinites is the largest
most common dasyclad, found in the Bellevue Limestone and through
the Ri ch mo nd Cr o u p (Fi gur es 5.1H, I). This alga resemble s a golf bal
size, shape, and its dimpled surface. In well-preserved specimens the d
pled surfa ce reveals a pattern of rho mbo ida l plates. Th es e plates arc actu
ex pa nd ed ends of br an ch es that radiate from a cent ral axis. Sever al spe
w e r e or ig in al ly de scr i be d u n d e r the n a m e Pasceolus, but Nite cki (1970)
ferred mo st of thes e to a sing le spec ies , Cyclocrinites darwini (Miller).
Th re e other genera of dasyclad algae occ ur in the Cinc inn at ia n (C
et al. 1996 ). Lepidolites dickhautii Ulrich has a sausage-like shape, abou
cm long, but is usually flattened (Figure 5.1G). Its surface has a scaly
pea ran ce, and i t oc cur s in the Kope Fo rmat ion . Anomaloides reticu
Ulrich is reported from the lower Fairview Formation and has a sim
scaly surf ace. H owe ver , it is cl ub- sh ap ed, and can reach several cent ime
i n le n gth. Ischadites circularis (Emmons) has been reported from
Fairview, Corryvi l le , and Mt. Auburn Formations (Halve 1948) but i ts tnomic status has not been recently reviewed.
One red alga, Solenopora richmondensis (Mil ler) , occurs in the Wh
w ater an d El k h o r n Fo r m a t i o n s of the up p e r m o s t Ri c h m o n d G r o u p (B lack-we
spherical in shape, less than 100 microns in diameter (micron = millionth of
a meter), and have proje cting spines that often branc h (Fig ures 5 . 1A -C ) .
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a meter), and have proje cting spines that often branc h (Fig ures 5 . 1A C ) .
Al thou gh acri tarch s are similar to the resting stages of the d i n o f l ag e l l a t e s
(division Pyrrophy ta), one of the major com po ne nt s of the mo de rn ma rin e
phytoplankton, they cann ot be diagno stical ly related to any particu lar phy-
toplankton group. Acri tarchs occur throughout the Cincinnatian and their
diversity is considerable: fifty-two species were recorded by Colbath (1979)
in a cor e from the Ko pe Fo rma ti on of Ind ian a; Jaco bso n (1978) reported
forty-four species from the Ci nc in na ti region ; and L oe bl ich and Ta ppa n
(1978) describ ed forty new specie s from the Cin ci nn at ia n of O hi o, India na,
and Kentucky. Jacobson (1979) found that fluctuations in ab un da nc e of dif
ferent acritarch species corresponded to paleoenvironmental variations in
relative water depth.
Despite the name , chit ino zoa ns are neither com po se d of the protein
chitin, nor do they def inite ly represent an ani ma l group. T h e actua l co m
position of the minu te (50 -20 00 micro ns) , bot tle -sh ape d test is simi lar to
chitin but is termed a pseudochitin. Chitinozoans are thought most l ikely
to be some type of phytoplankton, but their exact biological affinities are
unres olved (Jansonius and Jenkins 1978; Cros s et al. 1996). Li ke acri tarc hs,
chitinozoans are found throughout the Cincinnatian (Miller 1976; Jacob-
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PORIFERANS AND CNIDARIANS:
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SPONGES, CORALS, AND JELLYFISH
A l t h o u g h s p o n g e s are re g ar ded as t he least s p e c i a l i z e d , h e n c e m o s t p r i m i
t ive of mult icel led a nim al s , they play an essenti al role as "sanitary engi
neers" in aquatic environments , l iv ing as act ive suspension feeders or f i l ter
feeders (Plate 3A). By removing minute organic part ic les f rom the water,
sponges prevent dec ay prod ucts f rom po iso ni ng the en vi ro nme nt . This is
a long-running role, as sponges f irst appear in the fossil record during the
late Precambrian, over 540 mil l ion years ago.
T h e body of a spo nge lacks d ist inc t cel l layers , bu t is co mp os ed of dif
ferent spec ial ize d types of cel ls that perf orm dif fer ent li fe functi ons. T h e
fundamental sponge cel l is the col lar cel l , equipped with a waving f lagel-
lum that draws water into a con e form ed of micr ovil l i ( Fig ur e 6 .1) . T h e
Sponges
Figure 6.2. Reconstruc
tion of a living stromato-
poroid modeled on a
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poroid, modeled on a
living sclerosponge. A
section is removed to
show internal laminae of
the skeleton. Living tissue
occupies only the upper
most layer, with excur-
rent canals radiating from
oscula. Magnified inset
shows surface tissue and
microstructure of laminae. Compare to Figure
6.3D, below. Drawing by
John Agnew.
w h i c h typical s p o n g e cel ls carry out f i l ter f e e d i n g ( f i g u r e 6.2). Wastew
canals that converge on the oscula leave starburst patterns of grooves incalcareous skeleton that match with structures called astrorhizae in the fo
group kno wn as st ro mat opo roi ds (f ig ure s 6.2, 6.3A, D). These and o
similarities enabled the stromatoporoids to be recognized correctly as a
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mouth
droids, sea fans, sea whips, and soft corals that are often mistakenly thou
to be seaw eed s. Des pit e suc h a be wi ld er in g array of forms , cni dar ian s sh
l f t th t i di t th i l ti hi b f
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corallum
Figure 6.4. Reconstruc
tion of the Cincinnatian
solitary rugose coral,
Grewingkia canaden
sis, showing the polyp
with extended tentacles.
Drawing by Kevina
Vulinec.
several features that indi cate their c o m m o n relati onshi p as me mb er s of
of the simplest mul ti - ce l l ed an im al (me tazo an) phyla. Cnid ari an s st
apart from other animals in having two body forms, the polyp and
me dus a. The po ly p, as typified by a sea an em o n e or stony cora l (Plate
Fi gu r e 6.4), is cylindrical in shape and attached at the base to a hard s
stra tum . The body w all consis ts of on ly tw o cell layers (unlike the th
layers found in all other metazoans), separated by a non-cellular jelly la
called the mesoglea. There is a s ingle opening (mouth) into the body cav
th ro ug h wh ic h food is ing ested and waste is exp ell ed. A ring of tenta
surrounding the mouth serves for food capture and defense. The medu
or jellyfish has the same structure as the polyp (with thicker mesoglea)
is free-l iv ing, swimming by muscular pulsations with the mouth or ien
downwards. Both polyp and medusa forms arc present at different sta
du rin g the li fe cycle of so me cni dari an species.
T h e three m ajor classes of cni dar ian s differ in their expressio n of
p ol yp an d me d u s a s tage s . Hyd r oz oan s ( i n c l u d i n g hyd r oi d s , Por tu gu e se M an(true jellyfish) restrict the polyp to a larval stage and live mostly as me
sae. Anthozoans (anemones, corals , "soft corals") l ive exclusively as poly
and of ten as col on ies of mul tip le po lyp s that are gen eti c clon es of a sin
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A l t h o u g h the so ft p o l y p of r ug o s a n s is n e v e r p r es erv e d, r ug o s a n s are
known to be corals because the cal ice has mult iple radiat ing part it ions
Figure 6.6. Cincinnatian
colonial corals. A -E .
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cal led septa that are foun d in li vin g cora ls. Sep ta are secrete d by soft tissue
part it ions of the internal bod y cavity cal led m esen teri es . In l i v ing an th oz o-
ans, mesenteries serve important functions in digest ion and reproduction.
The nu mb er and arr an ge me nt of the septa are traits use d in the classif ica
tion of coral s. In rug osa ns the septa hav e a roug hl y four-fold sy mm et ry ,
compared to l iv ing corals that have s ix-fold symmetry.
Tw o sp ec ies o f s ol it ary rug o s ans o cc ur co m m o nl y in t h e C inc in na t ia n,
bo t h in t he R i c h m o n d i a n st ra ta ( F i g u r e 6. 5; El ia s 1982, 19 98 ). Grewingkia
canadensis (Bil l ings) is the largest and most common rugosan (Figures6.5D, E). Co ra ll a reach le ngt hs over 13 cm (5 in) bu t are gen er all y in the
range 10-60 mm (0.5-2 in); the diameter ranges from 22 to 40 mm (0.9-1,6
in) . Specimens are almost always found ly ing on their s ides and appear
h ig h ly abraded, encrus t ed , and bo red ( F ig ur e 6 . 5E) . Ex t ernal c o nc ent r ic
gro wth lines are rarely pre ser ved (Fig ur e 6. 5D , inset). Like so me liv ing
sol itary corals , Grewingkia p robab ly l ived uprig ht , part ly bur ied in sof t
s edim ent w it h t h e p o ly p ex p o s ed. So m e encrus t at io n and bo r ing t o o k
place during l i fe but continued af ter the coral was exhumed by storm act iv
ity and deposi ted on its side. B ry oz oa ns are the most c o m m o n enc rus ter s
Corallites as seen on ex
ternal surface or in cross
sections of corallum, atsame scale x 3.7. A - E
from Elias (1998). A.
Cyathophylloides, a co
lonial rugosan, Richmon
dian. B. Foerstephyl-
lum, a tabulate,
Richmondian. C. Cala-
poecia, a tabulate, Rich
mondian. D. Nycto-
pora, a tabulate,
Richmondian. E. Tetra-
dium, a tabulate, Rich
mondian. F. Coral bed,
Richmondian, Madison
County, Kentucky, length
of hammer 25 cm (from
Elias [1998, figure 5]).
G Protaraea richmon-
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to question whether tabulates were in fact corals. An extraordinary discov
ery of soft tissue pol yps preserved in a Silu rian tabul ate (C op pe r 1985)
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sett led the debate for most tabulates, a lthough some, l ike the Cincinnatian
Tetradium, are very simil ar to so me livin g spo nge s that buil d a ca lc ar eo us
skeleton with a tabulate structure.
Colonial corals occur exclusively in the Richmondian Waynesvi l le , Lib-
ertv, Whitewater, Saluda, and Elkhorn Formations. Within these formations,
there are as many as tour distinct horizons where colonial corals are concen
trated into "coral beds" up to about 4 m (12 ft) thick, traceable for great dis
tances along the outcrop belt of the Richmondian around the Cincinnati
Ar ch (Figure 6.6 F; Br o w n e 1964, 19 65 ; Ha tf ie ld 1968; Elias 19 98 ). F o u r
genera of massive coloni al tabulat es (Foerstephyllum, Calapoecia, Nyctopora,
an d Tetradium) and one colonial rugosan (Cyathophylloides) are fou nd in
these beds (Figures 6.6A-F). Another tabulate , Protaraea, occurs in the
Richmondian but did not form massive colonies. Instead, Protaraea e xc lu
sively encrust s the shells of bra chi opo ds and other objects ( Fig ure 6. 6G). In
John Paul Park, in Madison, Indiana, there is a unique, octagonal tool house
bu il t en tire ly of colonial corals gathered fro m the coral b ed exp ose d north
of the town (Fi gure 6.6H). Co lo ni al corals are also incor porat ed into stone
wa lls beside s o m e of the e legant houses in M a d i s o n .
Figure 6.8. Conulariid,
Conularia formosa
Miller and Dyer, Univer
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sity of Cincinnati collec
tions, Maysvillian, Cor
ryville Formation, ButlerCounty, Ohio, x 7.2.
is com par abl e to that of ma ny livi ng coral s in patch reefs. Hatfield (1968)
showed that the coral zone within the Saluda Formation acted as a low,
l b i di l l h fi i d b
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coral barrier su rro un din g a centr al lag oon where fine-grain ed ca rbo nat e
sediments accumulated. In this way the coral zone acted as do present-day
reefs to inf luence water mo vem ent and sedi men t deposit ion. Applic atio n
of the terms patch reefs and b io st ro me s to the Ric hm on di an coral beds is
therefore quite reasonable.
The inability of Ci nc in na ti an cor als and oth er pote ntia l reef buil ders
to construct major reefs has several possible explanations. First, corals were
present in the region only dur ing Ric hm on di an t ime ( We bb y 2002). T h e
absenc e of corals duri ng Ed enia n and May svi l l ia n t im e is pu zz li ng , be
cause of the similarity of the rest of the fauna th ro ug ho ut the Ci nc in na ti an .
Environmental conditions in the Cincinnati Arch region may have been
unsui table for solitary and coloni al corals du ri ng the Edenia n and Mavs vil
lian, but it is difficult to identify the factors responsible. Abundance of
fine-g rained sedi men ts and fre quent dis tu rba nce of the sea floor by stor ms
are two factors that mi gh t have restricted the pr es enc e of corals . Ho wev er,
b oth fa ct ors are pervasive t h r o u g h o u t the C i n c i n n a t i a n , an d i t is n ot cer
tain that e i ther decreased signif icantly during the Richmondian. Accord
ing to Elias (1982) solitary corals were introduced during an early Rich
T h e zoo log ica l af f init ies of con ula r i id s have been debat ed for
t im e. C hi ef l y on the basis of their four-part struct ure, conu lar i id
b e e n classif ied w i th the s c y p h o z o a n c n i d a r i a n s , w h i c h h ave a tetr
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(four-fold) body plan. There are l iv ing scyphozoans with a chit inous
and so me that live at ta ch ed by me an s of a stalk. So m e have arg ue
c on u la r i i d s shou ld b e r e c og n i z e d as a d i s t i n c t p hy lu m ( Bab c oc k
Ba bc oc k and F el dm an n 1986), but recen t work by Va n Iten and
(1996), and Hughes and others (2000), confirms that the similar i t i
tw ee n conula r i id s and scy pho zo an s are indicative of a c lose evolut
relations hip. Con ul ar i i ds arc found in mari ne strata of Ca mb ri an th
Triassic age . A sing le spe cie s, Conularia formosa M il le r and D ye r,
c or d e d f r om the Ci n c i n n at i an of the Ci n c i n n a t i Ar c h r e gi on .
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BRYOZOANS: "TWIGS"
AND "BONES"
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AND BONES
7The rocks in the Cincinnati region are loaded with fossils. Visitors to the area
commonly are struck by all the "tilings" in the rock that look like small twigs,or, with a stretch of the ima gi na ti on , smal l piec es of bon es (Fi gu re 7.1A). T h e y
are the most c o m m o n fossils in the bed ro ck of the area. In de ed , if yo u were
to pick up a fossil in the Cincinnati region at random, chances are that it
would be one of the se objects. But th ey are nei ther twigs nor bo nes . T h e y are,
in fact, the remains of a gro up of orga nism s called bry ozo ans (Plates 3D , E ) .
If you look at an un br ok en sur fac e of you r br yo zo an fossil with yo ur
trust} han d-l ens , you see that it is replete with tiny holes ( f i g ur e 7.1 B). If
you sh ift your f ield of v i ew to a bro ken su r f ace , th e tiny h o les are r ev ealed
to be minute tubes . Each one of those minute tubes was once home to an
Figur e 7.1. A. Frag
ments of bryozoan colo
nies are the most abun
dant fossils in the
type-Cincinnatian. Par-
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Bryozoans arc aquatic. Depending on the kind, some bryozoans live in
fresh water, but most live in salt water. In either case, the bryozoan subsists
on minu te orga nisms (protozo ans and so on) and tiny bits of orga nic matter
Figure 7.3. Cincinnatian
bryozoans. A. Colonies
of the bryozoan genus
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on minu te orga nisms (protozo ans and so on) and tiny bits of orga nic matter
susp end ed in the water. The bry oz oa n do es no t just wai t passively for su ch
food to fall into its mouth; it literally filters the food from the water. Bryozo
ans are acti ve filt er feeders . The indi vid ual an im al spreads the tent acle s of
its lophophore into a funnel-, bowl-, or vase-like configuration (Plate 3E;
Figure 7.2), and cilia that line the tentacles move in such a way that food
particles are carried do wn to the mout h. N ot onl y doe s the indi vidua l pol yp-
ide generate feeding currents for itself, but the colonies of at least some kinds
of bryozo ans generate currents that enh an ce fee din g in the colon y as a whole .
In at least so me kinds of br yo zo an s there are part ic ula r areas on t he col on y
that have polypides that direct currents away from the colony. These excur-
rent chimney's (Plate 3F) carry water that already has been filtered by the
lophophores away from the zoarium, and thereby draw unfiltered, nutrient-
laden water across polypides elsewhere in the colony (McKinney and Jackson
1989). Moreover, the very topology of the colony may facilitate the passage
of nutrient-filled water thr ou gh the colony and across its pol ypi de -li ne d, an d,
hence, lophophore- l ined, and, hence, food-gathering surfaces .
Fossil b ryozoan col oni cs co m e in a wid e variety of sizes an d shap es
f y g
Spatiopora characteristi
cally form a thin coating
on shells of orthoconic
cephalopods, MUGM
uncatalogued, Cincinna
tian, scale in mm. Note
that raised lumps on col
ony surface (monticules)
are elongated and
aligned with the axis of
host nautiloid shell. B,
C. Ctenostome bryo
zoan, Ropalonaria
venosa Ulrich. B. Col
ony on shell of brachio
pod Rafinesquina, CMC
IP 40061, Waynesville Formation, Butler Co.,
Ohio, x 9. Arrow indi
cates sac zooid. C.
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istics of on e "spe cies ," or eve n "ge nus ," give w ay to t hose of ano th er —a ll in
the space of a centimeter or two. Presumably, everyone in a single colony
is of the sam e specie s. Thus, the co nc lu si on is inevi tabl e: overall col ony
that has grown on the
pedicle valve of the bra
chiopod Rafinesquina
d it Ri h
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shape and details on its surface may not always be reliable indicators as to
w h o is re lat ed to w h o m . I n d e e d , there is a m p l e e v i d e n c e pr ovi d ed by pres
ent-day bryozoans that environment can play a significant role in colony
shape, at least in some taxa.
W e l l , i f the shap e of the z o a r i u m is not an i n co n tro ve r t i b le taxonomic
indicator, what, it anything, is? Is there an "inner truth" in bryozoan tax
onomy? It turns out that there is just that, namely, the internal structure of
the colony. Eac h zo ar iu m consists of the hard parts of all of the ani ma ls
that compr ise the colony. An individual bryo zoa n anim al is cal led a zooi d,
and the haul parts of that individual an ima l consti tute a zo oe ci um ( Fig ure
7.2). If one exam in es the hol es in a zoa ri um wit h a hand- lens or low -po wer
microscope, one commonly sees that the holes are not identical (Figure
7.1B). Th er e may be size classe s of larger holes and sma ll er holes ; there ma y
even be spine-like proje ctio ns in addi tio n to hol es. It wo ul d app ear that, in
at least so me colo nie s, not all the zooid s were ident ical . In so me present-d ay c olon i a l an i mals , the r e is p o ly mo r p hi sm , w i th d i f fe r e n t-shap ed or
different-sized individuals performing different tasks, for the good of the
colony the species or both That seems to have been the case among at
and overgrown it, Rich
ard Arnold Davis collec
tion, Bellevue Limestone,Cincinnati, Ohio, scale in
mm. G. Trepostome
bryozoan colony that has
grown on shell of pelecy-
pod Ambonychia and
overgrown it, Richard
Arnold Davis collection,Cincinnatian, horizon and
locality unknown, scale
in mm. H. Ring-shaped
bryozoan zoarium ("We-
ichold donut") presum
ably encrusting nautiloid,
CMC IP 51109, Richmon
dian, Hamilton Co., Ohio,
x 0.7. I. Cross-section
of ring-shaped bryozoan
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Figure 7.5. Reconstruc
tion of the lower part of
the zoarium of Het-
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erotrypa sp., diameter
about 28 cm. From
Waugh et al. (2004). The
open structure of the
zoarium would provide
exposure of interior feed
ing surfaces to water
flow, while the down
ward-arching fronds
would provide stabiliza
tion and attachment to
the substratum. Re
printed by permission of
Sigma Gamma Epsilon.
741, ). a wh op pi ng 26 inche s by 14 inch es by 6 inc he s (C uff ey an d Fin e 2005).
have bee n truly imp ort ant deni zen s of the Ci nc in na ti an sea f loor .
impression is nothing but enhanced when one focuses in on the deta
ju st w h e r e an d w i th w h o m they o c c u r .
A t m a n y t i m e s a n d i n m a n y p l a c e s the C i n c i n n a t i a n se a b o t t o
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A t m a n y t i m e s a n d i n m a n y p l a c e s , the C i n c i n n a t i a n se a b o t t o
pears to hav e be en soft mu d. Mo st of the kin ds of an im al s that we gen
find preserved as fossils do not seem to have "liked" soft, muddy bot
presumably because the mud did not provide solid footings upon whi
b u i l d a stable l if e. It w as al l too easy to be e n g u l f e d by the o o z e . M o re
the fine sediment was too easily swirled up into the water and clogge
spiratory a nd food- gat her ing apparati . l iven a cen tim ete r or two abov
mu d was more hospita ble . B ryo zoa ns ordinari ly did not grow their col
dire ctly on the surf ace of the m ud . But let a storm dro p a few shells or
me nt s of shells ont o the goo , and the sea floor was open for co lo niz a
So often, wh en on e is able to ex am in e the actual base of a colony —w
gr ow th c omme n c e d — on e d i sc ov e r s that the c o lon y w as fou n d e d on a
me nt of a shell of a bra chi opo d or p el ec yp od , i f not a com pl ete or n
co mp le te shell ( Fi gu res 7.3F, G, 7.4D). O n c e the sea floor was a bit
l ized, the bryozoans colonized and grew in earnest.
Once established, the bryozoans themselves added to the stabi l i
the sea floor in their immediate vicinity. First, as colonies grew, a ce
propo rtion of th em topp led over and their skeletal material b ec am e
W h e n the t i m e for s h e d d i n g was a t h a n d , t he tr ilobite a pp ar e n t l y
cli mb ed its way up into a suitable part of a bry oz oa n col on y and we dg ed
the projecti ons of its exos kel eton into the bryozo an (see Fi gu res 11 .6 E, F)
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Th is enable d the trilobite to pull itself out of the old exoske leto n and co m
men ce the har den ing of the new one , a l l the whi le bei ng hidden am on gs t
the bryozoan fronds from the eyes of wou ld -be predators. Al th ou gh this
must have been a convenient arrangement for the trilobite, it may have
b e e n le ss so for the b r yo zo an s . S hr a ke fou n d that, in s o m e i n sta n c e s , the re-
was pa th o lo g i c g r o w t h in the co lon y as the b r y o z o a n s grew up an d a r o u n d
the trilobite exoskeleton they had no way to dislodge.
T h e trilo bite/b ryoz oan association descr ibed by Do u g Shra ke is just on e
of a host of exa mpl es of the intera ctions of br yo zo ans and myr iad ot her crea
tures. On the one hand, a bryozoan larva would attach to almost anyone,
given suitable circ ums tan ces , and a co lo ny wo uld sprout. Bryozoan col onie s
have been documented as attached to, encrusting, overgrowing, or etched
into articulate brachiopods, inarticulate brachiopods, cephalopods, corals,
cornuli t ids, cr inoids, foraminifers, hydrozoans, monoplacophoran molluscs,
pele cypo ds, trilobites, and, of cours e, other bryo zoa ns, bo th o f the sam e andof different specie s (see chapter 16, Tab le 3). On the oth er han d, a nu m be r of
other orga nism s have bee n found att ach ed to or bor ed into bry oz oan s: cora ls,
i l b hi d i i l b hi d li id d l d
so me of the Wei ch ol d do ug hn ut s, there is a ring of wha t mi gh t be rec
lize d c eph al op od shell (F ig ure 7.31). Perhaps the apertural part of th
that com pri ses the shell of an orth oc on ic nautil oid cep ha lo po d broke o
c a m e to rest on the sea floor. Th en , on e or mo re bryo zo an larva e settl
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, y
this hard obje ct pro tru din g abo ve the oo ze . As the zoa ri um grew, it ass
the r ing-shape of the "se gm ent " of ceph alopod shell .
Su ch rings of cephalo pod shells have bee n descr ibed and figured
scientific literature (Teeter 1978), and similar things have been found
local rocks. However, the story may not be quite so straightforward
pro ble m is that so me of the We ich ol d doug hn ut s see m to be bryoz oan
parts all the way through—with no obvious remnants of cephalopod s
Frank McKinney, the well-known bryozoan worker , has seen ashaped co lony of the bryo zoa n genus Constellaria, with mud in the c
His interpre tat ion was that the c olo ny ha d slo wed the water and caus ed
to precipitate to such an exten t that gro wth of the co lon y was able to pr
only at the periphery (Mc Ki nn ey , pers. com m. ). However , this colo n
so me 8 to 10 inch es across (20- 25 c m ) — m o re tha n twic e as big as the
W e i c h o l d rings. Ob v i o u s ly , the p h e n o m e n o n n e e d s s o m e serious scie
study.
Figure 7.6. A. Detailed
studies of bryozoans re
quire carefully oriented
thin-sections or acetate-
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peels. Diagram shows
orientation of sectionsand terminology of inter
nal skeletal structures
used to identify species.
From Arens and Cuffey
(1989, figure 5), repro
duced by courtesy of
Roger J. Cuffey, with per
mission of the Pennsylva
nia Academy of Sci
ence. B. Left,
longitudinal thin section
of Heterotrypa fron-
dosa (d'Orbigny), CMC IP
40336, Bellevue Lime
stone, Cincinnati, Ohio,
R. J. Singh Collection.
Right, tangential thin
me an s of the gas in its ca me ra e (see cha pter 9). In addi tion , the bryo zoa n
ing migh t have increased the hydrodynamic drag on the ceph alo pod .
Th er e even m ay have b een sonic adva ntage s to h a vn i n g a coa t
b r y o z o a n s . Present-day " d e c o r a to r c r abs" arc c a m o u f l a g e d by the l
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anemones and such like that they carry. Indeed, the crabs delibe
"pl ant " oth er creat ures on their dorsal surfaces . Perh aps the bry ozo a
zoa he lp e d c on c e a l the Ci n c i n n at i an c e p halop od s that b or e the m
course, the zoarium would have covered, and thereby made v isually
less, any color patterns that the cephalopods had; but that is a stor
doe s not be lo ng in the chapt er abo ut phy lu m Bryozoa.)
Most br yo zo an sp ec im en s can be identified onl y on the basis of in
str uctu res, at least defin itivel y so. This me an s that, except for Su pe
it is necessary to cut them open—no small feat for the ordinary foss
lector. It is necessary to use a rock saw to make precisely oriented
thro ugh an individual spec ime n. Be caus e the bryo zoan co lony is pre
as part of the rock , it will n ot let en ou gh light th rou gh to sec intern
tails . This can be ov erc om e in two ways. T h e zea lou s and well -equ
paleontologist can cut and grind the specimen into slices so thin tha
b e c o m e transp are nt . These are c a l l e d th i n -s e c t i o n s , an d they arc
Studying Bryozoans
cia of wh ic h it consist s. A tra ns ver se secti on is orien ted at right an gl es to
the oth er two , for ex am pl e, across a bra nch of a give n colo ny. I n short, one
needs long itud inal , tan gent ial , and transverse sections of a colo ny to get a
com plet e pictu re of i ts internal stru cture and this com pl et e picture is es
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com plet e pictu re of i ts internal stru cture, and this com pl et e picture is es
sential to an u nd er st an di ng of just wh at kin d of br yo zo an is at han d and
ho w it grew and was con sti tut ed.
After the properly or ie nte d th i n - se c t i on s or pe e l s are m a d e , t h e o n l y
way the y c a n b e s tud ie d a d e q u a t e l y i s u n d e r the m i c r o s c o p e . T h i s , a g a i n ,
is a piece of equ ip me nt that ma y ch al le ng e one 's bud get .
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BRACHIOPODS: THE OTHER BIVALVES
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Brachiopods are among the most common fossils in the Ordovician rocks of
the Cincinnati area. Only fossils of bryozoans are more numerous to the
naked eye. In a study of ty pe- Ci nci nna tia n l imes tones , Ma rti n (1975) re
ported that brachiopods and bryozoans together constitute about 60 percent
of the fossi l frag ment s co mp ri si ng the lim est ones . There even are so me lay
ers, for exa mpl e, in the Bel le vue Limestone, in wh ic h the rock is a veritabl e
coq uin a, in this case consis ting of co mp le te and nearly co mp le te shells of
large, f lat brach iopo ds of a s ingle genu s. Th e se aptly na me d "shin gled
Rafinesquina bed s" co m m o nl y are tho ugh t of as rem ain s of very shallow- wa te r de po sit s r emi n isc en t of the shingled b e a c h e s of to day. A l t h o u g h they
have been living on E arth since the Ca mb ri an Period, brach iopo ds are not
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T h us , wh en a pe le cy po d relax es, its shell open s. by w ay of cont rast,
w h e n a b r ac h i o p o d re laxes, its shell ten ds to c lose . This has i m p o r t a n t
implications for how one finds pelecypods and brachiopods as fossils. Upon
death (the ult ima te relaxatio n), the indiv idua l valv es of a pe le cy po d tend
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p y p
to get separated from one another , because they gape apart, a l lowing cm-
rents, for example, to tear them asunder. On the other hand, the two valves
of a brachi opod shell more co m m o nl y rem ain to gether , and are foun d that
way by the intrepid fossil co l lector .
Brachiopod shells tend to be better preserved than are pelecypod shells
for anoth er reason, too. Most brac hio pod shells consist of ca lc iu m ca rbo n
ate, but so do pelecypo d shells. How ever, ca lc iu m ca rb on at e exists in mo re
than one form. Most bra chi op od shells arc of the mine ral cal cit e, wh ere as
pele cypo ds consist of or includ e aragon ite . The ato ms of ca lc iu m, car bon ,
and oxygen are arranged differently in aragonite and calcite, and the two
substances have di f ferent properties. Bec aus e of this , pel ecy pod s tend not
to be pres erve d as wel l as br ac hi op od s. The pract ica l result is that you may
find ma in well-preserved brac hio pods In the rocks of the Ci nc in na ti area,
but p e l e c y p o d s , with few e x c e p t i o n s , are pr eserved as internal m o l d s .
Brac hiopo ds are f i l ter feeders. T h e y extract small partic les of organ ic
matter from the sea water. These partic les are captu red by a ciliat ed struc
ture called the lophophore (Plate 3F; Figure 8 2A) 'The lophophore oc
of the op po sin g valve. B ec au se of the in terl ocki ng teeth and socket s, t
cal led dent itio n, it wo ul d be difficult for a wou ld- be devourer of brac h
flesh to twist the valves apart to get at supper.
A n i m a l s o f the o th e r m aj or g r o u p o f b r a c h i o p o d s , not surprising
t d i t i l t I th i l th ith t th
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termed inarticulates. In these animals, there are neither teeth nor so
No t having a real hi ng e, the task of ke ep in g the valves togeth er is a gc ha l le n ge for an i n ar t i c u la te . T he mu sc u latu r e i s a good d e a l mor e
plicated in inarticulates than in articulates—to keep the two valves
b e i n g twisted apart f r o m o n e ano the r .
In an art icul ate , there is a hin ge , wh ic h serves as a fu lc ru m. The
for muscles and the adductor muscles, in order to open and close the
operate against one anoth er abo ut the fu lcr um (Fi gure 8.2A). But an
ticulate has no such fu lc rum . T h e an im al ope ns its shell , not by con
ing diduc to rs, bu t by pul li ng the bo dy back towar d the rear of the
thereby causing the valves to gape sufficiently for the animal to fee
spire , and perform other necessary activ i t ies.
A n o t h e r c o m m o n dif ference b e t w e e n articulates and inarticulat
vo lves the c o m p o si t i o n of the shell. In most inarticulates the shel l co
not of ca lc iu m carb ona te, but, rather, of ca lc iu m pho sphat e. (N ote, ho
that this is not a univer sal r ule , for the shel ls of so me of the ani ma ls tra
alh called inarticula tes are- ca lc iu m car bona te, like those of the articu
diversity see n in this im por tan t gr ou p. His work has gu id ed and inspire d
much subsequent research and the discussion that follows
In ar t i c u la te s
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In ar t i c u la te s
Some present-day inarticulates (genus L i n g u l a and its relatives) spend
mu ch of their tim e in a burrow . Wh e n it co me s ti me to feed, the pedi cl e
ext end s so that at least the anter ior part of the shel l pr oje cts up int o the
wat er . In some b r ac h i o p od s , there is an o p e n i n g in the shell t h r o u g h w h i c h
the pedicle extends. In general, this pedicle foramen is in the anatomically
lower valve, wh ich , he nc e, is cal led the ped icl e valve. The ped icl e of so me
b rac hi o pod s may be a t tache d to a n o t h e r sh el l on the se a flo or; this m a yresult in pedicle attachment scars which consist of tiny, characteristic pits
in the other shell. In an inarticulate brachiopod, as in you, there is a mouth,
esophagus, stomach, intestine, and anus. In an articulate brachiopod, how
ever , there is no anus (f igure 8.2A). What might have been a complete
digestive tract is, in fact, a cul de sac. Thus, the only egress for waste mate
rial is back out the mouth. In present-day animals that have been studied,
solid waste is regurgitated as small pellets; these are then expelled from the
shell by rapid snapping of the valves.
A l t h o u g h the articulates ar e m or e readily n o t i c e d inart icu lates are not
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to the stratif ication, in what appears to be its l ife position, that is, the ani
mal's orie ntat ion in space du ri ng its l ife. The ind iv idu al in Fi gu re 8.3A is
a case in point.
It is only by great good fortune that that particular individual made it
Figure 8.3. A. Pseudo-
lingula sp., CMC IP
51994, Cincinnatian, ho
rizon and locality un
k S i
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into a m us e um . R alp h D ur y w as a C in c i nn at i ent o m o lo g is t of s o m e co n
siderable repute . H owever , his ac tua l l iv eli hoo d was in real estate. Ac co rd
ing t o C h a r le s D ury , C h ar les ' s so n and d irect o r o f t h e C i nc in na t i M u s e u m
of Natur al History for mor e than f ive de ca des , the sp ec im en in f i gu re 8 .3A
started its museum career in a real estate project . Charles Dury was having
a stone wall built. Being a meticulous fellow, he visited his sites on a regular
and freque nt basis. He ha pp en ed to ma ke one of the se visits to the bui ld in g
site shortly after a load of stone ha d be en d eli ver ed. In the cou rs e of ex am
ining the stone to be certain that it was up to his standards, he saw the
ch un k with the lin gu li de i n its l ife posi tion . So, inst ead of en di ng up as
part of a wa ll, the p ie ce of rock en de d up as part of the co ll ec ti on s of the
Cin ci nn at i Mu se um of Natu ral Hist ory , alo ng w ith Charles Dury 's in sect
co l lec t io n— but t h at i s ano t h er s t o ry ( Vul inec and D av is 1 9 8 4 ) .
A r t i c u l a t e s
T h e vast majority of the brac hiopod s that on e sees in the Or do vi ci an rocks
known, x 1.9. Specimen
oriented perpendicular tobedding, presumably in
life position with beak
downward. B. Trema-
tis millepunctata Hall,
CMC PT 585, brachial
valve interior, Cincinna
tian, horizon and localityunknown, x 3. C. Pet-
rocrania scabiosa (Hall)
encrusting on Heber-
tellasp., MUGM 29461,
Arnheim, Oxford, Ohio,
x 1.3. D. Petrocrania
scabiosa (Hall) encrust
ing on brachial valve of
Rafinesquina sp., Bruce
and Charlotte Gibson
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Figure 8.10. Environmen
tal distribution of bra
chiopods in the Cincin
natian Series. Shoreface
environments are equiva
l t t th h ll b
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lent to the shallow sub-
tidal (1-2 m or 3-6 ft);
transition zone environ
ments are deeper sub-
tidal (3-6 m or 10-20 ft),
and offshore environ
ments are deeper water,
with a maximum depth
of about 30 m (100 ft).The heavy lines indicate
the environments where
each genus is most abun
dant and thin lines indi
cate environments where
a genus is present at
lower abundance. From Holland (1997), in Pale
ontological Events.
Copyright 1997 Columbia
lop " you enjoy in your favorite sea food rest aura nt is an add uc tor mus cl e of
one of those pel ecy pod s. The add ucto r of a sca l lop is pow erfu l en ou gh , in
life, to snap the animal's valves together so swiftly that the creature can be
lifted above the sea f loor. Some scallops can even swim for some distance,
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although rather jerkily and in decidedly irregular trajectories .
Datt i lo (2004) has found evid enc e that one Cin ci nn at ia n brachi opod
with a c o n c a v o - c o n v e x shel l, Sowerbyella , was capab le of esc api ng from
burial b en e a t h s edi m ent s st ir re d up by storms, p r es u m a ble by s n a p p i n g its
valv es. Individuals of Rafinesquina may have had similar capabilities, be
cause convex-up specimens are found with a moat-like furrow around the
commissure that formed while the brachiopod was al ive (Meyer 2006).
Th es e recent f indings suggest that these br achiop ods wit hou t ped ic le atta ch
ment might have led much more active lives than previously realized.
Despite the high divers ity of type-Cincinnatian brachiopods, the distribu
t ion of specie s is not uni for m th ro ug hou t the strat igraphic suc ces sio n.
There are distinct associ atio ns of spe cie s an d shell typ es that ch ar ac te ri zedif ferent s trat igraphic intervals and even i ndiv idua l beds. Re ce nt research
In Steven Holland, in col la borat ion with Arn old Mil ler , David Meyer , and
Distribution of
Type — Cincinnatian
Brachiopods in
Time and Space
Even on a smaller scale, brachiopods reveal some very basic aspe
life on the Late Ordovician sea floor. Very densely populated lime
b e d s , f e a t u r i n g b r a c h i o p o d s l i ke Rafinesquina, Strophomena, and
manella, a r e v e r y c o m m o n thr ou gho u t the ty p e - Ci n c i n n at i an an
called shell pavements (also called shingled beds as noted above). In
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called shell pavements (also cal led shingled beds as noted above). In
shell pavements, the brachiopods are usually preserved with convex v
upward, sometimes covering the entire bed surface (Figures 8.4G,
8.8E). She ll p av em en ts can be as thin as a single layer of shells, or th
with the ent ire t h i c kn e ss up to a few tens of c e n t i m e t e r s c o n s i s t
stacked brachiopods. In some cases, the valves are vertical or ti lted at
ous ang les and pack ed closely tog eth er in an ed ge wi se shell bed. The
w is e shell p a v e m e n t s are g o o d e v i d e n c e of water m o v e m e n t i n the fo
w ave osci l la t ions b e c a u s e there are present-day e x a m p l e s of e d g e w i s e
b eds an d shale f r ag me n t s f o r m e d in very shal low water by wav e oscil la
II the hingelines or beak s of the bra ch iop ods were always direct ed d
ward in an e d g e w i s e b e d , it m i g h t be possible that the densely p ac ked
actually had lived in a manner similar to an oyster bed. However, an
of valv e orie ntat ion wi th in ed ge wi se shell bed s shows that valves d
show such a pattern and even can be predo min ant ly hingel ine- up
(Sei lacher 1973, pers. comm.).
There has been considerable debate as to how shell pavements
W ay nes v i l le Fo rmatio n as a for m of p a l e o e c o l o g i c s uccess ion (see F ig ure
4.S). In present-day settings, ecologic succession occurs when one assem
blage of ani m als or plants al te rs the ha bi ta t IN such a way that other species
can replace the so-called pioneer species. Harris and Martin (1979) sug
gested that thin shell ed brachio pods were pione er speci es that first col on iz ed
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gested that thin-shell ed brachio pods were pione er speci es that f irst col on iz ed
soft mu dd y patch es of the sea floor and p rovid ed a p av em en t on whi ch en
crust ing ani mals l ike bryoz oans and inart iculat e brachiopods coul d sett le .
Even tual ly other species cou ld take adva nta ge of the shell pa ve me nt and
thickets of bryo zoa ns, so that the diversity of the asse mbl age incr eased up
ward from the bo tt om of a p a v e m e n t bed. St o rm s frequently s m o t h er ed t he
shelly patches with mud, thus interrupting the succession until brachiopod
larvae onc e again could c olo niz e the barren m uds . some paleoecolo gistshave quest ioned whether paleoecologic succession comparable to present-
day succe ssion ca n be det ect ed in the fossil record be ca us e mos t stra tigr aphi c
changes in fossil assemblages represent a much longer time scale than the
scale of years to decades over which present-day succession occurs. Although
we st il l do not know how m u c h t im e was required for the fo rmatio n of c har
acteristic, thin typ e-C in ci nn at ia n shell pav eme nts , it is possible that they
formed over a short time scale. It also seems correct to view the brachiopods
as having a pivotal role in providing a hard substratum onto which encrust
i i l ld ttl th lt i th h bit t i th f
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MOLLUSCS: HARD, BUT
WITH A SOFT CENTER
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9Everyone knows molluscs—the oh-so-familiar snai ls and slugs, the c lams,
mussels, scallops, and oysters, the octopus, and the squid. But the mollusc
story is not a simple one . There are mor e kinds of mo ll us cs tha n of any
other gro up of an im al s, save the arth rop ods . So wha t links all the mo ll us cs
together?
The word "mollusc" is der ived from the Latin word "molluscus,"
me an in g "soft." This refers to the fact that eve ry mol lu sc has a soft, flesh y
body. But that, of c o u r se , is not the i m a g e c o n j u r e d up in the m i n d ' s e ye
at the ment ion of snails, cl am s, and oysters. In m os t of the mo ll us cs , thesoft parts are enc lo se d wit hi n a hard she ll. A n d it i s on th e basis of differ
ences in the shells that the moll uscs of the typ e- Ci nc in na ti an are di f feren
Of co urs e, even i n mo ll usc s that ha ve shells, t he hard parts are m
a part of the whole animal. In general, the body of a mollusc incorpo
five of wha t co m m o n l y are calle d bod y regio ns: th e head, the foot
v iscer al m a s s , th e m a n t l e - c o m p l e x , a n d th e gi l ls ( te c hn i c a l ly t e
cteni dia, from their co mb- li ke shape). The man tl e , the shell , and the
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t le-cavity togeth er com pri se the ma ntl e-c om ple x. The man tle is a shetissue that hang s do wn on e ach side of the bulk of the ani ma l or other
enc los es it. The shell, i f pres ent, is att ach ed to the outs ide of the m
and is secreted by it. To the inside of the mantle is the mantle-cavit
w h i c h are lo cated the gi l ls , i f the y are present .
T h e last phra se of the previo us par agr aph is an imp orta nt tip-off
all kinds of mo ll us cs have all f ive body r egio ns dev el ope d to the sa m
tent. Snails, for example, each have an obvious head, whereas clams do
Squids have well-developed gills; however, in terrestrial snails, there ar
ctenidia, and the mantle-cavity serves, in effect, as a lung.
It is only fair to admit that there is so much morphologic and ana
cal variation among the molluscs as a whole that it is difficult to poi
any trait that occurs in all molluscs. Some have shells, and some do no
mo st , the shell is ex te rn al , but in some i t is int erna l. So m e hav e gil ls
some do not. Some have heads, and some do not. And so on.
Regardle ss of whe th er on e is co nv in ce d that form follows functi o
consists of a nu mb er of small er grou ps of org ani sms ca l le d classes): c lass
C e p h a l o p o d a , c l a s s G a s t r o p o d a , c l a s s M o n o p l a c o p h o r a , c l a s s P e l e c y p o d a ,
c las s R o s t ro co nch ia , c las s Po ly p laco p h o ra, and c las s Scap h o p o da. Th ere
are some groups of moll uscs of wh ic h no sp ec im en s are kno wn fr om the
local rocks For example the class Aplacophora includes certain present
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local rocks . For example, the class Aplacophora includes certain present-
day ani mal s that are devoid of preserv able hard parts ; he nc e, their nam e,
w h i c h m e a n s " n o plate b ea r i n g . "
Snails , with their characterist ic coiled shel ls , are probably the easiest to Class Ga st ro po da —
rec ogn iz e of the mollusc s in the rocks of the Cin ci nn at i reg ion. Snai ls are Th e Snails
ve ry c o m m o n in m a i n e n v i r o n m e n t s of t oday's w o r ld , a l o n g wit h their re la
tives, the so-c alle d slugs and sea -slugs. ( An im al s of the latter two gro ups
either have no shell at all , or a small, internal one; thus, they are unlikely
to he preserved as fossils.)
Snails are the quintessential univalved mollusc (Figure 9.1). Each snail
has a single pr om in en t valv e. In most snai ls, this is a long , nar row, con ic al
tube that is coil ed off to one side, so that it re sem bl es a screw. Of co ur se, by now, y ou ha v e c o m e to e x p e c t that t h e sto ry is b o u n d to be fa r m o r e
co m p l icat ed .
Class Gastropoda—
The Snails
Figure 9.2. Cincinnatian
monoplacophorans and a
bellerophontid gastro
pod. A. Archinacella
area Wahlman, USNM
40615 a monoplacoph-
Mo st of the snail fossils in the rocks of the Cin ci nn at i region co
of mo ld s. The ac tu al shell mat te r has bee n diss olve d away, and all th
left is the sediment that originally fi lled or that surrounded the bu
shel l, or bot h. Ex cep ti on s to this gene rali ty are the snails of the ge nu
clonema; he r e , shel l matte r, n ot u n c o mm on ly , is present.
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40615, a monoplacoph
oran, Waynesville
Formation, Waynesville,
Ohio, a, dorsal, b, lateral,
c, anterior, all x 1.6. From
Wahlman (1992, plate 3,
figures 7, 9, 10). B. Hel-
cionopsis striata Ulrich
and Scofield, USNM
45827, a monoplacoph-
oran, Richmondian, Mar
ion Co., Kentucky, a, dor
sal, b, oblique-lateral, c,
lateral, all x 1.7. From
Wahlman (1992, plate 2,
figures 1, 2, 3). C. Cyr-
tolites ornatus Conrad,
USNM 265906, compos
G a s t r o p o d s o f t h e t y p e - C i n c i n n a t i a n
Snai ls (Figures 9.2, 9.3) are very common fossi ls throughout the Ci
natian Series in its type area, but, because they generally are preserved
as internal molds, then- can be overlooked, and precise identification
be di f ficult . N e v e r th e le s s , sna i l s undoubtedly placed an important ro
the e c o log y of the Ci n c i n n a t i an se a, e sp e c i a l ly i n some e n v i r o n m
w he r e t he y we re very a b u n d a n t . Hol l an d 's c o m p i l a t i o n o f C i n c i n n
fossils lists sixty-three species of snails in twenty-three genera as occu
abo ve the base of the K op e Fo rma tio n (H ol la nd 2005). Of thes e, si
species in nin e genera bel on g to the planispiral ly co i led belleroph o
that were revised tax ono mica lly by Wah l ma n (1992). of the othe r fou
genera and forty-seven species, the genus Cyclonema accounts for e
spe cie s, bas ed on the 1970 stud y by Th om ps on (1970). Spe ci me ns
additional p latyceratid, which belong in Naticonema, occur in the
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C inc innat ian are am o ng t h e o ldes t - k no w n cas es . Sp ec im ens o f Cyclonema
varicosum at tached to Pycnocrinus are kn ow n from the Lex in gto n Li me
stone, just below the Ci nc in na ti an (Felto n, pers . co mm .) . Wh et he r al l
cases of snails att ach ed to Pa leo zoi c crinoi ds are insta nces of co pr op ha gy
has been debated; other possibil it ies include parasit ism predation and
Figure 9.3. Cincinnatian
gastropods. A. Cy
clonema varicosum
Hall, CMC IP 51118, Cyn-
thiana Formation, Lexing
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has been debated; other possibil it ies include parasit ism, predation, and
commensal ism (Ba umil ler 1990; Morris and Felton 1993). (In co mm en sa l-
ism, indiv idua ls of the associa ted spec ies gai n so me adva nt age whil e the
host is una ffec ted. Gi ve n the size and l oca tio n of the snails on the cri noi d
host, it is difficult to see how the host co ul d rema in unaffe cted .) A mor e
complex associat ion between Cincinnatian gastropods, crinoids , and tubes
of Cornulites enc ru sti ng the snails was investigated by Morr is and Felton
(1993, 2003). These aut hor s sug ges te d that Cornulites gained an advantage
by en cru st i ng the sn ails tha t l ive d on the elevated c r o w n s of t he cr i noi ds ,
either by sha ri ng in the feca l feast or sim ply by virt ue of the ele vat ed posi
tion provided by the crinoid.
Most individuals of Cyclonema that are found are not attached to cri
noids, suggesting that the association with crinoids was not obligate, and
that specimens of Cyclonema were able to make their l iving in other ways.
We la ck direct e v i d e n c e for the f e e d i n g habits of f ree- l iv ing s p e c i m e n s of
Cyclonema, but the feed in g habits of presen t-day gas tro pods ma y prov ide
ton, Kentucky, x 1.7. B.Cyclonema humero-
sum, Steven Felton col
lection, showing aper
ture. Grant Lake
Formation, Brown Co.,
Ohio, x 1.2. C. Cy
clonema humerosum
Ulrich, CMC IP 51117,
Maysvillian, Cincinnati,
Ohio, x 1.6. D. Cy
clonema bilix lata (Con
rad), CMC IP 51116, Arn-
heim Formation,
Cincinnati, Ohio, x
1.7. E. Cyclonema
sublaeve Ulrich, CMC IP
51114, Fairview Forma
Figure 9.4. Gastropod-
rich beds. A. Miam-
itown Shale, Trammel
Fossil Park, Sharonville,
Ohio. Note shells with
geopetal cavities (lower
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part filled with sediment,
upper part filled with
calcite crystals). B. Mar
ble Hill Bed, Waynesville
Formation, Trimble Co.,
Kentucky. Scale in mm.
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im po se d onto on e an ot he r, so that th e featur es of the exteri or of a val
th os e of the inter ior are visibl e in ess ent ial ly a sin gl e sur fac e. The
mat ter of the livi ng pelecy pod so represente d must have bee n fairl
i n or gan i c mate r i a l , b e c au se the c omp osi te mold c ommon ly i n c lu
carbon fi lm where the shell substance once was (Figure 9.7A).
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A l t h o u g h preservation ma y provide a c lu e , the real wa y to id en tpe le cy po ds is by refer ence to the mo rp ho lo gy of the shell. The pl
sy mm etr y in a ty pe- Cin cin nat ian pelecypod is bet wee n the valves, wh
in an ordi nary articu late br ach iop od, th e plane of sy mm etr y runs
each valve perpendicular to the hinge (see Figure 8.1) .
Of cou rs e, the fossils in the rocks in and near Cin ci nn at i have
there for hun dre ds of mil l io ns of years. Ma ny thin gs migh t have ha p
du ri ng that span of tim e. S om et im es , for ex am pl e, the dead shell wa
ied in other than a living position. As soft sediments built up ever d
on the sea floor, the y b ec a m e co mp res se d into rock. In some ins tanc e
origina l bi lateral sym me tr y of the onc e-l iv i ng ani mal (and, hen ce,
shell) was distorted by the pressure, so that the shell appears skewe
such inst ance s, the orig inal symm etry may not be imm edi at ely obvi
Pe le cy po ds of th e Type — Cincinnat ian
Thus. Frey's work do cum ent ed the spread of pel ecy pod s from ma inl y c las
tic (shale-r ich) sedimentary environments in ear ly and middle Cincinna
tian t ime into carbon ate envi ron men ts in the late Ci nci nn at ia n. The abu n
danc e and diversi ty of pel ecy pod s in the ty pe- Ci nci nn at ia n demo nstr ate
that, even early in their evo lut ion ary history, pe le cy po ds oc cu pi ed mar in e
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environments in a variety of offshore settings, heralding their subsequent
diversi f ication and increasi ng ab un da nc e in the late Pale ozoi c and Mes o
zoic (A. I. Miller 1989).
By Late Ordovician time, pelecypods had exploited almost the full
range of living habits found in present-day forms (Figure 9.9). M ost c om
monly, Late Ordovician pelecypods used byssal threads for attachment to
objects on or within the sediment—similar to present-day mytilids (mussels).
Epibyssate forms, such as Ambon y c hia , attached at the sediment surface or
nestled with in the bra nch es of bry ozo ans . E ndobyssate forms, suc h as Modi-
olopsis and Pseudocolpomya, attac hed to shell frag ment s or sed ime nt grains
just below the se di me n t surface and exten ded the she ll for fi lter feeding.
Free-burrowing forms included deposit feeders (who fed on organic particles
wi th i n the sediment), such as Ctenodonta, and shallow infau nal filter feeders,
such as Ischyrodonta. As mentioned above, the ability to bore into hard sub
strata was first seen in the Late Ordovician Corall idomus (F i gu r e s 9 . 8 A -C).
d l d h d d l d h l h d
In the majority, the shell is a long, straight, conical tube closed a
narrow end . (Straig ht shells are said to be ort ho con ic. ) There are, how
so me in wh ic h the shell is curv ed or eve n coile d. In mo st of the kin
cephalopods present, there are partitions that go across the tube for p
its le ng th ; the se are ca ll ed septa (singula r, se ptu m) . The ch am be rs th
separated by the septa are called camerae (singular, camera). At the l
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end of the con ic al tub e that co mp ri se s the shell is a por tio n in wh ic h
are no septa. This is cal led the body cham ber . The cam era e are co nn
to one another by a tube that runs through all the septa from the
ch a mb er to the first cam er a at the tip of the co ne ; this inner tube is
the siph uncl e . The l ine al on g wh ic h a parti cular sep tum mee ts the
wall of the shell is c a l l e d a sutu re.
A l l p re se n t-d ay c e p h a l o p o d s l ive in the o c e a n s , and the re is
reason to conclude that the fossi l forms were marine creatures, too. D
li fe , the apt ly- nam ed body ch am be r was occu pi ed by the bulk of th
parts of the cep ha lo po d an im al , and the other cam era e apparentl y
tained gas. (In present-day Nautilus, the gas is similar in compositi
air, but without the oxygen.) Such a gas-fi lled shell would have served
f loat , b u oy i n g the an i mal u p i n the w ate r an d , d e p e n d i n g on how gas was in the camerae, allowing the animal to stay at a particular
dept h wit hou t the effort of sw i m m in g upwa rds or dow nwa rds . At
Figure 9.10. How nau
tiloids attained happi
ness. In the top diagram,
the center of gravity, G,
is inferred to lie in the
body chamber of the
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nautiloid, and the center
of buoyancy, 6, to lie in
the gas-filled phragmo
cone, forcing the animal
into a vertical, head-
down position. In this
position it could not
swim efficiently. In the
middle diagram, the for
mation of calcareous
cameral deposits in the
apical part of the phrag
mocone should result in
the center of gravity and
center of buoyancy shifting toward the midpoint
of the animal's length (G 2
Figure 9 . 1 1 . Cincinnatian
orthoconic nautiloids.
A - C from Davis and
Mapes (1996), courtesy
of the Ohio Department
of Natural Resources Divi
i f G l i l S
Ap p are n t ly , from the p o i n t of v i e w of o r g a n i c e vo l u t i on , this was
b en e f i c i a l s i tuation. At least in a n u m b e r of ev olu t io n ary l in e age s , c h
in the shell occurred that solved this problem in one way or another
greater or less extent.
Before proceeding further on this tack, we need to talk a bit
org ani c evo lut io n, per se —l es t we go astray into th in ki ng that, bec au
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sion of Geological Survey. A, B. Treptoceras
duseri (Hall and Whit
field), Cincinnatian, scale
bars = 1 cm. A. OSU
47422, internal mold of
part of phragmocone
and living chamber.
B. OSU 47417, part of
external shell, with dark
longitudinal lines which
are remains of what, in
life, were color bands.
C. Cameroceras in-
aequabile (Miller), internal mold of portion of
siphuncle, OSU 47420,
liv ing cephalopod did not "l ik e" its face in the mud , it delibera tely ev
its shell morphology and structure to avoid that condition.
As far as is k n o w n , the vo lition of o rganisms has no ef fec t on the o
evo lut ion of a lin eag e. Thu s, no cre atu re can will the orga nic evolu tion
des ce nde nt s to go into any par tic ula r dir ect ion , no matt er how b enefi c
enjoyabl e?) that dire ctio n m ig ht prove to be. R ather, it a particu lar dir
means that more creatures will survive to pass their genes on to future g
tions , th en the org ani c evol uti on of a li neag e w ill tend in that dire ctio n
mechanism is called organic evolution by natural selection, or natural
tion, for short. Some paleontologists and other biologists speak of evolut
"strategics" whereby a linea ge "solves" som e particu lar "probl em." The
figures of spe ech . They are, in tact, gener aliz atio n of what chan ges ac
are seen in the fossil record of the parti cula r lin eage . they definitely imply that org ani sms perc eive d a pro bl em and will full y evolve d to solv
Let us return to the type-Cincinnatian. One way to br ing the sh
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for 76 percen t of the 302 sp ec im en s found . S pe ci me ns of oth er spec ies are
much less abundant; included here are the cyrtoconic forms (having
curved shells) Manitoulinoceras tenuiseptum an d M. williamsae (8 percent),
Oncoceras delicatum (4 percen t), Zittelloceras russelli, and Z. williamsae (<1
percent), longiconic forms (having long, tapering, straight shells) tenta
tively assigned to genus Isorthoceras (6 percent) the endocerid Camero
Figure 9.12. Upper Cin
cinnatian (Richmondian)
nautiloids. A. Charac-
toceras baeri (Meek and
Worthen), MUGM 29591,
Whitewater Formation
Preble Co Ohio scale
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tively assigned to genus Isorthoceras (6 percent), the endocerid Camero-ceras inaequabile (6 percent, ha vin g a wide siph uncl e inf i l led wit h conic al
deposits), the orthocerid Gorbyoceras curvatum (<1 percent, an orthoconic
form), and the ascocerid Schuchertoceras obscurum (<1 percent, a form that
lost part of the phr ag mo co ne dur ing growt h to main tai n stabi l i ty) . T h e
material of the outer wall of eac h shell was prese rved on ly wh en en cru st ed
by b r yo zoan s , but the septa an d s i p h u n c u l a r s tru ctur es of the shell int eri ors
were replaced b y ca lc i te . B o d y c h a m b e r s an d p h r a g m o c o n e s w e r e inf il led
with c laystone, but , in s o m e cases, the c a m e r a e r e m a i n e d e m p t y or w e r e
infilled with calcite crystals. These preservat ional features led Frey to co n
clude that the nautiloid assemblage was buried in situ as complete, empty
shells . Rem oval of encr ust ing bryoz oans revealed the remark able preserva
tion of rem nan ts of col or patterns on the exte rior of the shell . In co ntras t
to shells of the pr esent -day Nautilus that are known to float after death and
drift with curren ts, the th inn er shells and septal m or ph ol og y of the se Or
dovician nautiloids suggest that dead animals sank to the bottom
Preble Co., Ohio, scale
bar = 2 cm. Compare to
present-day Nautilus,
Plate 4. B. Beloito-
ceras amoenum (Miller),
CMC IP 24415, internal
mold of phragmocone of
a cyrtoconic form, Whitewater Formation, Butler
Co., Ohio, x 1. C. Di-
estoceras eos (Hall and
Whitfield), MUGM 382,
internal mold of partial
phragmocone and body
chamber, Whitewater Formation, Preble Co.,
Ohio, scale in mm.
ro nm ent of pre dom ina ntl y lim esto ne deposit ion (Frey 1989,1995). The
two incurs ions of this "tropical faun a" into the Ci nci nna ti Arch reg io
first du ri ng the Edenia n, and the sec on d, alo ng with a host of oth er s
of invertebrates, dur ing middle-to-la te Ri ch mo nd ia n time (Frey 1989)
oldest Ri ch mo nd ia n eleme nts of this "tro pica l" nautiloid assemblag e ar
in the Waynesvi l le "Treptoceras duseri shale" in the form of specime
S h h t h d G b t (F 1985 1995)
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Sehuehertoceras ohscurum an d Gorbyoceras curvatum (Frey 1985,1995).
mondian limestone-rich units, such as the Drakes, Saluda, and White
Fo rm at io ns cont ain a divers e naut iloi d fauna that represents the peak
"Richmondian invasion." According to Flower (1946) and Frey (1995)
faun a totals sixty-five spec ies in t went y-fo ur gen era, of wh ich twen ty s
are co m mo n. So me characteristic forms are Narthecoceras dunni (Frey,
and Gharactoceras, Diestoceras, an d Gorbyoceras, shown in Figure 9.12
N a u t i l o i d T r a c e F o s s i l s ?
The late Rou sse au H. Flo wer, on e of the most prolific researchers on
zoic nauti loid ceph alo pod s, was inspired by type- Cin cinn ati an cephal
as the first docto ral stud ent (Ph .D ., 1939) of Ke nn et h E. Cas ter at the
ve rs it y of C i n c i n n a t i . In a 1955 paper , F l ow e r dr ew attention to the ta c
despite the ab un da nc e of ce ph al op od s in the Pa leo zoi c fossil record,
common. These are the gastropods (snai ls) , the pelecypods (c lams), and
the cephalopods (relatives of present-day Nautilus, squid, and octopus).
Type — Cincinnat ian representative s of eac h of these "major gro ups " are co n
sidered in some detail above. In addition to these better-known classes,
spe cim en s of four less we ll -k no wn classes often referred to as the "mi no r
moll uscs" also occ ur in the Ord ovi c ia n rocks of the Ci nci nn at i area Th es e
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moll uscs also occ ur in the Ord ovi c ia n rocks of the Ci nci nn at i area. Th es ear e the mon op lac op hor an s , the r ostr oc on c hs , the p olyp lac op hor an s (c hi
tons), and the scaphopods ("tusk shells").
M o n o p l a c o p h o r a n s
Monoplacophorans (there is no other "common name") are actually fair ly
diverse in the type-Cincinnatian, with twenty-three species in eight genera
(Figure 9.2; Wah lm an 1992; Holland 2005). Mo no pl ac op ho ra ns are similar
to the present-day gastropods called limpets in that each has a single, cap-
shaped or planispirally coiled shell and a muscular foot for locomotion.
Present-day forms like Neopilina graze on microbial mats through use of
the radula. Cyrtolites i s the most c o mm on t yp e - Ci n c i n n at i an for m ( f i g
ures 9. 2C , D) pres erve d either as a c alciti c s hell or as an inte rna l mo ld , butco mm on ly i t is encruste d with bryo zoan s. The keel ed, p lanispiral shell
with a d i a m o n d - s h a p e d ap e r tu r e is di st in ct ive
Scaphopods
Sc ap ho po ds so me ti me s go by the c o m m o n na me "tusk shells ." The
ano the r gro up of mol lus cs of wh ich s pec im ens potential ly could occ u
the typ e -Ci n c i n n at i an , b u t , as ye t , n on e hav e b e e n d oc u me n te d i n
scienti fic literature as ha vi ng co ni c from th e local rocks. Present-day
phopods each have a small , curved conical shell open at both ends,
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phopods each have a small , curved conical shell open at both ends, l ive partly buri ed in the sed im ent as deposit feeders. T h e oldest-kn
fossi l scaphopod, Rhytiodentalium kentuckyensis, was described from
Le xi ng to n Lim es to ne of Ken tuc ky, the for mat ion just below the base o
Ci n c i n n at i an (Poje ta an d R u n n e ga r 1979) . Poss i ble sc ap h op od sp e c i m
have been found in the type-Cincinnatian (Felton, pers. comm.).
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ANNELIDS AND WORM-LIKE FOSSILS
10
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Because worms are largely soft-bodied, their fossil record is rather limited.
Nonetheless , numerous foss i ls occur in the Cincinnatian that can be at
tr ibuted to the Phylum Annelida or related worms. Annel ids , the segmented
w o r m s , i nc lude p r e d o m i n a n t l y freshwater a nd te rrest rial l e e c h e s a nd eart h
w o r m s , and th e p r e d o m i n a n t l y m a r i n e p o ly c h ae t es . I n t h e m o d e r n o c e a n s
polychaetes are highly diverse and abundant and play many important
eco lo g ica l ro les . Th ro ug h o u t t h e C i nc in na t i an c o m m o n t o o t h -l ik e m icr o -
foss i ls cal led scolecodonts indicate that polychaetes were also components
of the Or dov ic ia n mari ne ecosyst em (Eriksson and Be rg ma n 2003). Al
though they resemble conodonts , another category of tooth-l ike foss i ls , in
size and form sco lec odo nts are distinct in ha vi ng a jet black app ea ra nc e in
h b l i l f d ( l ) h d f i i
. . . w e can easily imagine
that the ocean beneath
which the Cincinnati
group was deposited,
at times swarmed with
innumerable worms,
which have, so far as
we at present know,
left no traces of them
selves excepting their
jaws, tracks, and pos
sibly a few rude impres
sions of their bodies
specimens are very small, and have a distinctly segmented appearance
ure 10.1 B; R ob iso n 1987, figure 12.41F). Ulrich na me d this wo rm Protos
and desc ribed four speci es, all occ ur ri ng in the Ec o no my beds of the
nia n (now the Ko pe fo rm at io n) . Mil le r and F aber (1892b) des cri bed a
species also from the Edenia n, from "near the low water ma rk " of the
River, equiv alent to the Fulto n Me mb er of the Kop e Forma tion . A spe ci
in the collec tions of the Ci nci nn at i Mu se um C ent er is labeled from
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400-foot eleva tion , wh ich woul d plac e it in the Mavsvill ian C orr yvi ll e
mation. Although these fossils have not been restudied, their identifica
as wo rm s has not be en chal le nge d. In the Treatise on Invertebrate Pale
ogy. Protoscolex is listed with wo rm s for wh ic h the phylum is unc ert ain (H
ell 1962). However. Robison (1987) illustrated a specimen from the U
Or dov ici an of Ken tuc ky as a fossil ann elid .
The conic al en crus tin g fossil Gornulites i s a c ommon Ci n c i n n at i an ma
fossil of uncertain zoological position but with possible worm affinities.
nulites occurs throughout the Cincinnatian and several species have b
described. In the Treatise on Invertebrate Paleontology, Fisher (1962) descr
Cornulites as small tube s of ca lc iu m carb ona te, with a circula r cross-se
of 2 to 20 mm diam ete r and a le ngt h of 5 to 80 mm . Sm al le r Gornulite
Cornulites and
Other Small,
Conical Fossils
brate Paleontology, Fisher (1962) proposed that tentaculitoids and some
other small con ica l shells be incl ude d in an ext inc t c lass Cr ic oc on ar id a
(mea nin g "small , r inged cones") be lo ng in g to the Mol lus ca . Mor e recently ,
tentacul itoids have been treated as a c lass unto themselves , Tentacul i-
toidea, and not inc lud ed with any kn ow n gro up (Bergst rom 1996b). Cl us
ters of tentaculitoids are sometimes found on a bedding plane in parallell i b bl d b ( Fi A) T i
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al i gnm ent probably caused by water mo ve me nt ( Figu re 10.1A). Tw o specie s
are kn ow n from the type-Cincinnatian, Tentaculites sterlingensis and T.
richmondensis, bo t h f ro m t h e R ich m o ndian.
A n o t h e r gro up of small , ca lc a re o us , c o ni ca l foss il s th at c a n be fo und in
the Cincinnatian section are the hyolithids. Hyolithids are much smaller
than either Cornuli tes or tentacul it ids; those occurring in the Cincinnatian
are about 2 to 3 mm long. Unlike Cornulites or tenta culi tids, hyolithids hav e
a smooth shell with a roughly triangular cross-section and an operculum or
cap closin g the aper ture . In the Treatise on Invertebrate Paleontology, Fisher
(1962) placed the hyolithids in the Calyptomatida, regarded as an extinct
class of the Phyl um Mol lus ca. Al th ou gh there is un cert ain ty as to the correct
taxo nomi c classification of hyolithids, most workers co nt in ue to favor affinity
with the m ol luscs ( M a l i n k y et al. 200 4) . S o m e h yo l i t hids m a y h av e b e e nsimilar to the present-day planktonic pteropod molluscs, but others were
probably ben tho ni c and cap abl e of mo ve me nt al ong the sea floo r (Fisher
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ARTHROPODS: TRILOBITES AND
OTHER LEGGED CREATURES 11
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In terms of sheer ab un da nc e, spe cies divers ity, and exploi tat io n of habitats ,
arthro pods rank as the most succe ssful of al l l iv in g ani mal s . Mo re than
750,000 speci es (mostly insects) in habi t a cast ran ge of en vi ro nm en ts on
land , in the sea, an d in fresh water. Li vi ng ar th ro po ds inc lu de the inse cts,
crus t aceans , h o rs es h o e crabs , arach nids , cent ip edes , and m i l l ip edes . D ur
ing the Ordovician, arthropods had not yet invaded the land, but tr i lobites
w ere a b u n d a n t and diverse in the sea, a l o n g w i t h the eu r y p t er i ds , ostra-
codes, and a few other minor groups.
Desp ite their be wi ld er in g variety of form , al l arthr opod s share certai n
ba si c features . L ike their c lo s e relatives, the a n n e l i d w o r m s , a r t h r o p o d s
have a seg men ted body. Unlike the ann el i ds , the bod y and its ap pe nd ag es
actually protected the trilobite's stomach. Transverse glabellar lobes and
rows indicate fused segmentation of the head region. A prominent pa
compound eyes usually flanks the glabella. A sinuous facial suture crosses
cephalon alongside the eyes, separating the lateral free cheeks from the f
che eks . The facial suture provide d a line of bre ak age across the ce ph
during molting. For this reason, isolated free checks are commonly foun
wel l as the c r a n i d i u m . a s ingle un it c o m p r i s i n g the glabella and fi xe d che
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A pa ir of ge n al spines is often d e v e l op e d at the po steri or corners of the c e
lon, as seen in Flexicalymene meeki, Isotelus maximus, and Cryptolithus
sellatus f r om the Ci n c i n n at i an .
T h e segme nts of the thorax were conn ec te d by a thin int egu men t
al lo we d th e trilob ite to flex its body and in ma ny cas es to ac hi ev e co mp
enrollment l ike a modern pi l lbug (an isopod crustacean). After death,cay of the articulating integument often re leased individual thoracic
me nt s that res emb le bracke ts ({) wh en preserved. The py gi di um is c
nio nlv preserve d as a single unit be ca us e its segm ent s were fused.
c o n s e q u e n c e of mol t i n g an d p ost-mor t e m d e c ay , t r i lob i te f r agme n ts
ab u n d an t , b u t c omp le te , ar t i c u la te d sp e c i me n s ar e u n c o mm on . T he
considerable debate about whether complete specimens represent tr i lob
b u r i e d intac t, b e c a u s e s o m e m a y hav e mo l t e d w i t h o u t the exoske l
b r e a k i n g apart. Usual ly , h o w e v e r , art i c ula te d s p e c i m e n s , particular ly
Figure 11.2. Ventral views
of three Ordovician trilo
bites, showing recon
structions of the append
ages. Anterior at the top
in each. A. Flexicaly-
mene senaria (Con
d) i f
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rad). B. Composite of
Isotelus maximus and I.
latus, (exopodites omit
ted because they are un
known). C. Cryptoli-
thus tessellatus Green.
From Raymond (1920, figures 9, 16, 20) and
reprinted by permission
of the Connecticut Acad
emy of Arts and
Sciences.
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inclina tion of the anterior cran idia l border. Ross was unab le to verify the
other differe nces asserted by Foerste. A morp ho me tr ic analysis co nd uc te d by
Danita Brandt (1980) in her unpublished master's thesis led her to conclude
that only a single species, F. meeki, is valid, and that both F. granulosa and
F. retrorsa should be synonymized with F. meeki as intraspecific variants.
More recent work by Brenda Hunda (pers. comm.) supports the recognitionof F meeki F retrorsa and F granulosa as valid species
Figure 11.3. A, B. Flexicalymen
(Foerste), University of
Cincinnati collections,
Maysvillian, Corryville
Formation, Hamilton Co.,
Ohio, enrolled specimen,
h li idth 8
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of F. meeki, F. retrorsa, and F. granulosa as valid species.
F lexicalymene is commonly found as isolated part ial exoskeletons (cc-
phala , crani dia, f ree che eks , thora cic seg me nts , pygidia) in Cin cin na tia n
l imestones; complete specimens are less common and are usual ly found in
shales as either enrol led or extended individuals . On rare occasions, these
trilobites can be found in great numbers in yellowish shales known as but
ter shales. O n e of the most prolif ic trilobite dis cov eri es eve r ma de in the
C inc innat ian w as in s uch a s h ale w it h in t h e lo w er R ich m o ndian W ay nes
vi lle F o r m a t i o n d u r i n g c o n s t r u c t i o n o f an a p a r t m e n t c o m p l e x a t B o u d i n o t
Av e nu e a nd W e s t w o o d N o r t h e r n B o u l e v a r d in n o r t h w es t C i n c i n n a t i in t he
1950s. Literally thousands of F lexicalymene were collected from two shale-
beds 2.5 and 3 fe et t hick , a n d as th e e x p o s u r e w e a t h e r e d , tr i lobites b e c a m e
perc hed on pedestals of cla\ for eas\ p ic ki ng (Cast er, pers, c om m. ; Sc hw ei n-
f urt h 1 958) . Ta p h o n o m ic s t udies o f o cc urr enc es o f abu nda nt , c o m p l et e
il bi i i i i h l i d i h h h l f
cephalic width 28 mm.
C. Flexicalymene ret
rorsa (Foerste), CMC IP,
Ferree Collection, Rich
mondian, Arnheim For
mation, Highland Co.,
Ohio, x 1.4. D. Flexicalymene gran(Foerste), Mark Peter col
lection, Edenian, Kope
Formation, Brown Co.,
Ohio, x 2.5.
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pace of the trilobite (see Figu re 14.2B; Osg oo d 1970). Al th ou gh no speci
mens of Rusophycus pu di cu m f rom the Ci nc in na ti an have bee n found inter
secting worm burrows, the digging activity is consistent with predation on
small, infaunal organisms (Fortey and Owens 1999).
Despite the excel len t s tate of preservation found in Ci nc in na ti an Flexi
calymene, r emnan ts of the app end ages have never bee n found. S tur mer andBergstrom (1973) carried out x-radiographic studies that revealed preserved
Figure 11.4. I s o t e l u s
maximus Locke. A. En
rolled specimen, CMC IP
2250, Riehmondian, Arn-
heim Formation, High
land Co., Ohio, x 1.3.
B. Large hypostome,
CMC IP 33067 M il
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Bergstrom (1973) carried out x radiographic studies that revealed preserved
appendages in some trilobites, but similar studies by Brandt (1980) and by
Hug hes and Co op er (1999) detected no evi den ce of appe nda ges in Flexica
lymene. The study by Hu ghe s and Co o pe r reveale d pyritized material con
centrated within the body cavity of Flexicalymene that may have originated
as deca yin g soft parts. An appro xima te idea of the nat ure of the appe nd ag es
in C inc innat ian Flexicalymene species can be gained from the restoration of
the closely related F. senaria ( f igure 11 .2A; Raymond 1920).
Iso te lu s is the other highly characterist ic and widely distributed tr i lo
bi te of the C i n c i n n a t i a n (Plate 7 ; f i g u r e 11 .4 ). F r a g m e n t s of this large tr i
lobite are found in every Cincinnatian formation, in both l imestones and
shales . Complete specimens are quite rare, but in certain shale horizons,
p art icular ly in t h e W ay ne s v i l l e F o rm at i o n, nu m er o us co m p let e s p ec i m ens
h av e been f o und ( Sc h um ac h er and Sh rak e 1 9 9 7) . Sp ec im en s o f Isotelus
CMC IP 33067, Maysvil-
lian, Clermont Co., Ohio,
x 7. C . CMC IP 51,
Robert Nestor collection,
Maysvillian, Corryville
Formation, Clermont Co.,
Ohio, x 1.8. D. Ap pendages on ventral side
of a complete specimen,
USNM 33458, Riehmon
dian, Oxford, Butler Co.,
Ohio, x 0.75. This excep
tional specimen was orig
inally illustrated by Mick-
leborough (1883). Photo
courtesy of Loren
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ens 1999). Fortey and O we n s me nt io ne d several other un iq ue features of
the Isotelus hypostome that suggest its function as a rigid platform like an
anvil for the manipulation of bulky food: its forked shape, and development
of anterior win gs prov ide a larger surf ace area , and the f ine raised rid ges
on the inn er surfac es of the fork co ul d m a ke i t eas ier to hold the prey rigi dly
usin g the app end age s. This hy post ome is the most heavi ly calcif i ed part ofthe Isotelus exoskeleton and is often found as an isolated component
Figure 11.5 . A. Deco-
roproetus parviusculus
(Hall), CMC IP 46429,
Edenian (figured in Davis
[1992, plate 2, figure 23]
as Proetus parviuscu
lus), x 7.7. B. Triar-
thrus eatoni (Hall) Steve
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the Isotelus exoskeleton and is of ten found as an isolated component
(Figure 11.4B).
I n t h e C inc innat ian, larg e Rusophycus burrows have been attributed
to Isotelus on the basis of their size (R. carleyi, see Os go od 1970). Spe ci
me ns often show not on ly furr ows cre ate d by the app en da ge s, but also
impressions of the cephal ic and pygidial margins and pleurae. A remark
able specimen shows a horizontal worm burrow apparently truncated in
the appr oximat e locat ion of tr i lob ites mou th (see Figu re 14 .2A; Brandt et
al. 1995 )—a trace fossil reco rd in g the very act of pre dat ion . T h e trilobi te
evidently du g and drew itsel f do wn into a semi- cohe sive m ud subs tratu m
so as to impress the marg in s of its car ap ac e like a coo ki e cutter. T h e trace
shows impre ssion s of the basal seg me nt s (coxae) of the app en da ge s that
probably seized the prey along the ventral midline and worked it toward
the mou th. A s ingle excepti onal sp ec im en pre serv ing the app end ag es of
thrus eatoni (Hall), Steve
Brown collection, J. Rush
collector, Edenian, Kope
Formation, Hamilton Co.,
Ohio, x 4.6. C. Cryp-
tolithus tessellatus
Green, University of Cincinnati collections, Ede
nian, Kope Formation,
Hamilton Co., Ohio, x
3.7.
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Plate 1. Ordovician conti nents and oceans . The reconstructi on shown is for the Middle Ordovician, abo ut 458 million years ago,
about 5.5 million years older than the beginning of the Cincinnatian. Compare to Plate 2. The position of Cincinnati (*) on the
paleo-continent Laurentia was south of the equator. Map courtesy of C. R . Scotese , PALEOMAP Project ( www.scotese.com) .
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Plate 5. A. Scolecodont,
one element of an an
nelid worm jaw apparatus,
Nereigenys alata Eller,
CMC IP 1952, Fairview
Formation, Dearborn Co.,
Indiana, x 70. B. Cono -
dont, one element of anapparatus, Phragmodus
undatus Branson and
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undatus Branson and
Mehl, CMC IP 50705,
Kope Formation, Campbell
Co., Kentucky, x 227.
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Plate 9. Living echino de rm s. A. Stalked crinoids (sea lilies), Neocrinus decorus
Thomson, northeastern Straits of Florida, height about 1 m, 420 m depth.
B. Unstalked crinoid (feather star), Pontiometra andersoni (P. H. Carpen
ter), Palau Islands, 4 m depth, arm length about 12 cm. C. Ophiuroid
(brittle star), Ophiothrix sp., Caribbean Panama, disk diameter 5-10 mm.
D. Echinoid, Strongylocentrotus franciscanus, San Juan Islands, Washing
ton, diameter about 15 cm. E. Asteroid (sea star), Fromia nodosa Clark,
Seychelle s, Indian Ocea n, arm leng th abou t 40 mm. F. Holothuroid (sea
cucumber), Cucumaria miniata (Brandt), San Juan Islands, Washington,
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height about 15 cm. A. by Charles G. Messing, all others by D. L. Meyer.
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Plate 14. The Cincinnati an, by John Agn ew , 2007.
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however, pointed out that very little flow could have passed throug
minute pores and thus it is more likely that the pores had a sensory fu
to orient the animal into the current. It this is true, the pores may hav
the sites of sen sor y hairs that wer e not pres erv ed.
Triarthrus i s restr icted to the lowe rmo st Ko pe fo rm at io n but is s
cant for several reasons (f ig ur e 11 .5B). Triarthrus is the last of the
t r i lobi te s that w e r e p r omi n e n t d u r i n g the Cam b r i a n . Pyr i t i ze d sp e c
from the Up pe r Or do vi c i an Utica S ha le of N e w York are am on g the
l l d il bi f h i h d i l d i f h
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w e l l -p re se r ve d tr ilobites , f r om w h i c h d e t a i l e d r e c o n s tr u c t i o n s o f th
pen da ges and internal soft ana to my have been ma de (Cisne 1970;
t i n gton an d Al m on d 1987) . In the Ci n c i n n at i an , tw o sp e c i e s , T. eato
T. spinosus, are rec og ni ze d (B ab co ck 1996a) but preserved app en
have not be en foun d. Th e structure of the app end age s in Triarthrugests that it was a particle feeder, sorting food particles with the th
appendages, which then passed them toward the mouth along the v
axis (Fortey and O w e n s 1999). Enla rg ed, spin e-be arin g basal l im
men ts (g natho bases) ben eat h the ce ph al on acted l ike jaws to proce
food and transfer it to the mo ut h. T h e restriction of Triarthrus to th
shales of the lower Ko pe fo rm at io n, and i ts do m in an ce in some thin
b o t h s u g g e st that th is trilobite was u n i q u e l y ad apted to d e e p e r water
t l i th l i d t i l bi t (F t d O
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basis o f a d d i t i o n a l , a lb eit f r a g m e n t a r y d is cov er i es . A truly p h e n o
discov ery ma de in 1938 of exce pti onal ly well-preserved and nearly
plete specimens from a single bed in the uppermost Cincinnatian El
For ma t i on of the Ri c h mo n d G r ou p i n Ad am s Co u n t y , Ohi o , le d to a
unde rst and ing of the ani mal (F igur e 11 .8B). This material , which inc
male and female specimens, became the basis for a new species, M
ensis, descri bed by Ke nn et h F. Ca st er and Frik Kjell esvig -Wae ring inO n e additional eurypter id species, Eocarcinosoma batrachophthalmus
des cri bed from this bed on the basis of an isolated pr oso ma.
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des cri bed from this bed on the basis of an isolated pr oso ma.
Megalograptus ohioensis was one of the largest creatures in t he C
natian sea floor co mm un it y, re ac hi ng a leng th of over 50 cm. The fir
of app end ag es, the che lic era e, is smal l and located bene ath the head
next thre e pairs of ap pen da ge s bear wel l-d evel ope d spines (Fig ure 11.9
third appendages are most striking for their length and long spines di
toward the mid li ne (Fi gure s 11. 9C, D). E xactly how the eurypterid
these spiny appe ndag es is uncerta in. Cas ter and Kjel lesvig-Waering c
ered Megalograptus to have been a predator, and thus the appendages
had so me fun cti on in gr aspi ng prey. T h e basket-like structure of th
spines of the third appe nda ge s sugges ts that the ani ma l mi ght hav e
them through the sediment in order to extract prey in a sieving fashion
a few othe r eurypt erids have similar long spiny app end age s. Tubular ca
terids lived in so me wh at restricted or aty pica l mar in e en vi ro nm en ts , the
Cincinnatian occurrences argue strongly for association with the normal
marin e biota. Their chit inou s inte gum ent ma y ac co un t largely for their
rar ity in the Ci nc in na ti an . Th e extraordina ry quali ty and quant ity of eu-
rypterid preservation at the Adams County site may have resulted from
smothering of the marine fauna by volcanic ashfal l , because the 15 cm-
thick shale within which the fossils were concentrated was found to contain
b en ton i t ic clays (Caster and K j e l l es vi g -W ae r i n g 196 4) .
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Neostrabops
In 1952 Cas ter and Ma ck e desc ribed what they term ed a mav eric k mer os to me ,
Neostnibops martini, from a single specimen found in the Maysvillian Corry-
vi ll e fo r mat io n in C l e r m o n t C o u n t y , O h i o ( f i g u r e 11 .8D). Th is fossil co u ld
be ta ke n to be a tr il ob it e, but la ck s the ch aracteri stic lengthw ise di vi si on in to
three distinct lobes. It also resembles the aforementioned eurypterids in hav
ing num ero us narrow segm ents, alt houg h it lacks any demarca tio n of pre- and
postabdomen. Neostrabops doe s resemble oth er arth ropo ds kn ow n as aglaspids
from the Cam br ia n. Agl aspi ds are regarded as early offshoo ts of the evolut ion
ary lineage of mod ern horsesho e crabs, the well-k nown Limulus.
w i t h i n the re st o f t he C i n c i n n a t i a n was no t stud ied. B e r d an (19 84) re
on ostr acode s of the order Lep erd iti cop ida from the Mi dd le and
Or do vi c i an of Ken tu ck y and v ic inity. Th es e ostracod es are note wor
their s ize (som etim es > 1 cm long) and high ab un da nc e in single b
f ine-grai ned l im est one. T h e oc cu rr en ce of leperd iticop ids is restr i
f ine-grained l im est one fa de s depos ited in extrem ely shall ow subt
i n te r t i d a l e n v i r on me n ts p ar t i c u lar ly w e l l kn ow n f r om the M i d d le O
c i an Hi g h Br i d ge G r o u p of K e n t u c ky (Cr e ssma n an d N og e r 1976
un iq ue fa cies is absent from t he dee pe r water fa de s of th e lowe r and
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un iq ue fa cies is absent from t he dee pe r water fa de s of th e lowe r and
Ci n c i n n at i an b u t r e c u r s i n the Ri e h mo n d i an S u n se t M e m b e r of th
he im F orm ati on , in wh ic h four specie s are foun d.
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Figure 12.1. One skeletal
element of a modern
crinoid, showing the
porous microstructure
(stereom) typical of all
echinoderms. The arm of
a crinoid is composed of
a series of these ele
ments, connected by
muscles and ligaments.
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g
Comactinia sp., Carib
bean. Scanning electron
micrograph, x 79.
ECHINODERMS: A WORLD
UNTO THEMSELVES 12
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Echinoderms are among the rarest and most sought-after fossils in the Cin
cinnatian rocks. Not only are they complex in form and structure, but they
also possess a certain beauty and mystery that never fail to attract interest.
A n y o n e w h o ha s visi te d the seashore is familiar w i th l iv ing e c h i n o d e r m s such
as sea stars or starfish (asteroids), sea urchins, and sand dollars (both echi-
noids) (Plate 9), Ot he r l iv ing ech ino der ms found in deeper marin e waters are
the sea lilies and feather stars (crinoids), brittle stars (ophiuroids), and sea
cucumbers (holothuroids) (Plate 9) . T h e r e ar e ab ou t 6650 l iv ing species of
echinoderms, and over 3500 genera and 13 ,000 described fossil species.
The Ordovician Period marked a very significant time in the evolutionof ech ino der ms , bec ause m any dif ferent major groups (usually regarded as
I . . here salute the
noble echinoderms
as a noble group es
pecially designed to puzzle the zoologist.
L. H. Hyman 1955, vi
as sea ur ch in shells as exter nal , but in oth er case s, such as the arms
sea stars, the skeletal components, cal led ossic les, are c lear ly intern
neath a leathery "skin." In addition, because i t is truly mesoderm
ec hi no de rm skeleton has a uniq ue micr ost ruct ure not found in an
an im al group. Th e calc i te is for med aroun d me so de rm al ce l ls into
tricate three-dimensional latticework called the stereom (Figure 12.1
et al plates, spines, or ossicles thus have a highly porous structure inover 50 perc en t of the vo lu me ca n be taken up by pores . In life the
ing ce l ls occupy these pores, but after death, the ce l lular material
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leaving the porous skeleton. Buried in sediment, these pores are
infilled with secondary calcite, and the entire skeletal plate displa
typi cal rh om bi c c le ava ge of calc i te . Of te n the micros truc ture i s st il l
in thin or poli shed sectio ns. Thes e features have ena ble d many foss
no de rm s to be correctl y identi f ied, even th ou gh their body form
cons idera bly di f ferent from a ny of the f ive fam iliar l iv in g groups.
Echinoderms are also pecu li ar in lac kin g structu res l ike a
head, eyes, or internal systems such as a blood circulatory system or
tory syst em. Ins tead, they are uni qu e in ha vi ng an internal sys
b r a n c h i n g vessels that c o n t a i n n o t b l o o d , b u t a w atery fl ui d that c ir
dissolved oxygen and dissolved wastes and pressurizes the vessels
selves The vessels term ina te in charac ter ist ic structur es calle d tu
have been limited by the available space around the mouth, and thus the
pent ame ral pattern may have been the most efficient solu tion . O n c e pe n-
tameral structure be ca me geneti cal l y pr og ra mm ed in Echinoderms, i t per
sisted even in grou ps that gave up the ance str al m o de of life to be co m e
mobile sea stars or sea urchins.
Despite their ma ny "alien" features, Echinod erms are signifi cant as on e
of the invertebrat e phyl a most closely related to our ow n, t he chorda tes . For
a long time zoologists studying the embryonic development of echinoderms
have recognized close similarities in the early development of echinoderms
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have recognized close similarities in the early development of echinoderms
and chordates. Both groups have symmetrical cell division in the fertilized
egg, indeterminate development (embryonic ce l ls are not preprogrammed
to form a specific adult tissue), and the internal body cavity, the coelom,
forms in the sam e way in the emb ryo . Rece nt studies of mo le cul ar compo sition of ani mal phyla demo nstr ate that echinode rms are mu ch mo re closely
allied to hem ich ord ate s and cho rdat es than to any oth er gr ou p (Raf f 1996).
Co mp le te fossil echinoderms are indeed rare fossils in Ci nc in na ti an
strata, but the ab un da nc e of their isolated skeletal co mp on en ts suggest s that
they were very co mm o n me mbe rs of sea f loor co mm uni ti es durin g the Or
do vic ian . The reaso n for their rarity as co mp le te fossils is fo und in the na tu re
of the ec hi no de rm skeleton, co mp os ed of myria d tiny plates or ossicles, all
Figure 12.3. Variable
preservation of crinoids.
locrinus subcrassus
(Meek and Worthen).
A. Articulated individual
with partially disarticu
lated sections of stalk,
and matrix of disarticulated skeletal compo
nents. Upper Ordovician,
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Corryville Formation, Cin
cinnati, Ohio. University
of Cincinnati collections.
B. Two articulated
crowns, detached fromstalk, oriented parallel
but in opposite direction,
preserved on base of
bed. Upper Ordovician,
Corryville Formation,
Clermont Co., Ohio. CMC
IP 44362. Scale in mm. Bin upper right denotes
Figure 12.4 . Reconstruc
tion of Glyptocrinus
decadactylus in life po
sition. Calyx is in a hori
zontal position, with the
arms splayed into a filtra
tion fan. By analogy withliving crinoids, current
flow was from left to
i h D i b J h
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right. Drawing by John
Agnew.
Figure 12.5. Cincinnatian
crinoid columnals and
holdfasts. A, H. Cincin-
naticrinus varibrachialus
Warn and Strimple; A.
Articular surface x 9.6;
H. Lateral view of mature
section x 5.9. B, I. Ect-
enocrinus simplex
(Hall). B. Articular surface
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y.9.6; I. Lateral view*
12.6. C, J. locrinus sub-
crassus (Meek and
Worthen). C. Articular
surface x 5.2; J. iafer a/
view x 4. 4. D, G, K.
Glyptocrinus decadacty-
lus Hall; D, G. Articular
surfaces of internodal,
nodal respectively, x
4.4; K. Lateral view x 6;
note three cycles ofinter-
nodals E Merocrinus
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Figure 12.11. Cincinnatianrhombiferans. A, B. Le-
padocystis moorei
(Meek), CMC IP 24680,
Elkhorn Formation, Pre
ble Co., Ohio, x 4.3.
C. Cheirocystis fulton-
ensis Sumrall and Schumacher, holotype, CMC
IP 50402, Kope Forma
tion, Bracken Co., Ken
of cri noi ds, the disparids, the cla did s, and tlic cam cra tcs . Dispa rids h
small, monocyclic cup- or howl-shaped calyx with branching arms th
not be ar pin nu les . An elon gat e, plated tub e, the ana l sac. is often p
b e t w e e n the arms and ha s the anal o p e n i n g at its end. The most c o m
Cincinnat ian cr inoids , Cincinnaticrinus, Ectenocrimts, and locrinus are
parids (Figures 12.6,12.7; Plate 10). Cincinnaticrinus and probably Ect
mis had a unique, button-like holdfast composed of main tiny plates, found attached to brachiopod shells and other hard substrata. Before i
re co gn iz ed that this holdfa st be lon gs to these type s of crin oids , if was
the name Lichenocrinus with numerous species (Figures 12 5L N;
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, C ,
tucky, x 3.5. D. Recon
struction of late
Riehmondian sea floor of
southeastern Indiana and
southwestern Ohio,
showing Lepadocystis
moorei (A) attached to
bryozoans (B), brachio-
pods, Zygospira mod-
esta (C), and an edrioast-
eroid, Carneyella sp. (D).
A, B, D from Kesling and
the name Lichenocrinus with numerous species (Figures 12.5L-N;
1929; Warn and Strimple 1977). Quite often in paleontology isolated pa
one organism are described as distinct species before sufficiently wel
served fossils are found that reveal the entire animal.
Cincinnaticrinus an d Ectenocrinus are frequ ent ly foun d tog eth
the Ko pe For ma tio n, whe re their disa rticu lated col un ma ls can form
limestone beds. Sometimes the articulated stalks are packed tightl
gether l ike logjams where the collector should look closely for the s
deli cate crow ns ( Plate 10B). These "log ja ms " were probahlv form ed d
ancient storms that disrupted the sea floor.
locrinus is another common disparid crinoid in the Cincinnatian
is larger and more robust in structure than ( l incinnaticrinus and Ect
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Figure 12.12. Edrioaster-
oid Isorophus cincinna-
tiensis (Roemer), recon
structed as in life, with
food grooves open for
feeding. Drawing by John
Agnew.
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Figure 12.13. Cincinnatian
edrioasteroids A,B. Isorophus cincinna-
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Figure 12.17. Cincinnatianstylophoran carpoids.
A. 1 -6 . Enoploura p o -
pe/ Caster. 1 -3 . Holo
type, CMC IP 25993,
Corryville Formation,
Clermont Co., Ohio.
1. Ventral view. 2. Dor
sal view. 3. Lateral
view, x 2.3. 4- 6. Para-
type, CMC IP 25257, Cor
ryville Formation Hamil
ci nn at ia n in whi ch all of the arms are in the proces s of regen erati onure 12.6D). T h e y co nc lu de d that the regene rati on foll owed the loss
arms to an attack by an unknown predator. Ausich and Baumiller (
reported r egen era tio n of a col um n attac hed to the holdfast Licheno
dubius (known to be the holdfast of Cincinnaticrinus or other dispa
Do no va n and S ch mi dt (2001) il lustrated plur ic olu mna ls (sections of s
co l umnal s ) o f Cincinnaticrinus sho wi ng rou nde d overgrow ths of one
T h e y sugge sted that the over grow ths forme d after decapi tat ion o
cro wns by preda tion, le avi ng a "hea dles s" co lu mn . In l ight of the
kn ow le dg e of pred atio n dam ag e in l iving crin oids , it is most l ikely
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ryville Formation, Hamil
ton Co., Ohio. 4. Dorsal
view. 5. Lateral view.
6. Ventral view, x 2.3.
1 - 6 from Caster (1952,
plate 1). B. Reconstruc
tion of E. popei, from
Parsley (1991, text-figure
1). All reprinted by per
mission of the Paleonto-
logical Research
Institution.
predators also cau sed loss of cr ow ns and ar ms in Or do vi ci an crinoi d
tho ugh the identity of the culprits remain s unce rtai n.
R h o m b i f e r a n " c y s t o i d s "
Rhombiferans are stalked echinoderms that appeared in the Early Or
cian and b ec am e ext inct by the Late Devo ni an . The term "cystoid"
like) refers to the pla ted t he ca that has four to five cir clets of large
arra nged in a pen tame ral pattern. Rh omb ife ran s were original ly a
gro up of the cystoids but have be en elev ated to a separate class. T h e y
as suspension feeders , but unl ike crinoids, the feeding appendag
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(the n am e me an s "seated-star"). T h e sea star res em bla nc e derives fro
five usually curving food grooves or ambulacra! tracts that converge o
central mouth (Figure 12.12). Thin, overlapping calcitic plates called
ambulacrals take up the space between the ambulacra. A more rigid
of mar gin al p lates forms the r ing. Usua lly the amb ula cral and int er
lacral plates appear to have collapsed inside the marginal ring; thus
as sum ed that the multip late d theca was flexible in life. Rarely, unco ll
or "in fla ted " sp ec im en s are foun d that reveal ho w the an im al proba b
peared in l ife . Mos t Ci nc in na ti an edrioasteroids had a low do me shap
so me were mo re cylin dric al ( Sum ral l 1994). A total of six gen era and
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species are known from Cincinnatian strata (Figure 12.13).
Be ca us e edrioast eroid s are ext inc t, their mo de of life must be in
by a n a l o g v to l i v i n g e c h i n o d e r m s . Fdri oasteroids ar e always fou nd att
to a hard surface such as a brachiopod shell , bryozoan, or hardground
lower surface was plated in some species, but in Cincinnatian specie
parently only a soft tissue membrane served to adhere to the substr
possibly in the way sea anemones attach by their pedal disk. Some s
actually cemented to the substratum like a barnacle , but those adheri
a basal me mb ra ne ma y have be en capa ble of l imited mo vem en t. Be
the mouth and ambulacral tracts are directed upwards in edrioastethe y appa rent ly lived as passive filter feeders Th e ambu la cra l groov
over thirty millimeters in diameter, Bell (1976) was able to determine how
several species changed in morphology during growth. This information was
used by Me ye r (1990) to study the pop ula tio n pa lc oe co lo gy of three different
species foun d on a single pave men t. Sma ll i ndivi duals of the c o m m o n spe
cies Isorophus cincinnatiensis (Fi gure s 12.13A, B) clustered nea r the mar gin s
of articula ted (likely living) shells of the host bra chi opo d Rafinesquina. T h e
small edrioasteroids may have lived in a commensal relationship with the
l iving brachio pod, ta king advantage o f the feed ing currents generated by the
brachi op od and protect ion a lo n g the o v e r h a n g i n g m a r g i n of the ho st shell.
Because a single large Isorophus occupies almost an entire brachiopod shel l ,
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Because a single large Isorophus occupies almost an entire brachiopod shel l ,
it is l ikely that either mortality or relocation to avoid overcrowding depleted
the juvenile clusters. In time, the edrioasteroids may have outlived the bra
chiopod host, because the larger individuals are found on disarticulated,
abraded ( hen ce dead) host shells. Pa vem ent s foun d at different stratigr aphic
horizons in the Cincinnatian have dif ferent edrioasteroid species popula
tions (Sumrall et al. 2001).
A s t e r o i d s
Sea stars are exceptionally rare fossils in the Cincinnatian; often, specimens
h d i b h f di d N h
prey, and both may have produced the rare star-shaped burrows iC i n c i n n a t i a n c a l l e d Asteriacites (see Figure 14.3D; Branstrator 1975)
Ophiuroids
T h e ophiu roi ds (brittle stars or serpent stars) are dis ting uis hed from
asteroids by having the five arms sharply differentiated from the c
disk (Plate 9C ). The ar ms are qui te flexible be ca us e they are co mp os
a series of vertebra-l ike ossicles co nn ec te d by mus cle s and lig amen ts.
uroids can move quite rapidly on the sea floor by lashing the arms
some can flex the arms vertically as well The arms are equipped with
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some can flex the arms vertically as well . The arms are equipped with
feet that are used to gather organic particles either directly from the
ment or as suspended particles. Ophiuroids are usually considered
deposi t feeders, but so me are also cap ab le of susp ensi on fe edi ng andp r e d a t i o n . Taeniaster spinosus ( B i l l ings ) occurs in t he Cinc inn
(Ilotchkiss 1970; Figure 12.16B), and like asteroids, it is very rare. M
op hi ur oi ds ca n live in ver y de ns e ag gr eg at io ns on the sea floor. Rarely
have been found in t he Cin c in nat ia n bear in g dens e as sembl ages o
niaster, su gge sti ng that agg reg ati on be hav ior was achie ved in this very
me mb er of the group . In the only other kn own C in cin na ti an ophi
Protasterina flexuos a, pyritized tube feet have been reported in a spe
Because there appears to have been very l i t t le space between the
plated surfaces of the l iv ing cvcloeystoid, the animal somewhat resembled
a tambourine rather than a drum. Lacking internal space for organs, cyclo-
cystoids were restricted to feeding on minute organic particles. Smith and
Paul conclude that the particles were collected at the cupules by ciliary
action and conveyed via the ducts to the radial grooves that converged on
the central mouth. Although tube feet are not preserved, Smith and Paul
suggest that tube feet could have emerged from pores between plates on
the ventral surface and provided a means of locomotion. The eyclocvstoid
thus moved over the substrate and gathered organic particles using ciliated
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g g p g
cup ule s. Oth er specia lists, how eve r, do not ac ce pt the life orie ntat ion fa
vor ed by S m it h and Paul, and instead regard the opp osit e si de as l o w er m o s t
(Sprinkle, pers . co mm. ) Cycl ocy sto ids f irst appe ared in the Farlv O rdo vi
cian and last in the Late De vo ni an or Karlv C ar bo ni fe ro us (Sm ith and Paul
1982). A total of nin e genera an d forty- one speci es have b ee n des cri bed . In
the Cincinnatian, three genera and f ive species are known.
S t v l o p h o r a n s
f o r many paleontologists , s tvlophorans surpass even the cyclocystoids as
f h bi f il hi d F i i f i l i
groove entered an internal mouth, and wastes were emitted throuanal op en in g be tw ee n the spines. Propo nent s of the ealc i chor date
pretation of stv lo phoran s regard the app en da ge to be a true, wrigg lin
w i t h the m o u t h l o c a te d a t the o ppos i te e n d o f the t he c a . S t u dy o
p r e se r v e d ske le ta l mi c r ostr u c tu r e i n Ci n c i n n at i an V.noploura by C
and f isher (1981) revealed c lose similar i ty to typical echinoderm stc
furth er sup por tin g the c lassi f ication of stv lop horans with ech in od
Some enigmatic fossi ls cal led maehaeridians might also be re lated t
lophorans (see chapter 10).
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GRAPTOLITES AND CONODONTS:
OUR CLOSEST RELATIVES?
Graptol ites are among the most dist inct ive foss i ls found in Cincinnatian Graptolites
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Graptol ites are among the most dist inct ive foss i ls found in Cincinnatian
strata and are also uniquely s ignif icant . Craptol ites are commonly pre
served in shales in a highly f lattened condition, appearing like black pencil
mar kin gs with a saw- toot hed mar gi n (the n am e graptol ite in fact me an s"written stone"; Figure 13. lC) . In some Cincinnatian l imestones graptol ites
can be p res erv ed in an unco m p act ed , t h ree- dim ens io nal co ndit io n. Be
cause their skeletal s tructure (periderm) is organic these "inf lated" grapto
lites ca n be etc hed free of the matr ix usin g acid to reveal e xce pti ona l struc
tural details (F ig ur e 13.1B). Grapt oli tes rep resen t the skeletal she ath of a
colonial, soft-bodied marine invertebrate whose soft parts are not preserved.
Graptol ite colonies ex isted as f ree-f loat ing plankton (order Graptoloidea)
Graptolites
Figure 13.2. Cincinnatian
conodonts. Those shown
are among the most
common forms. All are
from the lower Riehmon
dian Stage near
Brookville, Franklin Co.,
Indiana. A, G, H. Plec-
todina tenuis (Branson
and Mehl). A. Pb ele
ment. G. M element.
H. Sc element.
\ l t h o u g h several spec i e s
of graptoli tes occur in the Cincinnati
the Ci nc in na ti Ar ch region, only a few are c o m m o n (Bergstrom 1997)
most c ommon an d c har ac te r i s t i c Ci n c i n n at i an gr ap tol i te i s Geniculo
tus (identified as Climacograptus in older l i terature; f igures 13.1B, C)
grapt olit e has a hiserial rh ab do so me and is very char acte risti c of the
fo r m at i o n , w he r e aggr e g at i on s of s t i pe s of te n oc c u r i n p ar a l le l a l i gn
on bed di ng surfa ces (f i gu re 13.1C). Tw o spec ies range from the
through Fairview formations (Bergstrom 1996a). ' Iwo hiser ial grapt
o c c u r i n t h e A r n h e i m f o r m a t i o n ( R i e h m o n d i a n ) : Orthogrciptus qucid
cronatus an d Arnheimograptus anacanthus (see Bergstrom 1996a). Se
species of Mastigograptus, a delicate , bush-l ike dendroid graptoli te ,
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B. Oulodus oregonia
(Branson, Mehl, and
Branson), Pb element.
C, D. Amorphogna-
thus ordovicicus Bran
son and Mehl. C. Sc
element. D. Pa ele
ment. E. Drepanoisto-
dus suberectus (Branson
and Mehl). F. Phrag-
modus undatus Branson
species of Mastigograptus, a delicate , bush like dendroid graptoli te ,
i n the K op e , Ar n h e i m, an d W ay n e sv i l l e for mat i on s .
The zoo lo gi ca l affinities of grapto lites were for m any years a m o n
most c ha l l e n g i n g p r ob le ms i n p a le on tol ogy . G r ap tol i te s w e r e c las
wi t h several dif ferent g r o u p s , i n c l u d i n g c e p h a l o p o d s , c n i d a r i a n s , b r
ans, and hemichordates, or were considered to be unrelated to any l
gro up (Bu lin an 1970). Res ear ch by the Polish paleont olog ist Rom an
zlowski (1966) noted several similarities between graptolites and the
hem ich ord at es cal l ed ptero bran chs that arg ue stronglv for a c lose e
tionary re lationship. Pterobranchs are a group of small , marine, tube-d
ing invertebrates that are classed together with the acorn worms in
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se c t i n g mi c r osc op e b e c a u se of the i r u n i q u e for ms an d a b e au t i fu l acolor (Plate 5B).
Co no do nt s have a wide range of shapes, i nclu ding single cone s, m
pronged "teeth," serrated blade-l ike shapes, and so-called platform
forms (Figure 13.2). Most conodonts have a tooth-like appearance, leadi
the assu mpt ion that they wer e used as teeth . How eve r, cono don ts also
rege nerat ion. This s ugges ts that at least so me con odo nt s were emb ed d
soft tissue by which they were secreted. Individual conodonts, called
ments, were given species names by earlier workers. However, the disc
of rarely preserve d ass emb lag es of different elem ent s in rock m atrix or
together led to the recognition that elements were arranged in bilat
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g g g
sym met rica l assemb lages of pairs of e lem ents , eac h of whic h is cal l
apparatus. Although few apparatuses are preserved intact, it has been
ble to d e t e r m i n e w h i c h e l e m e n t s fou n d in a s am pl e lik ely fo rme d an
ratus on the basis of statistical analysi s of the ratios of co m m o nl y asso
elem ents . M od er n taxono mists of con odo nts attempt to include the
seven different elem ent s of a given appa ratus u nde r a single species na
Be ca us e the tooth-l ike e leme nts apparent ly were the only miner a
parts of the org ani sm, th e identi ty of the con odo nt -be ari ng orga nism
the nat ur e of its soft part ana to my ha ve be en a m o n g the great myster
paleo ntol ogy A major brea kth rou gh c am e in 1983 whe n a fossil of a
Figure 13.3. Reconstruc
tion of the Ordovician
jawless fish, As t rasp is
desiderata Walcott,
from the Harding Sand
stone of Colorado.
Length about 13 cm.
From Cowen (2005, 86,
figure 7.4). Reprinted
with permission of Black-
well Publishing.
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preserved. A similar fossil f ish from Ordovician rocks in Bolivia showthese early fish had blunt, rounded heads, an elongated fish-like shap
a tail fin, but lacked bony jaws and separate fins. These jawless fish are
agnathans, but other Ordovician fossils represent the earliest jawed f
gn at ho st om es (S an so m et al. 2001). Thu s, a variet y of early fish had a
evolved by Cincinnatian t ime, but they are unknown from the Cinc
regio n. Ha d these early fish be en present in the Cin ci nn at ia n sea, it
seem reasonable to expect their mineralized plates to be preserved
limestones or shales. Although their absence may indicate that then - pre
an environment not represented in the type-Cincinnatian, the potent
their eve ntu al discover) shoul d not be over loo ked.
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their eve ntu al discover) shoul d not be over loo ked.
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TYPE-CINCINNATI ANTRACE FOSSILS:
TRACKS, TRAILS, AND BURROWS
The Ci nc in na ti an is re no wn ed for its ab un da nc e of well- preser ved shells
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and skeletons of Orclo\ ician marine invertebrates, and because these fossils
represent the rema ins of lon g-de ad or ga nis ms , at first gl an ce one wou ld not
expec t them to yield mu ch i nfor mat ion abo ut the activity and beh avi or ofthese anim al s dur ing life. Of cou rse , we can de du ce a great deal a bou t the
life habits of Or do vi ci an an im al s direct ly from the mo rp ho lo gy of shells
and skeletons (body fossils) by comparisons to their living relatives, but a
va st ran ge of e v i d e n c e abo ut a n c i e n t b e h av i o r also c o m e s from a c o m
pletely different source, namely the trace fossils that are both abundant and
diverse in Cincinnatian strata.
Trace fossils are evi den ce of the activities of anc ien t orga nis ms pre
Figure 14. 1. A. Repich-
nia of the trilobite Isote
lus, Asaphoidichnus
trifidum Miller, CMC IP
37569, Edenian, Kope
Formation, Cincinnati,
Ohio, x 7. B. Repichniaof the trilobite Cryptoli
Linnaean binomial system was developed for trace fossils because they once thought to be body fossils, and this procedure has persisted (Sim
1975). Ideally, the ichnogenus might be defined to represent a parti
behavioral pattern w h i le the i ch n o sp ec i e s rep res ents variations on th is
tern, a lthough this procedure has not been uniformly applied (Bro
1990). Trace fossils provide information about unpreservable soft part s
tures of an im al s that are kn ow n as skeletal fossils. I n the Ci nc in na ti an ,
ex am pl es of this are the varieties of Rusophycus, the resti ng trace of trilo
with impressions fo r m e d by the d i g g i n g act iv ities of the leg s (F ig ur e 1
Tr ac e fossils also au g m en t ou r record of diversity by preser vin g activiti
entirely soft-bodied species that are otherwise unknown in the record.
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T h e greates t sig nif ica nce of trace fossils is, how ever , the inform
they yie ld abou t the beh avi or of long -dead anima ls. The G e rm a n pa
tologist Ru dol f Richt er pione ered the analysis of anci ent beha vior
trace fossils by studying traces made by shallow marine organisms in
cent sed im en ts of the No rt h Sea. In ma ny cases mo de rn tracks and
could be compared to fossi l ized traces. A remarkable book by Wil
Sch afe r (1972) s um ma ri ze s the work of Richt er and m an y subs equen t
m a n stu dent s of the Nor th Sea traces in Eng lis h. Ad ol f Sei lac her
classified trace fossils into behavioral categories that facilitated the i
preta tion of an cie nt en vi ro nm en ts on the basis of trace fossil assembl
of Paleodictyon from other localities. Al th ou gh it may be the impres sion of a
patterned object being rolled along the substratum, Osgood concluded that
the Ci n c i n n at i an Paleodictyon is a trace fossil. Seilacher (1977) interpreted
patterned traces of this type to represent complex burrow systems rather than
gra zin g patterns. These bur row netwo rks are used by so me infauna l orga n
isms for the "fa nn in g" of mic rob es benea th the sea floor.
Fod ini c lm ia are (c edin g traces of deposit feeders operat ing from a
f ixed burrow. The most co m m o n Ci nc in na ti an fod inic hni a are the var iet
ies of the b ran chi ng burro w Chondrites as descr ibed by Osgood. Chon
drites, ty pe-A ap pea ri ng on the verti cal edg es of beds of fine -grain ed car
b t bl t l t ( 8 di t ) t i
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bon ates , resembles n arro w rootlets (0 .8 mm average diameter), sometimes
pen etr at ing the entire thi ckn ess of a bed (P'igure 14.3A). On be dd in g
plane s, this form o ccu rs as a circ ula r patte rn of closel y spa ced h oles. Chon
drites, type-B can be seen on both bedding planes and vertical sections.
C o m p a r e d t o Chondrites, type- A, typ e-B has tubes of greater dia met er (1-4
mm), and branches that propagate to a greater extent horizontally than
vertical ly ( F i g u r e 14 .3 B) . Chondrites, type-C occurs as densely interwoven
radiating branches, e i ther paralle l or oblique to bedding (Figure 14.3C).
A n o t h e r c o m m o n an d interestin g C i n c i n n a t i a n trace classif ied b y O s
good as fodinic lmia, or possibly domichnia, is Trichophycus venosum Mil ler
Diplocraterion i s the most c o m m o n d omi c hn i a l t r ac e foss i l o f the Cnatian. Osgood (1970) recognized three ie lmospccies of Diplocrateri
seen on vertical joint surfaces, /). cf. luniformis (Blanckenhorn) has a
I l-shape with a rounded base. Narrow, elongated slots on the upper s
of a bed repr esent the base of the 11. In the mor e c o m m o n /). cincin
sis (Osgood) the U expands with depth in di f ferent ways (f igures
14.4 \i. f aint dow nw ar d- cu rv in g striations (Spreiten) connect the ar
the U. These so-called protrusive Spreiten may represent the dow
propa gati on of the bur row as the org ani sm grew. Alternatively, the
ism ma}' have burrowed deeper in order to maintain a constant dep
low a con sta ntl y er od in g sed ime nt- wat er interfac e. The reason for the
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expa nsi on is unclear . S om e expa nsi ons occ ur at an interface with c
material , yet others expand within the si l t-sized layer (f igure 14.4A)
led Osgood (1970) to suggest that the organism may have sensed a che
ch an ge in the sed ime nt before enc ou nt er ing a cha ng e in grain siz
spe cul ate that the exp ans ion of the U mi gh t have deve lop ed aft
w o r m - l i k e o r g a n i s m b u r r o w e d to its preferred d e p t h . Lateral e x p a n s
the U enabled the worm to grow in length while maintaining i ts de
T he thi r d Diplocraterion i c h n o s p e c i e s , D. biclavata (Miller), has
of short extens ion s deve lo pe d at the base of the U as blind po uc he sf l i l i f i h l l l d "
Figure 14. 2. A. Cubich-
nia and sderite impres
sions of trilobite Isotelus,
Rusophycus carleyi (J. F.
James), CMC IP 46411,
Maysvillian, Corryville
Formation, Clermont Co.,
Ohio, x 0.6. Note impres
sions of cephalic margin,
genal spines, pleurae,
pygidial margin, as well
as coxae (medial paired
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p
lobes, flanked by cres-
centic impressions made
by legs. Also, at top, note Paleophycus burrow
terminating at approxi
mate position of trilobite
mouth, with superim
posed scratchmarks.
B. Flexicalymene
meeki (Foerste), CMC IP37574 Maysvillian Cor
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a crus ta cea n that took adv an ta ge of the bry oz oa n host for prot ectio n
as for an eleva ted fe ed in g posi tion . Exc ava tio n of the interior of the br
b ran che s probably r edu ce d the struct ural integ ri ty of th e co lo nv and re
it more susceptibl e to brea kag e du rin g storms. Trypanites borings on
ans usually occur in clusters and were probably formed when dead,
colonies were exposed on the sea floor (Erickson and Bouchard 2003)
A n o t h e r typ e of t race f o u n d i n C i n c i n n a t i a n b r v o z o a n c o lo n i e s
act ual ly a bor in g but rather recor ds the pre sen ce of a soft- bodied or
that l ived as an en do sy mb io nt with in the brvo zoa n skeleton, term
cl au st ra ti on by Pal me r and Wi ls on (lcjHS) (fi g ur e 14 . 4 0 ; see Figure
Af te r se t t l e m e n t by a larva of the e n d o s y m b i o n t on to the l iv ing
b r o o a n o o e c i a gre a r o u n d the o r g a n i s m In c o n f o r m a t i o n to its
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b r v o z o a n z o o e c i a gre w a r o u n d the o r g a n i s m In c o n f o r m a t i o n to its
result ing in distinctive pits arranged in rows, named Catellocaula
by P a lm e r and W i l s o n (19S8). t i n l i k e b o r i n g s , the m a r g i n of these l ined by zooe cia l walls . T h e mor ph ol og y of the pits and their arra ng
in rows suggested a colonial , stoloniferous organism, most l ikely a tu
Tapanila (2005) has proposed that a new behavioral category, Imped
be used fo r s u c h cavit i es that l oc a l l y in hi bit the n o r m a l skeletal g r o
the host. Catellocaula vallata represe nts on e of the oldest kn ow n ex
of this end os ym bio ti c behavior .
Cubichnia are temporary traces made by mobile animals Althoug
vagile organi sms and suspension feed ers to seek shel ter in bu rrows. As depth
increases within the shallow n e a i shore /one , condit ion s of lowe r water mo ve
ment permit food particles to settle, and thus deposit feeding activity in
creases, resulting in fodiniehnial traces. In deep sea environments there is
little need for permanent shelter, and so dwelling and resting traces are not
form ed; instead, organi sms tend to gra ze the sedi men t surfa ce for food , creat
ing the meandering pascichnial traces. Kepiclmial traces are found in all
su b mar i n e e n v i r on me n ts , b e c au se or gan i sms ar e a lw ays goi n g some w he r e
and leaving trails, no matter what the setting! Seilacher's original concept of
trace fossil facics distribution has been widely tested, substantiated, and ex
p an d e d to i n c lu d e asse mb lage s c har ac te r i z i n g n on -mar i n e e n v i r on me n ts
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and partic ular substrata such as har dgr oun ds and woo d (Bro mle y 1990). C a n
the ichnofacies concept be applied to C in ci nn at ia n trace fossi ls , and what
can this tell us about the Late Ordovician marine environment?
Osgood (1970) listed thirty ichnogenera and forty-four ichnospecies
from the Cincinnatian. Recent additions and revisions bring the total to
thirty-four ichnogenera and fortv-seven ichnospecies (Holland 2005; Tapa-
nila 200^). The fo ll ow ing list sho ws the dist ribu tion of the se ich no spe cie s.
C u b i c h n i a 5
These are usually preserved as hyporeliefs . One such structure, "Bla
cus," prob abl y represe nts cast ing s of impr ess ions left by enroll ed tr
and current scou r aro und the m (O sg ood 1970). Ano the r , "Dystactop
is a fan-shaped pattern of f ine concentr ic r idges and grooves, and wa
th ou gh t to be an algal frond. Os go od co ncl ude d that i t forme d by r
of a cr inoid st em as i t was buried . At several Ci nc in na ti an local i t ie
fectly c irc ular impre ssions of co nce nt r ic r ings oc cu r on the upp er
of a bed (F ig ur e 14.4F ). Ca st er interpret ed these to be bo dy fossil m
a porpitid jellyfish and described them as Palaeoscia floweri (see
1942 ) . How e v e r , Osgood e xami n e d ad d i t i on al sp e c i me n s an d c on s
that thes e con cen tr ic r ings coul d have form ed un der the inf l uence
rents by rotational swe epi ng of so me kind of org anic dw ell ing tube e
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rents by rotational swe epi ng of so me kind of org anic dw ell ing tube e
ded in the sedime nt. Alternatively, so me l iv ing poly chae tes create a f
trace that resembles Palaeoscia, and thus Osgood re legated thesep e c u l i a r C i n c i n n a t i a n " t r a c e s " t o incertae sedis. Sta nle y (1986) sup
Osgood ' s asse ssme n t that Palaeoscia is a trace fossil, but controvers
tinues over interpretation of Palaeoscia and simi lar con cen tr i c r i
structures in the geological record (Ewing and Davis 1967, 274-275
et al . (2001) l isted Ci nc in na ti an Palaeoscia as a jellyfish, and asserte
Osgood was incorrect in regarding it as a trace fossil, but gave no ba
this evaluation
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Figure 15.1. Divisions of
type-Cincinnatian strata.
Geologists have tradi
tionally defined sedimen
tary rocks on the basis of
time, usually inferred
from fossil assemblages,
and on the basis of rock
type. The Cincinnatian
has been traditionally
divided into three stages,
shown at the far left, and
there is currently dis
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agreement over the rela
tive durations of these
three stages. A fourthstage, indicated by "G"
and called the Gama-
chian Stage, is not pres
ent in the Cincinnati area.
The divisions based on
rock type are shown at
the right. Most of those
PALEOGEOGRAPHY ANDPALEOENVIRONMENT
Steven M. Holland
Earth sc ientists reconstrucl conditions during the ancient past from a wide
i f h i f i l k d f i l A l h h l
15
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variet y o f ch i cs from m i n e r a l s , rocks , a n d fo ss i l s . A l t h o u g h n o p l a c e on
Earth today is exactly l ike the Cincinnati area during the Late Ordovic ian,
comp aris ons with mode rn envi ron men ts offer valua ble insights into the
interpretation of these c lues. O f mo de rn en vir on men ts , the Persian Gu l f
is perh aps the mos t simi lar to the Lat e Or do vi ci an of the eastern U nit ed
States in terms of its cl im at e, th e size of the sed im en ta ry basin , the gcntlv
dipping sea floor, the mix of carbonate sediment and clay, and the occur
renc e of storms that rework and depos it sed im ent .
T hr e e maj or oc e an s se p ar ate d the se c on t i n e n ts . T he Iap e tu s
separated Laurentia and Siberia-Kazakhstan from Baltica to the sout
Pale ote thys Oc e a n lay b e t w e e n G o n d w a n a an d the c on t i n e n ts of
and Siberia-Kazakhstan to the west. The massive Panthalassic Ocea
ered alm ost al l of the Nor the rn Hem is ph ere and wou ld have dwar
day's Paci f ic Ocean.
Global sea level was high during the Ordovic ian, and althou
position is difficult to constrain, the current consensus is that it was
200 meters hi gh er tha n prese nt-d ay sea level (see Fi gur e 1.3). Several
con tri but ed to such a hi gh pos itio n of sea level. Rates of sea floor spr
w e r e h i g h f o l l o w i n g the b r e a k u p of an older, L ate Prote rozoic su pe
nen t cal led R od in ia , ca us in g the averag e elevat ion of the sea floo
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hig her th an nor ma l. This raising of the "bo tt om of the buck et"
ocean waters to spill onto the continents. In addition, the lack of po
caps in m ost of the Ord ov ic ia n also wo ul d ha ve raised sea level rela
today because water in modern glacial ice caps such as Antarctic
G r e e n lan d i s p r od u c e d f r om sn ow ge n e r ate d b y e v ap or at i on f r o
ocean. Because of this high sea level , low-lying areas on the con
w e r e f looded with o c e a n w aters , m u c h l ike the f looding of the prese
continental shelves, but to a much greater extent. During much
Or do vi c i an , most of L am en ti a was su bm erg ed, with the exce ptio n
W a r m te m pe rat u re s at the poles in hibi ted the form at i on of ice ca ps ,such as today's continental glaciers on Antarctica and Greenland and the
sea ice over the Arc tic Oc e a n . In the last mi ll io n years of the Ord ov ic ia n,
a tmosp he r i c p CO 2 , levels dropped precipitously, triggering the rapid growth
of pola r glaci ers and a g eol og ica lly brief 160 met er global sea level fall. Th i s
fall in sea level drained the seas from the Cincinnati area, producing an
erosional div ision bet wee n the Or do vi c ia n and Si lur ian strata cal led an
u n c on for mi ty .
In addition to this end-Ordovician fall in sea level, evidence for six
cycles of global sea level ch an ge is preserved in the Ord ov ic i an near C i n
cinnati ( f igure 15 .1) . The evidence for these cycles comes from packages
of rock kn ow n as deposi t ional seq uen ces which are bou nd ed by unco nfo r
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of rock kn ow n as deposi t ional seq uen ces , which are bou nd ed by unco nfo r
mitie s, or surfaces that record the eros ion and wea th er in g of se dim en ts .
Eac h depo sit iona l se qu en ce begi ns with a relatively thin inter val of rock
that records local de ep en in g of the oc ea ns and end s with a mu ch t hick er
interval of rock that records progress ive sh al lo wi ng of the oce an s. Th es e
same sequences can be recognized across the United States and in Estonia.
The fact that these sequences are not just local features is strong evidence
that they reflect global sea level changes rather than local tectonic changes.
In the Cincinnati area, these si\ deposit ional sequences also contain evi
Figure 15.2. Destruction
of benthic communities
by storm-generated
waves and currents. Dur
ing calm, pre-storm con
ditions, benthic commu
nities of organisms
develop on the sea floor.
Under storm conditions,high winds generate
large waves that stir up
fine-grained bottom sed
iments into suspension.
Stronger wave and cur
STAG E III . CA LM CO NDI TI ON S, POST -STO
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Stronger wave and cur
rent forces can displace
benthic organisms, andsuspended sediment can
clog feeding and respira
tory mechanisms of or
ganisms, and even
smother entire benthic
communities. Mobile or
ganisms can escape if
b i l i t t
Cyc lic al ch ang es in the charact er of storm beds are well develo ped in
some Cincinnatian deposits, such as the Kope formation. At a broad scale,
these roug hly meter-thick cycle s consist of a mudst one- rich unit and a lime
stone-rich unit (see Fi gur e 4.6). T h e m uds ton e-r ich unit consis ts of 3—5 cm
beds of normally graded mu dst on e, with u n c o m m o n thin lami nated si lt st one
beds. Limeston e-r ich unit s consist of shel ly l i me ston e be ds , with a les se r
am ou nt of thin mud sto ne and siltstone beds. Th e alternation b et we en these
two units was originally thought to reflect changes in sea level, but recent
studies suggest that these cycles may instead reflect changes in the average
frequency and intensity of hurricanes over tens of thousands of years.
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Co mp ar e d to man ) mod e r n c ar b on a te se t t i n gs , Or d ov i c i an l i me ston e of
the Ci nc in na ti area is unu sua l in several regards. Mos t mo de rn and an
cient war m water car bon ate depos its cont ain a wide variety of grai n type s,
inc lu din g skeletal grains (the shells of org ani sms ), ooi ds (small, sph ero ida l,
concentr ical ly laminated grains), peloids (ovoid grains produced primari ly
as fecal pellets), and intraclasts ( pieces of se mi -c em en te d carbo nat e sedi
ment that have been eroded and redeposited). In the type-Cincinnatian,
ooids are absent, peloids are uncommon, and intraclasts occur sparingly
Oceanography
lectiv ely sugges t a decre ase in the intensity of up we ll in g in the
O r d o v i c i a n .
A d d i t i o n al e v i d e n c e from the ri ch fossil faunas supports the in ter
tion of co ol wate rs, fol low ed by a return to wa rm wate r in the latest O
cian. D ur in g the Late Or dov ic i an, the western United States and C
straddled the equator . Th ei r carbo nate sed imen ts are typical of m
w a r m water sett ings, so their faun as are interpreted to re fl ec t w a r m
conditions. These areas contain abundant corals and stromatoporoids
a diverse array of bra chi opo ds and trilobites. In part icular, col onia l ru
and tabulate corals (for example, Tetradium), solitary corals (Grew
Streptelasma), several brach iopo ds (Glyptorthis, Plaesiomys, Rhyncho
Hiscobeccus, Lepidocyclus, Holtedahlina, and Leptaena, for exam ple) ,
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bi te s (Ceraurinus), an d diverse ce ph al op od s are characteristic of this
water faun a. T h e s e o r g an i s ms are abse nt from E d e n i a n an d Maysvstrata in the Ci nci nna ti area, but appear in the Ri ch mo ndi an as the
stones beg in to reflect a return to war m water, low-n utrient conditi ons
Type-Cincinnatian rocks differ from typical carbonate platform de
in ano the r signi ficant aspec t: the ab un da nc e of terri geno us mu d, that i
pro du ced b y the wea th er in g of silica-rich min eral s such as feldspar
earliest influx of this mu d closely coi nci des with t he be gi nn in g of nu
i h l t d it t th b f th L i t Li t
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Tidal flat environments today are flat, nearly featureless areas that
for m be tw ee n the low tide li ne and the hig h tide lin e. The se area s are
covered daily by tides, but are subjected to extreme variations in salinity
and temperature on a daily basis.
In U pper Or do vic ia n rocks of the Ci nc in na ti Ar ch , t idal f lat envir on
ments are preserved as laminated to burrowed dolomite and dolomitic
l imes tone co nt ain ing small am ou nt s of c lay (F igu re 15.4A). T h e pres ence
of dolo mite suggest s strong levels of evapo rati on w hi ch wo ul d hav e drawn
Figure 15.3. The four
principal sedimentary
environments of the
type-Cincinnatian. Cin
cinnatian seas generally
deepened northward
from shallow water envi
ronments in central Ken
t k t d t
Figure 15 .4 . A. Out
crop photograph of finely
laminated dolomite de
posited in a tidal flat en
vironment. B. Outcrop
photograph of rubbly
weathering, nodular
limestone and mudstone
deposited in a shallowsubtidal environ
ment. C. Outcrop pho
tograph of interbedded
limestone and mudstone
deposited in a deep sub-
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p p
tidal environment, with
the limestone beds recording deposition dur
ing hurricanes. D. Out
crop photograph of
mudstone with thin beds
of limestone and silt-
stone, all deposited in an
offshore environment.
va si ve b u r r o w i n g of the se d i m e n t by soft-bodie d or gan i sms. A l t h o u g h storms
certainly reworked the sediment and deposited the characteristic well-sorted
layers of shells overlai n by layers of mu d that are prese rved i n s om e pla ces ,
subsequent burr owin g mixed these layers, pro duc ing pods of shell-r ich and
shell-poor material . Preferential cem ent at io n of these ch ur ne d sed ime nts
produce d pockets of well -cem ente d shells material surro unded by non -ce
mented zones rich in clay.
Shallow subtidal l ime ston e in the Ci nc in na ti area is loca lly r ich in
phos phat e , particular ly in the Maysv i l l ian . Mu c h of this pho sph ate oc cur s
as infillin gs of bryozoan zo oe ci a, the por ous skele tons of echino derms, and
the larval shells of pelecypods, gas tro pod s (such as Cyclora), a n d m o n o p l a -
coph oran s. The pres ence of this phos phat e indicates large am ou nt s of de
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cay ing organi c matter within the sedi men t. By dissol v ing piec es of shal low
subtidal limestone in vinegar or dilute hydrochloric acid, one can see the
rich fauna preserved by this phosphatization. Shallow subtidal rocks are
broadly distr ibuted ove r the C i n c i n n a t i A r c h a n d o c c u r f ro m the s o u t h e r n
edg e of the Ord ovi c ia n outcro p belt in southern Ken tuc ky to the north ern
limit of Ord ovi c ia n rocks in central Oh io and Indi ana. T h e Be lle vue , Mt.
A u b u r n , O r e g o n i a , an d W h i t e w a t e r F o r m a t i o n s al l a c c u m u l a t e d within
shallow subtidal environments.
As on m o d e r n shelves , storm depos its ar e the m ost c o n s p i c u o u s
of the d e e p su b t i d a l e n v i r on m e n t i n the ty p e - Ci n c i n n a t i a n , w i th r
e q u al p r op or t i o n s of th i n to me d i u m- b e d d e d she lly l i me ston e , lam
si l ts ton e s , an d mu d ston e (Fi gu r e 15.4C). Burrowing is much less i
here tha n in de ep subtidal en vir onm ent s, result in g in thicker and
lateral ly co nt inu ou s l imest one beds. Bed s of s i l tstone are co m mo n
p le d or d i sp lay i n te r n a l p lan ar or hu mmoc ky lami n at i on ge n e r at
strong storm currents and waves. At t imes, storms occurred with suff r e q u e n c y that the y c ommon ly e r od e d thr ou gh the mu d ston e laye
ping the deposit from the previous storm, such that the shelly layer
on e storm was depos ited di rect ly on the shel ly bed of the previ ous
T h i s p h e n om e n on , kn o w n as am alg am ati on , p r od u c e s th i ck laye r s o
i h b l i l i f h i di id l
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stone with subtle internal erosion surfaces that separate individual
b e d s , t h e r e b y p r o d u c i n g w h a t are k n o w n as m u l t i - e v e n t b e d s . I n
cases, a on e-foo t thick bed of l im est one m ay record ha lf a doz en
events. De e p subtidal rock s are as broa dly distr ibuted over the Cinc
A r c h as s h a l lo w subt idal rocks. The Fa i rvi ew, C o r r y v i l l e , S u n se t , an
e r ty For mat i on s ac c u mu late d i n d e e p su b t i d a l e n v i r on me n ts .
D e e p su b t i da l r oc ks of the typ e -C i n c i n n at i an c on ta i n an ab u
and diverse fauna. Preservation is commonly better than in shallow
tidal rocks with less overall disarticulation breakage and abrasion
quarrymen were the River Quarry Beds (now called the Point PleasantForm atio n) and the Hill Q ua rr y Beds (now call ed the Fair view For mat ion ).
A l t h o u g h the River Qu a rr y Beds n ear C i n c i n n a t i ar e n o w largely u n d e r
the Ohio River , whose level was raised during the construction of dams,
they can still be seen n ear Point Pleas ant, Oh i o , alo ng the crest of the
Cincinnati Arch. Many of the Hill Quarries can still be seen in the bluffs
south of the Universi ty of Ci nc in na ti and f lan king the Mill Cr ee k Valley.
Of fs ho re en vi ro nm en ts on mo de rn coasts l ie below t he wave base of
most storms, but are sometimes affected by the most severe storms and
extend to depths of several tens of meters. In these modern settings, deposi
t ion is dominated by muds, which can accumulate when currents and
waves ar e weak. Rare, e x c e p t i o n al l y strong storms are c a p a b l e of m o v i n g
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, p y g p g
shells and sediments even at these depths and produce thin storm beds,
alt hou gh these mak e up a minori ty of the deposits . Offs hor e env iro nm ent s
are adjacent to and somewhat deeper than deep subtidal sett ings.
In the type-Cincinnatian, offshore rocks contain a greater proportion
of mu ds to ne (c om mo nl y near two-third s), but in oth er regards are quite
simila r to the deep subtida l (F ig ur e 15.4D). The less freq uent oc cu rr en ce
of storm beds in offshore depos its indica tes less frequen t dis tu rba nc e by
storm-generated waves and currents As a result amalgamation is much
de ep wat er sett ings is the pr es en ce of the bl ind trilobites Cryptolithu
Triarthrus in offshore strata. Crinoids are also numerous and are frequ
ar t i c u la te d . T h e most c o m m o n ge n e r a ar e Cincinnaticrinus and F
crinus, w ho se ossic les ma y co mpr ise entire beds of l imes tone. G iv e
tens of ki lo mete rs over wh ich such beds can be traced, the num be r
noid individuals must have been astronomical . As in the deep sub
trace fossils are numerous in beds of siltstone. Chondrites, Diplocra
Trichophycus an d Paleophycus are al l common. The tr i lobite burrow
phycus i s a lso c ommon , an d e xamp le s of Rusophycus made by Isotelu
Cryptolithus have been reported, but ones produced by the calym
Gravicalymene an d F l e x i c a l y m e n e a r e m u c h m o r e c o m m o n .
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LIFE IN THE CINCINNATIAN SEA
16The Ecological Theater and the Evolutionary Play is a boo k of fas cin ati ng es
says about the complex interactions between the environment and the organ
isms inh abi tin g it. It was writt en by the eco lo gis t G. E. Hu tc hi ns on in 1965.
The Ecological Theater
and the Evolutionary Play
G. E. Hutchinson 1965
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Hutchinson's title, an extrapolation of the Shakespearean metaphor, provides
a useful an alo gy by wh ic h to view the Ci nc in na ti an as a Series of acts in the
evolutionary play. In chapters 5-14 of our book we introduced the cast of
characters, the players on the stage, of the Late Ordovician sea that covered
the Cin cin nat i Arch region. Hav ing read these chapters, the reader shou ld be
able to rec og niz e the character s and know so me th in g of their rol es— in par
ticula r their mod es of life and fe ed in g habits.
In scientific terms this is the realm of autec ol og y the rela tion ship of
G. E. Hutchinson 1965
Figure 16. 1. A. Internal
mold of nautiloid, Trep-
toceras duseri (Hall and
Whitfield), with crinoid
Xenocrinus baeri
W h a t cast was on stage fo r a g i v e n act? An ecologis t c an observe an d s
living organisms in the field, but the paleoecologist must deal with
b lages of d ead re mai n s preserved in s e d i m e n ta r y rock . Fac to rs af fect in
preserv ation disc usse d in chapt er 1 are of the utm ost imp or ta nc e her
these as sem bla ges represen tative of act ual life assemb lag es or are they
ass emb lag es repres enti ng mi xtu res of org anis ms that lived at different
or places and accumulated gradually over t ime (t ime-averaged assemb
or sudde nly in so me quick event? How mu ch information is missinthe fossil record be ca us e of preservat ional bias? Fossil assem blag es are
in favor of org ani sms wit h preserv able re mai ns with hard parts like
skeletons, and exoskeletons, and they are biased against organisms l
hard parts. Criteria such as those presented in Table 1 in chapter 1 c
applied to answer these questions. Th ro ug ho ut this book , exam ples
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applied to answer these questions. Th ro ug ho ut this book , exam ples
cin nat ian fossils preser ved in life positio n or in direct associa tion with
isms of oth er speci es provide ev id en ce by wh ic h to dist ingu ish life
b l ag e s f r o m d e a t h as se m b l ag e s . K e e p i n g these iss ues i n m i n d , w
proceed to examine how fossi l assemblages vary through the Cincin
and what this reveals about Cincinnatian paleosynecology.
Ev er sin ce som e of the earl iest studi es of Ci nc in na t ia n fossils and
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the section, toward higher values moving up-section. As noted in chapt
and 15, analyse s of lith olog ic featur es such as the shale-to -lim eston e rati
b e d d i n g thickness demonstrates th at water depth decreased from the ba
the top of the forma tio n. The D C A sho we d that the com pos iti on of
assemblages also reflects this trend and may even provide a more sen
mea su re of dep th c han ges th an the chara cter of the rock reveals.
In fossil ass emb lag es from the lowe r Kop e (deeper water), the
ab un da nt fossils are the slend er cri noid s Ectenocrinus and Cincinnnus, t he s mal l , t h in- s hel l ed brachiopod Sowerhyella, and the trilo
Cryptolithus and Acidaspis (see Fi gu re s 11.5,11.6). Higher in the Kope
l arger brachiopod Dalmanella, br an ch in g bryozoans, a nd the trilo
Flexicalymene an d Isotelus b ec om e the most abun dan t taxa. The brachiopods Zy
Dalmanella assemblage at higher levels. At the lop of the Kope, the l
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g g p p
concavo-convex brachiopods Rafinesquina and Strophomena are the
abu nd ant brachiopods, an d the larger crinoid Glyptocrinus replaces
smal ler crino ids. In an earlier study of the K op e to Fairvie w to Bell
Formations, Diekmeyer (1998) found s imilar transit ions from tax
smaller, more delicate animals in the Kope to larger, more thick-sh
and robust an ima ls ( Pl aty stro ph ia, more massive bryozoans) in the ov
in g Fair view . Sh e inter pre ted this to be a result of a co nt in ua ti on o
h l l i i i t i t d d i th d it i f th K q
hut the influx cu lm in at es withi n t he C5 seq ue nc e wh er e fossils of over fifty
new genera of corals , brachiopods , bry oz oan s, moll usc s, tr i lobites, and
ec hi no der ms appear (Ho llan d 1997; Hollan d and Patzkowsk y 2007; f i g ur e
16.3; see cha pte rs S and 9). many of the new taxa were not pre sen t du ri ng
the Ed e n i an an d M ay sv i l l i an S tage s of the Ci n c i n n at i a n , a l t hou g h some
n e w c om e r s r ep r ese nt sp e c i a t i on w i thi n lon g- r an gi n g Ci n c i n n a t i an taxa
suc h as the brachiopods Platystrophia an d Strophomena (Holland 1997).
Ne w taxa appear ed in al l deposit i onal en vir onm ent s across the spe ctr um
of the Ci nc in na ti an depth gra dient, and the an ima ls occ up y the entire
rang e of fee din g typ es and life habits (Ho ll an d 1997).
In the C4 seq uen ce, s om e e le men ts of the older , depth-rela ted ass em
b lages, su c h as Hebertella an d Platystrophia, are present in the shallow
btid l d R fi i d Z i i th d btid l
The invasion was not
limited to particular fa
cies , trophic groups,
or life-habit groups;
rather the Riehmondian
Invasion was a major
ecological revolution
affecting all aspects of
the Cincinnatian seas.
Steven M. Hol land
1997, 320
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subtidal zone, and Rafinesquina and Zygospira in the de ep er subtid al zo ne
(figure 16.3). In the C5 sequence, a Dalmanella b r ac h i op od asse m b lage i sagain present in the offshore env iro nm ent s, hut sp ec im en s of Zygospira
also are present. Sp ec im en s of Rafinesquina, Platystrophia, an d b r an c hi n g
b r y o zo an s o c c u p y the d e e p e r subt idal z o n e , but the Hebertella-Platystro-
phia assemblage is gone from the shallow subtidal zone, where an assem
blage of c o lon i a l cora ls ap pe ars fo r the first t i m e. Th e d e p t h gr ad ien t is
re-established in the C5 sequence, but it is much more crowded with taxa
plot and o ut co me of the play. In terac tion s bet we en o rga nis ms are of m
im po rta nce in eco lo gy and lead to the con cep t of an ecos yste m.
For an ecologis t , an ecosys tem enco mpas ses all the che mic al , phy
and bio log ical aspects of the envi ro nm ent , in clu din g the sources of en
and nutrients entering the environment and the way living organisms
this ener gy to survive and repro duce . The Sun is the prim ary energy s
for the vast majority of eco sys tem s on Ea rt h, the only kn ow n except ion
ing the recently discovered deep sea vents, where hydrothermal fluidsin nutrients sustain microbial life that is, in turn, the basis for unique
systems. In shallow seas like that of the Ci nc in na ti an , we can ass ume
planktonic as well as benthic algae harnessed solar energy as the pri
pro duc ers . Individuals of a ll of the anima l groups consti tuted con su m
feeding either directly on the primary producers or on other consumer
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order to unders tand the natu re of Ci nc in na ti an eco syst ems, we must u
stand how the consumers were interrelated in what ecologists call a
ch ai n or, mo re realistically, a fo od we b. How did Or do vic ia n mar ine ec
tems c om pa re to those of the present-day shal low sea? Di d the nature o
interrelationshi ps am o ng org anis ms and th e form of the food web play a
in det erm ini ng the diversi ty and abu nda nce of organism s in the Ci nci
tian sea? In our analogy, did the interplay among characters actually d
i th t t d i ti l t?
species derives some benefit, whereas the "host" is unaffected by the presence of the symbiont. In so me cases a living an im al of on e spec ies mi gh t
be associated with n o n - l i v i n g r e m ai n s of a n o t h e r sp e c i e s , as in the c ase of
hermi t crabs oc cu py in g shells of dead snai ls (Dav is, Ma pe s, and Klo fak
1999; Davis, Fraaye, and Holland 2001).
W e c o m p i l e d avai lable in for mat io n o n associat i on s o f f os si ls o f C i n c i n
natia n speci es into two su mm ar y tables. Table 2 sho ws pot enti al predator-
prey asso ciati ons deriv ed from direc t fossil evi de nc e, and Table 3 sho ws all
other associations reported am o ng individual s of Ci nc in na ti an species or
other groups.
Pr e d ator -Pr e y In te r ac t i on s
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The list of potent ial predator-prey asso ciat ions in the Ci nc in na ti an is quit eshort (Tab le 2), and the natur e of the evi de nc e is varia ble. Th ro ug ho ut the
fossil record there are rare but nota ble eases of fossil ized st om ac h cont ent s
that are the strongest evi de nc e for the diet of an extin ct an im al . The only
possible instance of this in the Ci nc in na ti an is the oc cu rr en ce of ostra codes
preserve d wit hin the corall a of the rug ose coral s Grewingkia an d Streptel-
asma (E lias 1984). The ost raco des are mos tly art icul ated and locat ed near or
Cincinnatian crinoids also show evidence of damage and regener
of the cal yx , arms , and co l u m n that is very likely the result of pred
(Ausich and Baumiller 1993; Donovan and Schmidt 2001; Baumiller
Gahm 2004). However there is no ev id en ce as to the speci fic predato r re
sible. We speculate that nautiloids might be the most likely culprits, in
of their ab un da nc e and potent ial behav ior.
O t h e r I n t e r s p e c i f i c I n t e r a c t i o n s
Table 3 shows that virtually all the major invertebrate groups found i
Ci nc in na ti an ha ve recorded associations of individu als of taxa bel o
to a wi de variety of gro ups . T h e type or nat ure of the associ ation s r
from the use of eith er a livi ng host or dead re ma ins as a substr atu
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encrustation, boring, or habitation (Figure 16.1) , to possible commens
or parasitism. To the extent that a host was, by definition, living at the
of the associatio n, i t appears that interaction s betw een individuals o
cin nat ian speci es were very c o m m o n. Deta i le d discussion of the nat
the associations l isted can be found in the chapters concerning the
no mi c gro up of eac h host.
We present Ta bl e 3 w i t h a s ign if icant cave at . In cases in w h i c h a
i i l d i f i tt h d t i di id
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THE pr imary producti v i ty of the world 's oc ea ns dur in g the Ph an ero zo ic
Eon. As mentioned ear l ier , we assume that pr imary production in the
Ci nc in na ti an sea was based on m icro phy top lan kto n as well as so me ben
thic macro alg ae. It is a paradox that despite the high ab un da nc e of ben thi c
suspension fe edin g invertebrates in the Ci nc in na ti an fauna the onl y pre
served microphytoplanktonic organisms are the acr i tarchs. Were acr i tarchs
the only suspe nded food so urce for a bot tom faun a dom ina ted by ab und an t
and diverse suspension feeders? We can speculate that perhaps other mi-
crop lan kto nic algae existed —o n es that lacked pres ervabl e org ani c ce l l
wal ls or m i n e r a l i z e d te sts. O r , p er haps c lay partic les s u s p e n d e d in the water
column served as substrata for bacteria that formed a "marine snow" that
nourished benthic suspension feeders. But, speculation aside, the fossi l
record of ma rin e plan kto n definit ely in dicat es that the diversity of taxa of
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p y y
p lan kton i c or gan i sms i n the Ci n c i n n at i r e gi on d u r i n g the Ci n c i n n at i an wa s about fif ty sp ec i e s ( C o l b a t h 19 79 ), r e a c h e d a P a l e o z o i c m a x i m u m of
three- to four hu ndr ed spe cies of aerit arch s, the n suffered a de cl in e du ri ng
the late Pal eoz oic and early Meso zo ic . It then beg an to incr eas e du ri ng the
Jurassic and Cretaceous, with the appearance of present-day groups such
as d i n of lage l la te s , c a lc ar e ou s n an n op lan kton , an d d i a toms (T ap p an an d
Lo ebl ich 1973). If the diversi ty of p la nkt oni c orga nis ms is correlate d in
Figure 16.3. Time-envi
ronment diagram for theCincinnatian Series. The
vertical axis shows the
timescale and six major
shallowing-upward se
quences of the Cincinna
tian (see chapters 4 and
15). G = Gamachian Stage(not preserved in Cincin
nati region). The horizon
tal axis shows the major
environments of the Cin
cinnatian (see chapter 15).
Offshore environments
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are located towards the
present north in Ohio and
westward in Indiana, and
shoal and lagoon environ
ments are located toward
the present south in Ken
tucky (see Plate 12). FWB
= fairweather wave base,
b
hard pressed to f ind mai n go od anal og ues . Of cou rse, exti nct ion has swept
away v irtually al l the taxa of the organi sms that inhabit ed the Ci nc in na ti an
sea, including entire major groups such as graptoli tes, conular i ids, tr i lo
bites, e u ry pt er i ds , an d edrioasteroids . H o w e v e r , tod ay w e c a n f i n d rather
restricted regions where the sea floor is covered with bryozoans, such as the
shelf of f Sou th Australia ( Bradst ock and Co rd o n 1983; H ag em an et al .
2000). New Zealand, isolated in the southwestern Paci f ic , harbors the
greatest diversi ty of l iv ing b rach iopo ds; e l sew her e these ani ma ls are but
mi n o r c omp o n e n ts of sha l low w ate r c om mu n i t i e s . C lo se r to the are a of the
Ci nc in na ti an , the San Juan Islands in the Paci f ic Nor th wes t suppo rt a di
vers e and ab u n d a n t fauna that i n c l u d e s m a n y P a l e o z o i c e l e m e n t s s u c h as
e c hi n od e r ms, b r yozoan s , b r ac hi op od s , so l i tar y c or a ls , an d sp on ge s . How
ever, these relicts are a mere fracti on of a mo re diverse fauna rich in mo l
gospira; trilobites; ca =
calymenid, Ct = Cryptoli-
thus, Is = Isotelus; ostra
codes = os; gastropods =
ga; p e le c yp od s; Am =
Ambonychia, by = inde
terminate pelecypod, Ca
= Carotidens, mo = modi-
omorphid pelecypod; cri
noids. Ci = Cincinnaticri-
nus, Ec = Ectenocrinus,
Gl = Glyptocrinus, Xe =
Xenocrinus; graptolites =
gra. Italicized fossils are
h d
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ever, these relicts are a mere fracti on of a mo re diverse fauna rich in mo l
luscs, crus tace ans, and f is h—all gro ups that proli ferated in the post- Paleo zoic . Present-day tropical reefs far surpass those of the Or do vi c i an in
diversity and abundance, and yet there are certain reef-related habitats that
mirror , to some extent, the Paleozoic . In Australia 's Great Barr ier Reef ,
deeper , soft sediment bottoms located below the coral-dominated shallow
reefs have a Paleozoic aspect, r ich in algae, sponges, soli tary corals , bryo
zoa ns, and cr inoids (Me ssi ng et a l . 2006). Indiv idual s of the cha mb er ed
those invading region
during the Richmondian
invasion. Information de
rived from Holland and
Patzkowsky (2007); see
this publication for distri
bution of additional
fossils.
Table 3. Associations
among Individuals of Cincinnatian Taxa
A R R A N G E D B Y " H O S T "
Host Associate Type of
Associat ion
References /
[Annotations
PROTISTA
foraminifers bryozoan bi oi mm ur at io n;
epizoism
Wi ls on , Palm
and Taylor (1
PORIFERA
stromatoporoids pelecypods boring Pojeta and P
(1976);
Wi ls on an d
Palmer (1988
CNIDARIA
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CNIDARIA
hydrozoan bry ozoan encrustation (post
mortem);
dwelling inside
empty shell
Wi ls on , Palm
and Taylor (1
corals alg ae, fungi boring Elias and Lee
(1993)
A R R A N G E D BY " HOS T "
Host Assoc i a te Type of
A s s o c i a t i o n
Re fe r e n c e s /
[ A n n o t a t i o n s ]
bryozoans encrustation; ?
epizoism
Anstey and Wi l
son (1996)
[Corynotrypa on
interior of bra
chiopod shell]
bryozoans en cr us ta tio n; ?
post-mortem
Anstey and Wi l
son (1996)
[Cuffeyella on
interior of bra
chiopod shell]
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bryozo ans en cr ust at io n Ulrich (1879)
bry ozoans en cr us tatio n Ulrich (1883)
brachiopods ep iz ois m Alexander an d
Scharpf (1990)
Cornulites ? commensalism Morris and Rollins
(1971)
A R R A N G E D B Y " H O S T
Host A s s o c i a t e T yp e o f
A s s o c i a t i o n
References /
[An n otat i on s]
Cornulites epizoism; ?
commensalism
Morris and Rol
(1971)
cornulitids encru stat ion and
intergrowth
Baird, Brett, an
Frey(1989)
pele cypo ds boring Wilson and
Palmer (1988)
Sanctum bo ri ng Erickson an d
Bouchard (200
Sphenothallus epizoism Bode nbend er
al (1989)
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al. (1989)
trilobites aegism Shrake(1989)
"worms" —
Trypanites
bo ri ng s Palmer an d Wi
son (1988)
CORNUUTIDS
cornulitids bryoz oans encrustation and
intergrowth
Baird, Brett, an
Frey (1989)
A R R A N G E D B Y " H O S T "
Host Associate Type of
Assoc iat ion
References/
[Annotat ions]
Mollusca
"nautiloids" —
actinoceroids
endoceroids
nautiloids
bry ozoans
bry ozoans
epizoism; ?
commensalism
encrustation
presumablyepizoism;
? commensalism
Frey (1988, 1989);
Baird et al. (1989)
Davis and Mapes
(1996)
bryozoans encrustation Ulrich (1879a)
bryozoans en cr us tat io n U. P. James
(1884b)
b
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bryozoans en cr ustat ion Ulrich (1883);
Bassler (1953)
bryozoans en cr us ta tio n (p os t
mortem);
dwelling inside
empty shell
Wi lson, Palmer,
and Taylor (1994)
Cornulites ? epizoism
? commensalism
Richards (1974)
A R R A N G E D B Y " H O S T "
Host Ass oci ate Type of
Associat ion
References/
[Annotations
Echinodermata —
crinoids
byroni ids parasitism Wa rn (1974) ;
We lc h (1 976) ;
Malinky et al.
(2004)
Echinodermata —
crinoids
Cornulites commensalism Morris and Ro
(1971);
Morris and Fe
(1993, 2003);
Richards (1974
Echinodermata —
crinoids
gas tro pods commens alism Bowshe r (1955
Morris and Fe
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crinoids Morris and Fe
(1993)
Echinodermata —
crinoids
"WORMS"
(see also:
cornulitids)
ga st ro pod s ? parasitism ? Baumiller and
Gahn (2002)
SUSPENSION HERBIVORE CARNIVORE
PRIMARY
PRODUCERS
chitinozoans (?) cephalopods acritarchs
conodonts eurypterids
graptolites
trilobites (pt)
SUSPENSION
trilobites (pt)
MOBILE
ATT ACH ED
LOW
stromatoporoids (pt)
tab., rugose corals (pt)
br yo zo an s
DEPOSIT HERBIVORE CARNIVORE
monoplacophorans mono plac opho rans trilobites (pt)
gastropods gastropods asteroids
trilobites ostracods
ostracods
ophiuroids, asteroids (pt)
A U N A
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craniate brachs
rhynch. brachs
bivalves
cornulitids
edrioasteroids
cyclocystoids
ATTACHED
ERECT
sponges
conulariids
stromatoporoids (pt)
tabulate corals (pt)
SUSPENSION DEPOSIT CARNIVORE
SHALLOW
PASSIVE
rostroconchs
SHALLOW
Ungulate brachs
bivalves
bivalves
polychaetespolychaetes
E P I
F A
PRIMARYSUSPENSION HERBIVORE CARNIVORE PRODUCERS
gastropods bo ny fish cephalopods dinoflagellates
malacostracans mammals chondrichthyans coccolithophores
mammals bony fish diatoms
reptiles
mammals
F A U N A
SUSPENSION DEPOSIT
MOBILE
biv alvescrinoids
ophiuroids (pt)
asteroids (pt)
holothuroids
gastropodsostracods
malacostracans
ophiuroids
asteroids (pt)
echinoids
holothuroids
ATTACHE D
sponges
corals
br yo zo an s
HERBIVORE CARNIVORE
chitons gastropodsgastropods cephalopods
ostracods malacostracans
malacostracans asteroids
echinoids
P E L A G I C
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E P
I F
LOW
br yo zo an s
brachiopodspolychaetes
bivalve s
ba rn ac les
ATT ACHED
ERECT
sponges
corals
bryo zo an s
crinoids
RECLININGcorals (pt)
bivalves
SUSPENSION DEPOSIT CARNIV
SHALLOW
PASSIVE
gastropods
biva lves
echinoids
bi val ves bi va lv es
SHALLOW
AC TIVE
lingulate brachs
biva lves
polychaetes
bivalve s
polychaetes
echinoids
gastropod
malacostr
polychaet
EPILOGUE: DIVING IN THECINCINNATIAN SEA
many paleon tologi sts, ourselv es in clu ded , b ec a me fascinat ed wit h fossils
and emba rke d on scientific careers long befor e we ever en co un te re d liv ing
mar ine an ima ls. For ma ny of us, the greatest thril l has be en our firs t en
counte rs with l ivi ng representatives of the ani ma l grou ps we kne w firs t only
as grey, lifeless for ms en ca se d in rock. Bot h of us ha ve be en pr ivi leg ed to
ex am in e f irstha nd livin g relatives of an im al s of our favorite grou ps of fos
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ex am in e f irstha nd livin g relatives of an im al s of our favorite grou ps of fos
s i ls—crinoids for Meyer and nauti loid cephalopods for Davis . Our experi
ences have fueled a curiosity that affects practically anyone who contem
plates the fossil r ich ness of the Ci nc in na ti an or other co mpa ra bl e
fossil iferous strata. Many times, in the field, we stand on a Cincinnatian
outcrop where fossils are abundant in almost every rock, and we wonder:
what did the C i n c i n n a t i a n sea act ual l y look li ke? How did these creatures
We d on masks and snork els for a quic k reconnoiter, an d sl ip ove
side. We have anchored over very shallow water, and the bottom appears
a meter or so beneath us. As we take a closer look at the bottom, we n
that it is irregula r, with low m ou nd s separ ated by pat ches that are more
T h e mou nd s are actually c lum ps of large, r ibbed brachiopo ds, Platystro
ponderosa. Livi ng ani mal s with articulated shells are intermin gled with
rated valves, some broken and worn smooth. It is the environment
millions of wars in the future, will be preserved as the Mt. Auburn Me
of the Gran t Lake Lim est one . We easily scoo p up a sample of spe cim e
Platystrophia because they have no pedic le attachments.
Cl ea rl y there is m u c h of interes t to see here , but we retur n to the
b e c a u s e snorkel d i v i n g is not ad e q u a t e for p r ol on g e d explorat ion. We
not been able to hold our breath during our dives as long as we norm
d o — w e had to c o m e up quickly , gasp ing for air. A che ek of our air qu
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d o w e had to c o m e up quickly , gasp ing for air. A che ek of our air qu
monitor reveals the reason: the Late Ordovic ian atmosphere has on
fract ion of the ox yg en con ten t of present- day air, perh aps as little as 1
cent. Fort unatel y we have brou ght alo ng some sophisticated div ing
that will let us fi ll our diving cylinders with compressed air in which
have boosted the oxygen content to its present-day level, 21 percent
cause the rest of our com pres sed gas mixt ure is predo min ant ly nitro
the ambulacra] grooves recall the reconstructions made by Bruce Bell .
W i t h a rock pick we easi ly break off a p i e c e of the h a r d g r o u n d w i th e d r i o
asteroids att ach ed, and we ba g it for study in the lab. The ha rd gr ou nd gives
way to an area c ov e r e d w i th th in-shel led Rafinesquina b a c h i o p o d s , f o r m i n g
shell pavements l ike those we found in rock units with names l ike Cor
ryvi l le , Bellevue, and fairv iew in the distant future from which we came.
T h e c on c av o -c on v e x b r ac hi op od s r es t w i th the c on v e x v a lv e e i the r d o w n
or up, and main are encrusted with small bryozoans or edrioasteroids.
Brachiopods of taxa like Zygospira a n d p e l e c y p od s of ge n e r a l i ke Cari-
todens are attach ed. O th er c lam s, of taxa l ike Modiolopsis, poke up through
the se d i m e n t b e t w e e n she lls . S mal l Flexicalymene an d Acidaspis tri lobites
glide over the surface, and here and there a crinoid has the stem coiled
around a bryozoan. This is a diverse habitat.
B t th t b th f l i i i l t d
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But the water mass above the sea f loor is surprisingly empty compared
to the sc en e in presen t-da y shall ow seas. Today, fish are ev er yw he re in the
sea, fi lling a wide variety of ec ol og ic al roles. Wh e r e are the fish in this
Or d ov i c i an sea? On l y smal l n au t i lo i d c e p ha lo p o d s ar e j e t t i n g ab ou t ,
agai nst a bac kdr op of pul sat ing jellyfish. At clo se ran ge , we ca n pic k ou t
very smal l strings s u s p e n d e d i n t h e water w it h c l u m p s o f m i n u t e te n tacle s
arra nge d in vertical series. These are graptol ites. Clo se r to the bo tt om som e
A gu l ly leads us b ac k into d e e p e r water. H e r e we find va st areas co
w i t h b r a c h i o p o d s of taxa l ike Rafinesquina and Strophomena or b r an b r y o z o a n s . T h e r e are i n t e r v e n i n g p a t c h e s of m u d a b o u t e q u al i n ar
the shelly patch es; this mus t be the env ir on me nt of the Fairview,
continuous, even beds that wi l l produce about 50 percent l imestone
50 percent shale .
In the dis tan ce we spot a ridge of bry ozo an s sta ndi ng alm ost a
abo ve the sur rou ndi ngs , and we hea d toward it. Co u l d we he app roa ch
shar p drop-off lea din g to dee per water? Ap pr oa ch in g closer, we can
richnes s of life aro und this ridge, and we sense a gent le curren t f lowing
the bottom, parallel to the gradual slope, which intensifies as we reach
ridge. T h e depth ga ug e reads 15 meters (50 feet), and we have des ce
b e l ow the dep th w h e r e the small surface waves st ir re d the b o tt om .
cur ren t flow takes over and fol lows the con tou rs of the slope. Th e rid
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actuall y a mo un d bui lt entire ly of the br anc hin g bryozoa n colonie s of g
Parvohallopora; it proje cts ou tw ar d fro m the slo pe, The cur ren t is div
and gains velocity as it f lows over the ridge. Crinoids forming a dense c
are of ge nu s Glyptocrinus; their stalks are coiled around the bryo
b r a n c h e s , a n d the c r in oid s stand ab o v e t h e m lik e a fo re st canopy.
crowns are splayed out in feathery fi ltration fans that are all aligned pe
of northern C an ad a! Never thel ess, we have ma na ge d to gath er up several
smal ler trilobites, and may be their D N A fina lly will resolve the que sti on of
how trilobites are related to other arthropods.
We venture out beyond the bryozoan ridge onto a seemingly lev el plain
with patches of b rac hiop od p av e m e n t and b r yozo an s . The b r ac h i o po d s are
noticeably smaller forms like Dalmanella an d Sowerhyella. Bryozoans are
delicate, twig-like colonies with fewer sheet-like or massive forms. Flexicaly
mene trilobites are here , hut are sm all er th an tho se we foun d in sha llo wer
wa ter; so m e n e w tr il ob it es of gener a like Cryptolithus and Triarthrus cruise
on the mu ddy patches, leaving gro oved trails. E xt en din g upward from som e
br yozoan thic kets ar e very s lender yet l o n g - s t e m m e d cr inoids of the tax a
Ectenocrinus and Cincinnaticrinus. Al th ou gh they h ave fewer arm s than the
other crinoids we have seen today, they splay them into conical filtration fans
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in alignment with the gentle current (not open into the flow but with theconcave side bearing the food grooves downcurrent). Some sediment patches
are sands entirely com po se d of frag ment s of disarticu lated cr inoids , with o dd
bu ndles of lon g , sti ll-articu lated stei ns . The sands have broad, s inu ou s rip pl e
marks like those along a beach. How could such ripples form at our present
dept h of 30 meters (100 feet)? We sense no wave m ot io n and on ly slight cur
rent; however, as experienced divers, we know that severe storm conditions
oxy gen -po or air. We haul o urselves over the gun wa le , peel of f our we
and just lie in the boat, catching our breath, our minds racing wi
sights we have seen. What a dive!
Sud denl y, we sense the warm th of the afternoo n sun, amid the
ness of a crisp au t u m n day. We are sta ring at th e fossil-co vered sur
a bed of Or do vi c i an l im est one l i ttered with rusty aut um n leaves. W
si t t in g on the b an ks of S ton e l i c k Cr e e k , i n Cl e r mo n t Co u n t y , Ohi o ,
lunch on a f ie ld tr ip with our students—and maybe you!
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APPENDIX 1. RESOURCES: WHERETO GO FOR MORE INFORMATION
There are many textbooks in paleontology, but we restrict the following list
to some of the most rec ent as wel l as one old er, clas sic wo rk.
Fossil Invertebrates ( Bo ar dm an et al. 1987)
Principles of Paleontology, 3rd ed. (Fo ote and Mi ll er 2007)
Invertebrate Fo ss il s ( Moore et al. 1952)
Paleontology
textbooks
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Invertebrate Palaeontology and Evolution, 4th ed. (Clarkson 1998)
Ohio Fossils (La Rocque and Marple 1955)
Fossils of Ohio ( Fel dman n and Hac kat hor n 1996)
Exploring the Geology of the Cincinnati/Northern Kentucky Region, 2nd
ed. (Potter 2007)
Publications of
Geological Surveys
relation and Resources," http://www.es.mq.edu.au/MUCEP/igcp
Ma cq ua ri e Universi ty, Sydney , Austral ia (accessed Febr ua
2008).
K e n t u c ky G e o log i c a l S u r v e y , http://wwvv.uky.edu/KGS/, University o
tucky (accessed February 18, 2008).
K e n tu c ky Pa le on tology S oc i e ty , ht tp :/ / w w w .u ky.e d u / Othe r or gs/ K PS
cessed February 18, 2008).
Oh i o G e o log i c a l S ur v e y , http://www.ohiodnr.com/geosurvey Divis
G e o lo gi c a l S u r v e y , O hi o D e p ar tm e n t of N atu r a l Re sou r c e
cessed February 18, 2008).
U n i ve r s it y o f C i n c i n n a t i D e p a r t m e n t o f G e o l o g y , http://www.u
geol ogy / (acce ssed Fe bru ary 18, 2008).
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Th es e are gu ide boo ks to f ie ld trips pert ain ing to the Ord ovi c ia n g
of the Oh io , India na, and K ent uck y regions. Most conta in detai led
logs and directions to geological localities, as well as detailed descri
of exp ose d stratigraphic sections. Loca li t ie s l isted in older gui debo ok
no longer be accessible.
Ca st er 1961b; Hatt in e t al. 1961; Pop e an d Mar ti n 1977; Hay et al
Field guides
S a w y e r P o i nt G e o l o g i c a l T i m e l i n e , C i n c i n n a t i , O h i o http :/ / w w w
.c incinnati-oh.gov/crc/pages/-5708-/ (accessed February 18, 2008)
T hi s t i me l i n e b e gi n s w i th the L ate Or d ov i c i an an d c on t i n u e s thr ou gh
the fou n d i n g of Ci n c i n n a t i w i th e ac h p av e m e n t b loc k r e p r e se n t i n g on e
mil l ion years. Imp ort ant geo log ica l events are eng rav ed on block s at ap
propriate intervals.
T r amme l Foss i l Par k , S har on v i l le , Ohi o http :/ / w w w .shar on v i l le .or g/
fossilpark.aspx (accessed February 18, 2008)This park is dedic ated to edu cat ion about Or do vi c i an geology and pale
ontology (see Figure 1.8).
Paleo ntolo gical Society. The Pale onto log ical Soci ety is the largest pal eon
tological orga niza tion in the United States. It publ ishes both the J ournal
Scientific societies
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tological orga niza tion in the United States. It publ ishes both the J ournal
of Paleontology an d Paleobiology. A ser ies of edu cat ion al bro chu res abo ut
fossils can be downloaded from their website http://paleosoc.org/ (ac
cessed February 18, 2008).
Paleo ntolo gical Re searc h Institution, Ithaca, Ne w York. PRI publ ishe s bot h
the Bulletins of American Paleontology an d Palaeontographica Americana,
as well as a popular magazine, American Paleontologist. Their website is
and institutions
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APPENDIX 2. INDIVIDUALS AND
INSTITUTIONS ASSOCIATED WITH
THE TYPE-CINCINNATIAN
T h e fol lo win g is a l ist of the na me s of indiv idual s and ins ti tutions associ
ated with the Cincinnati region, and, especial ly, i ts geology and paleontol
ogy. So me of the individual s l isted were me mb er s of the Cin ci nn at i S cho ol ;
most were not.
There are some potential problems with this list. In some instances,
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there are two people with similar, but different names, but who may not be
different people. For example, different sources refer to a J. H. Hall and a
John W. Hal l associated with the Ci nc in na ti S oc iet y of Nat ura l History, and
there is I. Harris, I. H. Ha rris, and I. M. Harri s, all of Wa yn es vil le , O hi o .
G e or ge Val lan d i n gham an d G e or ge Val lan d i gham ar e a lmost c e r ta i n ly the
same person, and the latter probably is the correct spelling, but maybe not.
Ci nc in na ti an and other tr i lobites. An th on y was on e of the hosts
Charles Lyell , probably the foremost geologist on Earth, v isi ted the Cnati area in the 1840s. Ant ho ny was autho r, alo ng with U. P. Jame
paper on echinoderms in the local rocks, and he descr ibed an un
sp ec im en of a fossi l ce ph al op od from the ty pe- Ci nci nna ti an. In 18
resigned from t he academ y, and, in 1863, he be ca m e curator of con ch
at Harvard Universi ty, under Louis Agassiz . According to Clench (193
A n t h o n y ' s ". . . c h i e f interest w as in freshwater m ol l u sk s , an d he bu i
series of A m er i ca n freshwat er form s that for its tim e was superior
collection in this country" (Anthony 1838,1839a, 1839b, 1847,1848; An
and James 1846; Brandt and Davis 2007; Coan, Kabat, and Petit
Hendrickson 1947; Johnson 2002; Lyell 1845).
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Dr. Austin, a physic ian from Wilmington, Ohio, was l isted as a loc
lector by Nickles (1936). His collection went to the United States Na
Museum, which published a paper by him on fossi l zones in the Ric
dian rocks of the ty pe- Ci nc in na ti an area (Austin 1927; Becker 1938; N
1936; Shideler [1952] 2002).
Austin, G e o r g e M.
25), and, in conjunc tion with Paul Mo hr , anot her Ci nci nn at i col lector , un
dertook long-term and extensive excavations for crinoids from Carboniferous
rocks at Craw ford svil le, In diana ( Van Sa nt and La ne 1964).
D r . Br i d ge w as b or n i n N or w ood , Ohi o , ad j ac e n t to Ci n c i n n at i , an d he
receive d his bache lor' s de gr ee from the Univers ity of Ci nc in na ti in 1913.
His higher education was e lsewhere, and professionally he was associated with the Uni te d St ates (Geologica l Survey an d the Uni ted States Nati onal
M u se u m i n W ash i n g to n , D . C. He w as a lon g-t i me assoc i a te of Ulr i c h
(Becker 1938; Croneis 1963).
Dr. Richard M ah an Byrnes was a foun din g me mb er <>l the Ci nc in na ti Soc i
Bridge, Josiah
(1890-1953)
Byrnes, Richard M.
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ety of Nat ura l History, and he served as its cura tor of mi ne ra lo gy fr om 1871
until 1880, when he was thrice elected president and, thus, served three years
in that posit ion. He was a me mb er of the society 's C o mm it t ee on Geol og ica l
No me nc la tu re (S. A. M ill er et al. 1879) and was a co-au tho r of the eul og y of
C. B. Dyer. He was listed as a local fossil collector by Nickles (1936), and he
had a large co ll ect io n of fossils, but was inte rested in mi ne ra ls , in terrestrial
(1835-1892)
David Christy was a Cincinnati-area geologist , anti-slavery writer , pr
and newspaperman. His geological letters were published in 1848,
nally in a newspaper, the Cincinnati Gazette, but th en separately. In ar
1858 his collection was sold to Miami University, Oxford, Ohio, for $
(Becker 1938) or, according to Shideler ([1952] 2002, 2), for $5000, "
mos t un hea rd of su m for such a purpo se, in tho se days." T h e same c
tion must have been a financial benefit to at least one other party,
again according to Shideler ([1952] 2002,4): "I have mentioned the Ch
collection, acquired by Miami Universi ty. Some years ago when the
ver sity o f C h i c a g o was c l e a n i n g h o u s e w e w e r e g i ve n s o m e o f thei
w a n t e d m ate ri a l . I n c l u d e d w e r e fo ss il s di st i nct ly m a r k e d ' C h r i s t y C o
t i on . ' T h e Ch r i s ty c o l le c t i on had d i sap p e ar e d f r om M i a mi w he n
insti tution was c losed between 1873 and 1885. Chicago had bough
co ll ect io n from James Hal l wh o ope rat ed out of Al ban y, N.Y. Just
Christy, David
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swiped the collection and sold i t to Hall hasn' t been determined.
material from the Christy collection has come back in similar w
(Becker 1938; Christy 1848; Merrill [1924] 1964; Shideler [1952] 2002).
(S e e T h e Ci n c i n n at i S o c i e ty of N atu r a l Hi s tor y)The Cincinnati
to its ow n bui ldi ng on Gi lb ert Av en ue , in the southw est cor ner of Ed en
Park (Ano n. 1978). At the sam e time , the na me of the orga niz ati on wasch an ge d to the Ci nc in na ti M u s e u m of Natu ral History and, with the ad
d i t i on of a p lan e ta r i u m, the Ci n c i n n a t i M u se u m of N atu r a l Hi s tor y an d
Plan e ta r i u m. W i t h the ab a n d on me n t of the G i l b e r t Av e n u e fac i l i ti e s an d
ab sor p t i on i n to the Ci n c i n n at i M u se u m Ce n t e r a t Un i o n T e r mi n a l i n the
1990s, what was or iginall y the Cin ci nn at i So ciet y of Natu ral History ceas ed
to exist as a separate entity.
Listed as a local fossil collector by Nickles (1936).
Ele cted a me mb er of the Cin ci nn at i Soc iet y of Nat ural History in 1878 and
Cook, W. E.
Cooper, Edward M.
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served as curator of pal eont olo gy of the soci ety (A non . 1879,1885a).
Elizabeth A. Dalve, known to her friends as Bettina, was an illustrator and
part-t ime mu se um assistant in the Dep ar tm en t of Ge ol og y at the Universi ty
of Ci nc in na ti . S he prepar ed a list of the stratigra phic occ ur re nc es of the fos
Dalve, Elizabeth A.
me nt of Ge ol o gy at the Univ ersi ty of Ci nc in na ti . Th e y are st il l goi ng
(Brand t and Da vis 2007;
D a l v e 1951).
(See chapter 2)
(See chapter 2)
O n e of the "ma na ge rs " of the Weste rn M u s eu m at the t im e of i ts fou
a b o u t 1820 ( [S i l l i man ] 1819).
Dyche, D. T. D.
Dyer, Charles Brian
(1806-1883)
Embree, Jesse
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(See chapter 2)
Professor J. C. Kales, of Ce nt re Co ll eg e. D anv ill e, Ke ntuc ky, was the p
Faber, Charles
(died 1930)
Fales, J. C.
Towa rd the end of i ts exist ence, the Wes te rn Mu s e u m in Ci nc in na ti was
best k n o w n as a c h a m b e r of horrors; in that b u s i n ess it had a ri va l in theform of Fra nks ' Mu s e u m (Tu cke r 1965, 32).
Lawyer; listed as a local collector by Shideler ([1952] 2002).
Listed as a local fossil collector by Nickles (1936).
G e or ge G r ah am w as on e of the or i g i n a l me m b e r s of the W e ste r n Ac a d e my
of Nat ura l Sc ie nc es in 1835, and he was one of the seven s urv ivi ng me mb er s
of the ac ad em y wh en i ts assets were don ate d to the Ci nc in na ti Soc iet y of
Franks' Museum
Fulton, Robert
Gault, Wm.
Graham, George
(1798-1881)
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Nat ural History in 1871. Hi s co ll ec ti on of fossils, shell s, and pla nts, unfo r
tunately, was destroyed in a fire (Anon. 1878; Hendrickson 1947; James,
Howe, and Norton 1881).
A u t h o r of the species Nereidavus varians Grinnell, 1877, which was based on Grinnell,
info rma tio n abo ut James H all of N e w York is vol um in ou s and fa
(Becker 1938; Caster 1951, 1982, 1985; Croneis 1963; Merrill [1924]
Sherborn 1940; Shideler [1952] 2002).
J. W. H all , Jr., is listed as curat or of pal eon tol og y of the C in ci nn at i So c
Nat ura l Histo ry from 1874 to 1877, and as corre spo nd ing secretary of th
ety in 1877 and 1878 (An on . 1878). Lat er tha t year, he don at ed so me fo
the soci ety (An on. 1879). He also served the society as curator of min e
(An on. 1885b). He co-a uth ored a pap er wit h a num be r of me mb er s
Ci nc in na ti S ch oo l and other loc al fossil coll ecto rs (S. A. Mil ler et al.
J. W. Hal l, of C ov in g to n, is listed as a loc al fossil collec tor by Nic kle s (
Of M di li t d l l f i l ll t b Ni kl (1936)
Hall, John W., Jr.
H d W E
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Of Ma di s on , so listed as a loca l fossil co ll ect or by Nic kl es (1936).
(See chapter 2)
Hammond, W. E.
Harper, George
W. (1832-1918)
"Andrew Herrl inger , later an attorney in Cincinnati" is l isted as a local
fossil collector by Nickles (1936).
It was Reverend He rze r wh o intro duc ed E. O. Ulr ic h, then seven years old,
to the wo nde rs of fossil col le ct in g (C ro ne is 1963; Ul ri ch 1891).
Dr. Hi l l was associated with the Cin ci nn at i Soci ety of Nat ura l History in
the 1870s, in cl udi ng vario us terms as curato r of arc hae olo gy, cura tor of
conchology, and librarian (Anon. 1876, 1878). He also is listed as a local
fossil collector by Nickles (1936).
Herrlinger, Andrew
Herzer, Henry
Hill, H. H.
lb k
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Professor R. H. Holb roo k, of the Nat ion al No rm al Universi ty, Leb an on ,
W a r r e n C o u n t y , O h i o , lent th e m at er i a l u p o n w h i c h Stromatopora sub-
cylindrica originally was based (U. P. James 1884a).
L. M. H ose a was co-propriet or and c o-ed itor of the Cincinnati Quarterly
Holbrook, R. H.
Hosea, L. M.
Kemper, Willis
Lathrop, J. P.
Lesquereux, Leo
(1806-1889)
Lawyer; listed as a local collector by Shideler ([1952] 2002).
Do na te d a collec tio n of fossi ls to the Ci nc in na ti Soci ety of Natur al H
(Anon. 1882).
(Charl es) L e o Lesq ue reu x was bo rn in Switz er la nd in 1806 but mo v
the Uni ted States in 1848. He bec am e, perhaps , the foremos t paleo bot
of his day in the Un ite d States. He work ed for a nu m be r of the state ge
cal surveys, i ncl udi ng the I l l inois survey of A. H. Wo rt he n (q.v.), and
w o r k e d fo r t h e gr eat su rv ey of th e A m e r i c a n W e s t h e a d e d b y F . V . H a
(q.v.). Le squ ere ux aut hore d two papers on wha t were then conside red
land plants in the rocks of the typ e- Ci nc in na ti an . He died in 1889
E d w a r d O r t o n (q v ) was on e of his pallb ear ers (Bar tlet t 1962; Mer ril l
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Letton's Museum
E d w a r d O r t o n (q.v.) was on e of his pallb ear ers (Bar tlet t 1962; Mer ril l
1964; Lesquereux 1874,1878, 2006; Tri tt 2006).
Messrs. Letton and Willet founded a museum in Cincinnati in 1818
about the same time that the Western Museum was init iated by Dr. D
T h e Eng li sh ma n, Cha rl es Lyell , was one of the founders of the sc ien ce of
geology as we know it today. In 1842 he accepted an invitation from the West
ern Ac ad em y of Natural Scie nces to co me to Ci nci nna ti . His hosts were John
Locke, Robert Buchanan, J . G. Anthony, and Joseph Clark, and they v isi ted
localities in the area, including Big Bone Lick, Kentucky, world famous for
its remain s of Ice Ag e ma m m a l s (Hen dri cks on 1947; Lyell 1845).
Sidn ey Smit h Lyon was bor n in Cin ci nn at i . He was on e of the assistant
geologists in the geol ogi cal survey of Ke nt uck y by Da vi d Dal e O w e n (q.v.),
and h e author ed or co-au tho red a nu m be r of sc ienti f ic pape rs of fossi l
ech in od erm s of India na, K entu cky, and Oh i o (S. A. Mi l l er 1882a).
W W M a t h e r was c h i e f g eo lo gi s t o f t he f i r s t g e o l o g i c a l su rv ey o f O h i o
Lyell, Charles
Lyon, Sidney S.
(1808-1872)
Mather William
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W . W . M a t h e r was c h i e f g eo lo gi s t o f t he f i r s t g e o l o g i c a l su rv ey o f O h i o
(1836-1838). Assistant geologists included John Locke and Charles Whit
tlesey (Merrill [1924] 1964; S. A. Miller 1882a).
Of Ox fo rd , so listed as a local fossil col lec to r by Nic kl es (1936).
Mather, William W. (1804-1859)
McCord, D. A.
Miller, Samuel
Almond (1837-1897)
Milne-Edwards,
Henri
Misener, John
Misener, S. R.
(See chapter 2)
He n r i M i ln e -Ed w ar d s an d J u le s Hai me d e sc r i b e d the Ci n c i n n at i an
zoan s n ow kn ow n as Parvohallopora rugosa an d Dekayia aspera (C
Davis, and Utgaard 2002).
List ed as a col lec to r of fossils at Ri ch m o nd , In dia na, by Foerste (1917)
n a m e d Conchopeltis miseneri a n d Conularia miseneri after him.
Of R ic h mo nd , Indi an a, so listed as a local fossil col lec to r by Nic kle s (
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Mohr, Paul Mo h r was associat ed with the Cin ci nn at i Soc iet y of Nat ural History
1870s (S. A. Miller et al. 1879), and the cephalopod Orthoceras mohr
named after him (S. A. Miller 1875b). Mohr also was involved with Fred
Braun, another Cincinnati col lector , in excavations for cr inoids from
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Owen, David Dale
(1807-1860)
Patterson, W. J.
Plummer, John T.
D av i d D al e Ow e n , a l t hou gh n ot a me m b e r of the Ci n c i n n at i S c hoo
mu ch work in the region, and, ind eed, in m uc h of the Am er ic an Mid
He had c on n e c t i o n s w i th the W e ste r n Ac ad e m y of N atu r a l S c i e n c e
t h e C i n c i n n a t i a n b r y o z o a n , Fistulipora oweni, was named after
(Becker 1938; Croneis 1963; Hendrickson 1947; U. P. James 1879c, 1
Merrill [1924] 1964; S. A. Miller 1882a).
W i l l i a m J. Patterson w as lis ted as a loc a l foss il co l le c to r by W e t h e r b y (
an d b y N i c k l e s (1936) . T h e Ci n c i n n at i an c r i n oi d , Glyptocrinus patte
w a s n a m e d aft er h i m (S. A . M i l l e r 1882b).
Dr. P l um m er was the auth or of an early paper on the ge ol og y of the
about Richmond, Indiana, in which fossi ls are discussed and i l lust
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Probasco, Henry
(1820-1902)
about Richmond, Indiana, in which fossi ls are discussed and i l lust
(Plummer 1843).
Henry Probasco was in the hardware business in Cincinnati . He w
origi nal m em be r of the Cin ci nn at i So cie ty of Nat ural History, and h
J . M . Ri c har d son , o f W i lm i n g to n , Cl i n ton Cou n ty , Ohi o , fou n d the or i g i n a l
spe cim ens of the crino id species Glyptocrinus richardsoni Wetherby, 1880.
Dr. Ri ddell, a fa mou s botanist, was part of the faculty of the Me di ca l De par t
ment of Cinc inna ti C ol le ge, a short-l ived medi cal col lege foun ded by Daniel
Drak e (Writers' Progra m of the Work Projects Adm inis tra tio n in the State of
O h i o 1943). Abo ut the sa me ti me, Dr. Ridde ll was a curat or of the West ern
A c a d e m y of Natural S c i e n c e s , b e g i n n i n g in Apri l of 1835 (He n d ri c kson 1947).
Carl Lu dw ig Roem ing er was born in G er m an y and earned a med ica l degr ee
(but with a geo lo gic al thesis!) at Tu bi ng en . At the ti me of the 1848 revol u
tions, he moved to Cincinnati , which, at that t ime, had a large German-
speaking population, and he practiced medicine there for some twenty-five
Richardson, J. M.
Riddell, John L.
Roeminger, Carl
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speaking population, and he practiced medicine there for some twenty five
ye ars . He studied w hat are n o w k n o w n to b e foss il b r y o z o a n s — a s spa re t i m e
from his medical duties would al low. It was he who named Rhombotrypa
quadrata (Roeminger , 1866). About that same time, he moved to Michigan,
w h e r e he s i m u lt an e ou s l y c o n d u c t e d a m e d i c a l pr ac t i c e , wa s a profes sor of
geol ogy at the Universi ty of Mi ch ig an , and was Mich iga n's state geologist .
have been an extraordinary individual; for example, from 1874 to 188
ta ug ht at Har var d, was state geo log ist of Ken tuc ky , and wa s presiden
mining company in Montana, al l at the same time. In the upper echel
his profession, he served as president of the Geological Society of Amer
1895 and was a m em be r of the N ati ona l Ac ad em y of Sci enc es , but he als
a contributor to the Atlantic Monthly and to Scrihner's magazine and
five romantic dramas in blank verse. Although Shaler did author a paper
involved brach iopo ds of the Cin ci nn at i region , he was not real ly a me
of the Ci nc in na ti Sc ho ol (B eck er 1938; Cas ter 1951,1982; Cr on eis 1963; H
1907; Livingstone 1987; Shaler 1876; Shideler [1952] 2002).
W i l l i a m H e n r y Shi de ler w as b o r n i n W e s t M i d d l e t o w n , O h i o , an d grad
in 1907 from Mi am i Universi ty, Oxf ord , Ohi o. Im med iate ly upo n rece
his doct orat e at Co rn el l Univer sity in 1910, he retu rne d to his al ma m
Shideler, W. H.
(1886-1958)
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y 9 ,
w h e r e he sp en t t he re st o f his lif e on the faculty. He w as r e c o g n i z e d
except ional teach er , and , even tho ugh his publica tion record was not e
sive (Shideler 1914,1 916,1 918,19 39), his kno wl ed ge of the local rocks an
sils was extraordinar y, and his inf lue nce on coh orts of students and col le
was pr o fo u n d. A l t h o u g h n ot really a m e m b e r of the C i n c i n n a t i S c h o o
of As he rm an n, Sch le mm er, Vaup el, and others. He was e lected to me mb er
ship in the Ci nc in na ti S oci ety of Nat ura l History in 1885. H is large coll ect ion
of typ e -C i n c i n n a t i an b r yozoan s , i n c lu d i n g th i n -se c t i on s , w as b e q u e at he d to
the University of Ci nc in na ti . He served as curator of mic ro sco py at the society
and authored several scientific papers, although not on fossils (Anon. 1885a;
Caster 1982; Shideler [1952] 2002; Twitchell 1885,1887,1934).
(See chapter 2)
G e or ge Val l an d i gh am, of Ci n c i n n at i , w as an ac t i v e c o l le c to r a f ter w h om
Cyrtoceras vallandighami S. A. Mil le r, 1874 was na me d. He also col le ct ed
on e of the sp e c i m e n s on w hi c h the ge n u s of c ar p oi d e c hi n o d e r ms , Enop-
Ulrich, E. O.
(1857-1944)
Val landigham,
George L.
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p g p p
loura, was based (Wetherby 1879a). This is certainly the George L. Val-
landingham listed as a local fossil collector by Nickles (1936).
D e sc r i b e d the f ir s t-n ame d typ e - Ci n c i n n at i an b r y ozo an , Constellaria con- Van Cleve, J. W.
D r . W e l c h p r a c t i c e d m e d i c i n e i n W i l m i n g t o n , C l i n t o n C o u n t y , O h i o
or i g i n a l sp e c i me n s of Glyptocrinus richardsoni Wetherby, 1880, wer
W e l c h ' s c o l l e c t i o n ( W e t h e r b y 18 80 , 246; Foerste 18 93a , 18 93b) .
Char le s W e sse ls fou n d the sp e c i me n of Lepidocoleus jamesi describe
Faber , when he named the genus Lepidocoleus (Faber 1886,17).
T h e W e ste r n Ac ad e m y of N atu r a l S c i e n c e s w as fou n d e d b y D r . D
Dra ke in 1835. In 1840 the Natur al S ci en ce Sect ion of the Cinc in na
ciety for the Prom oti on of Useful K no wl ed ge , of whi ch John Go ul
thony was secretary, merged with the academy. In 1871 the rema
me mb er s of the Western Acad emy of Natu ral Sc ien ces donated a
assets of the acade my , in cl udi ng money , book s, and their col l ecti on, t
Welch, L. B.
Wessels, Charles
The Western
A c a de m y of
Natural Sciences
(1835-1871)
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Ci nc in na ti Soc iet y of Nat ural History, wh ic h had been found ed the
ous year (Anon. 1878,5; Hendrickson 1947).
A so ci et y w as f o r m e d in 1819 or shortly b efo re by a g r o u p , i n c l u d i n g D
k ( ) b l i h i i i i h
The Western
M
Merri l l [1924] 1964; Wh itf ie l d 1878). T h e "R. P. Wh it ef ie ld " in Nic kl es
(1936) must be the same person.
A l t h o u g h primari ly associated with the C l e v e l a n d area, C o l o n e l W h i t t l e s e y
ma de an agricul tural survey of Ham ilt on Co un ty , Oh io , in 1844. Prior to that,
he had been in cha rge of the to pog raph ic work for W. W. M ath er's 1836-1838
geo log ical survey of Oh io . Lat er he was party to an apparen tly vici ous disput e
as to wheth er New ber ry or Whi tt l ese y shoul d have be co me state geologist of
O hi o in 1869; New ber ry was so appointed (Ha nsen and Col l i ns 1979; Merri l l
[1924] 1964).
(S e e L e t ton ' s M u se u m)
Whittlesey, Charles
Willet
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J u dge W i l l i am W i lso n , o f L e b an on , W ar r e n Co u n t y , Ohi o , fou n d the typ e -
sp e c i me n of Prioniodus dychei U. P. James, 1884, and the three specimens
on w hi c h Polygnathus wilsoni U. P. James, 1884 was bas ed. He , D y c h e , an d
James were friends (U. P. James 1884c, 147-148).Wilson W m W
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GLOSSARY
How to pronounce scientific terms can be a real bugaboo. In the followingglossary, and elsewhere in this volume, we have included occasional advice
on how to pronounce terms. As you know, lexicographers have developed
a sch em e of sym bol s to indica te ho w they feel parti cul ar letters, syllables,
and words should be pro no un ce d. We hav e tried to ke ep the use of suc h
symbols to a minimum. We hope that, in so doing, we stil l have managed
to help you pronounce words in a way useful to you.
Unfortunately, not all scientific terms are included in dictionaries, not
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y, ,
even in the premier dictiona ry of the Engli sh la ng ua ge, The Oxford English
Dictionary (S imp son an d We in er 1989). T h e na me s of gen era an d spe cie s are
ve ry ra re ly inc luded, except in s om e s pec i a l i zed works. O n e that we have
found helpful is The Biologist's Handbook of Pronunciations (Jaeger 1960),
and there other similar works. Alas! Space here does not permit us to list all
ad ap er tu ra l (adj.) in ce ph al op od s, the direct ion toward the op en in g o
shell (and, h en ce , away from th e apex of the shell); so me workers p
to u se the syn on ymou s te r m " ad or a l" (cf.: adapical) .
ad ap ic al (adj.) in ce ph al op od s, the direc tion toward the apex of the
(and, he nc e, away from the op en in g of the shell) (cf.: adapertural
ad du ct or mu s cl e (n.) on e of the musc les of a bra chi opo d or a pel ec
w h e r e b y th e a n i m a l c l o se s its shel l (cf: d iductor muscle; adju
mu sc le ) .
adj ust or m us cl e (n.) on e of the mus cle s of a brach iop od whereb y the
mal mo ve s its shell with res pect to the ped ic l e (cf: adductor mu
d i d u c tor mu sc le ) .
ad ora l (adj.) adap ertu ral (q.v.).
ae gi sm (n.) "Associa tions for prot ectio n, either thr oug h cam ou fl ag e o
rounding vegetation, residence upon or within another animal, or
transport within the protective umbrella or aegis of a larger partner areco m m o n in the anim al king dom " " this term collects under its ba
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co m m o n in the anim al king dom . . . . this term collects under its ba
all those associations which are not principally concerned with food a
sition. . ." (Morton 1989,10) (cf: commensalism; mutualism; parasit is
ag e (n.) geo lo gi c tim e unit , equ iva le nt to stage in the time-s tratig raphic
sification; th e hierar chy of geo log ic ti me units, fro m largest to sm
is: eon era period epoch and age (cf : eon epoch era period)
b i n o m e n (se e: s pe c i e s n a m e ) .
b i o c l a u s t r a t i o n (n.) t he p roc ess w h e r e b y a soft -b odied o r g a n i s m that in
fests another organism or colony is embedded by the skeletal growth
of that other orga nism or colony. T h e word was coin ed for instance s
in which a particular parasitic organism, suggested to be a hydroid or
a colonial asc idiacian tunicate , was engulfed by the bryozoan colony
on wh ic h it was gro wi ng ; the result is a patt ern of hol es call ed Catel-
locaula vallata Palmer and Wilson, 1988.
b i o f a c i e s (n.) character ist ics of a s e d i m e n t a r y rock based on fo ss il c o n te n t
(cf.: facies, lithofacies).
b i o h e r m (n.) a feature that is e lev ated a b o v e th e sea floor a n d w as pr o
duced by organisms (Davis 2004, 283); a f ine, upstanding example is a
coral reef, (cf.: b iostrome).
b i o i m m u r a t i o n (n.) u su a l l y an e x te r n a l m o l d of a soft-bodied o r g a n i s m
that was ove rgro wn by a calca reo us org ani sm, fo l lowed by deca y of thesoft bodied organism
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soft-bodied organism.
b i o s t r a t i g r a p h i c u n i t (n.) a b o d y of rock c h a r a c t e r i z e d by fo ssi ls of a pa r
ticul ar species or gr ou p of spe cie s; at a give n local ity, this is repre
sented by the thickne ss of strata so chara cter ize d, ( c f : bio zo ne, l i th-
ostratigraphic unit , t ime-stratigraphic unit) .
b i o s t a t i g a p h (n ) st d of d is t ib t i on of fo ss il s in s e d i m e n t a ocks
ca st (n.) a replic a of an orga nis m ma de from a mold of that org ani sm
mold ) .
C ha e t o g n a th a (n. ) a smal l p hy lu m of mar i n e i n v er te br ate s l i v i n g m
as plankton, commonly cal led "arrow worms"; fossi ls are known
the Pennsylvanian, and possibly the Cambrian (Valentine 2004).
ch it in (n.) a co mp le x org anic materia l pro duc ed by arth ropod s, fo
ample, by insects; the comparable material produced by mollusc
te r me d c on c hi ol i n , a l thou gh some p e op le u se the te r m " c hi t i n
refer not only to chitin, in the strict sense, but to conchiolin and o
such organic materials .
ch it in oz oa ns (n.) a gro up of ma rin e plan kto nic microfossi ls with gene
flask -shap ed organ ic walls; of unc ert ai n aff init ies, but presu mab ly
mals (Jansonius and Jenkins 1978).
C ho r d at a (n.) an i mal p hyl u m c har ac te r i z e d b y the n otoc ho r d , i n c l
the vertebrates.Ci n c i n n a t i Ar c h (n. ) a br oad u p w ar p i n g of s tr ata e xte n d i n g fr om
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Ci n c i n n a t i Ar c h (n. ) a br oad u p w ar p i n g of s tr ata e xte n d i n g f r om
tu c ky thr o u gh Oh i o .
Ci n c i n n a t i an (n. ) ge ol ogi c age te r m for the L ate Or d ov i c i an Ep o c
N o r t h A m e r i c a .
Ci n c i n n a t i a n S e r i e s (n. ) t i me -str at i gr ap hi c te r m for the Up p e r Or d
c i an i n N or th Ame r i c a
co pro li te (n.) a sp ec im en of fossi l ex cre men t (from the Gr ee k "ko pro s" +
" l i thos , " me an i n g " d u n g" + " r oc k" ) .cor rel ati on (n.) process of de te rm in in g the corr es po nde nce of strata in
different sections or locations.
co qu in a (n.) a kind of sed ime nta ry rock that consists alm ost exclusi vely of
shells , shell-fragments, or both (from the Spanish for " l i tt le shell" ;
p r on ou n c e d : ko-ke e n -u h) .
cotypes (n.) in taxonomy, when two or more type-specimens are desig
nated for a speci es, and all of th em are con si der ed to be equa ll y repre
sentative of the species, they are cal led coty pes . So me ti me s, in forme r
years, th e te r m " s y n t y p e " w as u s e d (cf.: holotyp e ; p ar atyp e ) .
cy cli cit y (n.) in stratigraphy , the regu lar repetit ion of a patt ern or or der in g
in sedimentary strata.
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de at h as se mb la ge (n.) fossi ls in a strat um that represent rem ain s of org an
isms that died at so me tim e before f inal burial , and usual ly ac cu m u
lated from various or iginal locations [cf.: l i fe assemblage).
den ti t i on (n.) col l ectiv e na m e give n to the teeth and sockets al on g the
hi n g e l i n e of an ar t i c u la te b r ac h i op o d or p e l e c y p od ; the te e th an d
D
do rs al (adj.) on or toward the top of the an im al ; the no un for m o
concept is "dorsum." (Note that what is functionally dorsal in a g
an im al ma y not cor res pon d to the anato mi cal dorsal . For exa mpl
humans, the head is anatomically anter ior , but, as you walk aro
y o u r h e a d i s a t th e top of y o u r b o d y a n d , h e n c e , is f u n c t i o n al l y
sal"; cf: anterior; distal; lateral; medial; posterior; proximal; ventr
d u ro p h a g o u s (adj.) said of predat ors that cru sh the shells of their pre
ec ol og y (n.) 1 . the study of the en vir on me nt; 2 . the env iro nme nta l n
of org ani sm s of a giv en taxon or gr ou p of taxa (cf: au te c o
syn e c ology) .
ec os ys te m (n.) a l l the org ani sms in a give n t im e and place, a l ong wit
ch em ic al and physica l aspects of the envi ro nm ent in whic h they
in that time and place.
E de n i a n (n ) term for the low erm os t stage of the Ci nc in na ti an Series
E
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E de n i a n (n.) term for the low erm os t stage of the Ci nc in na ti an Series
the e arl i e s t age of Ci n c i n n at i a n Ep o c h ,
e n d o sy m b i o n t (n .) or gan i sm l i v i n g w i th i n the b od y of an ot he r l
or gan i sm.
eo n (n.) largest geo lo gic t im e unit , as in Phan ero zo ic Eo n, wh ic h
era (n.) a geol ogic t ime unit , as in the Paleoz oic Era, wh ic h includes , am o ng
others, the Ordov ic ia n Period; the hierarch y of geol ogi c t ime units , fro m
largest to smallest is: eon, era, period, epoch, and age (cf.: age, epoch,
eon, period).
eus ta ti c (adj.) gl ob al , refer ring to ch an ge s of sea level .
eve nt ho ri zo n or ev en t be d (n.) a sed ime nta ry layer for med by a short-
term depos it ion al pro cess or dis tur ban ce, su ch as a layer of volcanic-
ash, or a storm bed.
exo sk ele to n (n.) shell or car apa ce that encl ose s the bod y of an an im al .
ex te rn al m o l d (n.) a mol d of the exterio r of an ob jec t (cf: internal mold;
steinkern).
fac ies (n.) any charac teri stic s of rock s us ed for re co gn it io n of lateral or
vertical var i at i on s , a n d u s u a l l y r e f le c t i n g the p r o c e ss e s f o r m i n g t h e
rock (cf: biofacies; l i thofacies)
F
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rock (cf: b iofacies; l i thofacies).
fa cu lt at iv e (adj.) refers to an in sta nce of an org an is m that is a party to an
association that may be convenient, but is not required for the survival
of the org an ism ; if , on the othe r ha nd , a gi ven parasite is un ab le to
exist outs ide of its part icu lar assoc iat ion wit h its hos t, the assoc iat ion
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in te rn al m o l d (n.) a mol d of the interior of an objec t; the interna l mo ld of
a shell sometimes is cal led a ste inkern (q.v.) (cf.: external mold),
is oc hr on ou s (adj.) equi val ent in age.
K-b ent oni te (n.) a potas sium- rich c lay min era l for med by ch em ic al a ltera- K
tion of vo lc an ic ash depo sit ed in salt water,
k i n g d o m (n.) the le v e l i n the L i n n ae an hi e r ar c hy ab ov e p hy lu m .
lag d e p os i t (n . ) se d i me n tar y ac c u mu lat i on r e mai n i n g a f te r e r os i on of L
finer-g rained size fractio ns or ty pes of se di me nt .
lat era l (adj.) on or tow ard the side of th e an im al (cf: anterior; distal; dorsal;
medial; poster ior; proximal; ventral) .
Lebensspur (pl., Lebensspuren; n.) a trace fossil (= ichnofossil). ("Lebens-
spur" is a Ge r m a n word that l iterally me an s "tra ce of l i fe"; the Ge r m a n
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spur i s a Ge r m a n word that l iterally me an s tra ce of l i fe ; the Ge r m a n
w or d " S p u r " is related to t he a r c h a i c E n g l i s h w ord "spoor.")
life as se m bl ag e (n.) fossils in a str atu m that represent rem ai ns of org an
isms that were living at the time of final burial, usually at or near the
site of burial (cf: death assemblage).
m e d i a l (adj.) on or tow ard the mid dl e of the ani ma l, that is, towar d
p lan e o f sy mm e tr y of the an i ma l . S o m e p e op l e , to av oi d p ote
confusion with the statist ical term "median," use the term "me
(cf.: anter ior; distal; dorsal; lateral; poster ior; proximal; ventral) .
m eg ac y cl e (n.) repeate d set of sed im ent ary strata of abo ut on e to two
ters in thickness.
m e m b e r (n.) l i tho strati graphi c subdiv ision of a form atio n.
m es o de rm al (adj.) per tai nin g to the mi dd le layer of t issue in the bod y
of tr iploblastic an im al s (those ha vi ng thr ee embr yo nic t issue la
e c t o d e r m , m e s o d e r m , a n d e n d o d e r m ) .
m e t am o r p h i s m (n. ) p r oc e sse s of c h e m i c a l or p hys i c a l a lte r at ion of
und er the i nf l uenc e of heat , pressure, or bo th (cf: d iagenesis) .
m et az oa ns (n.) a lternative term for "an ima ls. "
Mi la nk o vi tc h cycl e (n.) repeate d pattern of events or strata on a frequ
simi lar to peri odic i ty of on e or mor e variations of the Earth's o
such as obliquity or eccentr ic i ty. Named after the Yugoslav mathe
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c i an M i lu t i n M i lan kov i tc h (1879-195 8) .
m i n e r a l i z a t i o n ( = r e p l a c e m e n t ) (n.) an a l t e r n at i v e f or " r e p l a c e m
(q .v.).
m o l d (n.) an imp ress ion or ot he r neg ati ve replica of the intern al or e
nal parts of an orga nis m form ed by se dim en t enc lo sin g the orga
on tog e n y (n. ) e v e r y thi n g that hap p e n s to an or gan i sm f r om the b e gi n n i n g
of i ts l i fe to deat h. I ncl uded wit hin "on to gen y" are emb ryo log y, devel
op me n t , gr ow th, matu r at i on , an d so on (cf.: t a p h o n o m y ) .
order (n.) the level in the Linnaean hierarchy below c lass and above
family.
Or d o v i c i an Bi od i v e r s i f i c a t i on Ev e n t (n. ) ap p e ar an c e of n u m e r o u s maj or
groups of skel eton ized ma ri ne invertebrate org ani sms duri ng the Or
dovic ian Period (syn. , Ordovic ian Radiation).
Or d o v i c i an Pe r i od (n. ) ge olo gi c t i me u n i t fo l lo w i n g the Ca mb r i an Pe r iod
and prece din g the Si lur i an Period.
ou tc ro ps (n.) surface exposur es of bed roc k for med by natural processes .
pa ck st on e (n.) a l im esto ne cha rac ter ize d by frag men ts in grain-to -grain
contact, with intersti t ial calcareous mud (cf: grainstone).
pa le ob at hy me tr y (n ) dep th of an anc ien t bo dy of water
p
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pa le ob at hy me tr y (n.) dep th of an anc ien t bo dy of water .
pa le on to lo gy (n.) the study of anc ien t l i fe based on ev id en ce provid ed by
fossi ls (from the Greek "palaios," meaning "ancient," + "on, ontos,"
me an i n g " b e i n g, " " th i n g," or " that w hi c h has e x i s te n c e ," + " logos ,"
meaning "discourse," and, by extension, "study of"); the word also
pl an kt on ic (adj.) sus pen ded wit hin or floati ng up on a bod y of water,
transportation exclusively or primarily by currents and other m
me nt s of the wat er itself (cf.: benthic; nektonic) .
po ly mo rp hi sm (n.) the existence i n one species of individuals of
than one size, shape, or nature; in bryozoans, for example, there
be several different p o l y m o r p h s in a g iven colony, e a ch of w h i c h ,
sumably, performed a different funct ion (cf.: d imorphis m) .
po st er io r (adj., n.) on or towa rd the rear of the ani ma l. ( Not e that wh
functionally posterior in a given animal may not correspond to
ana tomi cal posterior. For exa mpl e, in hu ma ns , the back is anat
ca lly dorsa l, bu t, as yo u wal k ar ou nd , yo ur back is at the rear of
bo d\ an d. h e n c e , i s funct ional ly "posterior"; cf.: . i n t e r i o r ; distal; do
lateral ; medial ; proximal; ventral .)
pr ed at io n (n.) a relationshi p in wh ich one an im al cons um es anot her o
ism; the consumer is called a predator, and that consumed is calledprey (cf: aegism; commensal ism; mutual ism; parasit ism; symbiosis
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pr im ar y pr od uc er s (n.) org ani sms at the base of a food cha in that
organic compounds such as carbohydrates from simple compon
suc h as carb on dio xid e and water by the proces s of photos ynthe s
priority (n.) in taxonomy, the convention that, when there is an o
R i c hm o n d i a n (n. ) the you n ge st s tage of the Ci n c i n n a t i an S e r i e s ; a lso the
y o u n g e s t a g e o f th e C i n c i n n a t i a n E p o c h ,
r oc k- str at i gr ap hi c u n i t (se e l i thostr at i gr ap hi c u n i t ) .
sc ien ti f ic n a m e (see: speci es nam e).
sea f loor spr ea di ng (n.) proce ss by wh ic h new oc ea ni c crust is for med by
upw el l in g of mol ten m ater ial a l on g a f issure or r i ft .
se is mit e (n.) rock layer for med or altered by ear th qua ke shock.
ser ial sec ti on in g (n.) cut ti ng a fossi l , for ex am pl e, a brach io pod , into a
n u mb e r of th i n , e q u al l y sp ac e d s l i c e s of kn ow n or i e n tat i on . Fr om
the se slices, it is poss ible to rec on st ru ct the int erior of th e she ll; in ef
fe c t, on e i s d oi n g a p a le on tolo gi c a l C A T sc an . In sp e c i me n s i n w h i c h
the interior of the shell is fi l led wi th rock ma tri x, this ma y be the on ly
practical way to see what is inside the shell.ser ies (n.) a t ime-s tratig raphic subdivis ion of a syst em, such as C in ci nn a
s
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t ian Series or Upper Ordovic ian Series (cf.: stage; system).
se q u e n c e b o u n d a r y (n. ) s tr a t i gr ap hi c hor i zon ma r ki n g the b e gi n n i n g or
ter mi nat ion of strata dep osi ted du ri ng an interval of sea level rise or
fal l , usually in dica tin g a cessatio n of sed ime nta tio n or interval of ero
st ei nk er n (n.) petri fied sed im en t infi lli ng of a shell or bo dy cavity, a
ternal mold (from the German "Stein," meaning "stone," + "Ke
m e a n i n g " k e r n e l " ) ; a l t h o u g h a G e r m a n n o u n , i t c o m m o n l y a p p
neither capital ized nor in i tal ics (cf.: cast; mold).
sto rm cy cle s (n.) a layer of sed ime nta ry rock pro duc ed duri ng a s
stor m, co m m on ly i t consists of an interval repr esent ing intense di
b a n c e o f th e sea fl oor fo l l ow e d b y an interval r ep re se n t i n g w a n i n
storm activity.
st ra ti gr ap hy (n.) the study of lay ered or stratified rock s (cf: biostratigrap
str o mato l i te s (n. ) hn e K lami n ate d se d i me n tar y ac c u mu l at i on s r e su
from trap ping and bind in g of sed ime nta ry partic les by cyano bact
u su al ly she e t- l i ke , d ome -shap e d or c o lu mn ar i n for m; some t
called "algal stromatoli tes," because cyanobacter ia once were tho
to be algae.
str om ato por oid s (n.) dom e-s hape d, col umn ar, or bran chin g calcareous
etons, usually with f inely lami nate d mierostru cture, formed by cal
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ou s sp on ge s , oc c u r r i n g i n Or d ov i c i an thr ou gh Cr e tac e ou s r oc ks .
su bs pe ci es (n.) a level in the Lin na ea n hierarc hy just bel ow spe cies ; s
t imes cal led variety.
su bs tr at um (n.) a surfa ce on whic h an org ani sm mi gh t be atta che
mi gh t be the surfa ce of a strat um ro ck shell or oth er hard ob
rentian plate; nam ed for the ' l ac on ic Mo un ta in regio n of eastern Ne w
York w h e r e this d e f o r m a t i o n fi rst w as r e c o g n i z e d .
ta ng ent ia l (adj.) with respec t to bry ozo ans , referr ing to a secti on cut paral
lel to the surf ace of the zo ar iu m an d close en o ug h to the surfac e of the
colo ny to sho w the ma tur e features of the zoo ec ia in cros s-section ;
such a tangential section is perpendicular to a longitudinal section
(Bassler 1953, G15, G18) (cf.: longitudinal; transverse).
ta ph o no m y (n.) 1 . every thi ng that ha ppe ns to an org ani sm after i t dies; 2 .
the study of those ph en om en a that tend to destroy the rema ins and
traces of a liv ing th in g as well as of tho se that tend to pre serve t hos e
r e mai n s an d tr ac e s ( f r om the G r e e k " tap hos ," me an i n g " gr av e " or
" t o m b , " + " n o m o s , " m e a n i n g " l a w " ) .
tax oba ses (sing. , taxobasis; n.) charact er istic s on wh ic h the ta xo no mi c po
si t ion of a gro up of org ani sm s is bas ed (p ro no unc ed : tax-uh- base-
ease).
ta xo n (pl., taxa; n.) a for mal bio lo gic al gro up to wh ic h an or ga ni sm is as
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signed; humans, for example, belong in a taxon at the family-level of
the L i n n ae an hi e r ar c hy c a l le d Ho mi n i d a e .
ta xo no my (n.) the sc i ence of ass igni ng orga nism s to their proper biol ogica l
groups (alternative names are c lassi f ication and systematics).
i ( ) di k f d b f
un iv al ve d (adj.) said of a shell tha t cons ists of a singl e valve (cf.: bival
p se u d ob i v a lv e d ) .
v a r i e t y (n.) in fo r m e r t i m e s , e q u i v a l e n t to t he t a x o n o m i c rank of sub
c i e s , b u t n o lon ge r r e c ogn i ze d w i thi n the zoologi c a l c ommu n i ty
te r n at i on al Co mm i s s i on on Z oo log i c a l N o me n c la tu r e 1999,19) ; n o
days co m m o nl y used speci f ical ly for kind s of p lants bred delibera
by hor t ic u l tu ri s ts .
v e n t ra l (adj .) on or toward the b o t t o m of the a n i m a l ; the n o u n form o
concept is "venter." (Note that what is functionally ventral in a given
ma l may not cor resp ond to the an at om ica l ventral. For exa mp le, in
mans, the belly is anatomically ventral , but, as you walk around, y
bel ly is at the fr ont of yo u r b o d y an d , h e n c e , is fu nc t ion al ly "a nte ri or"
anterior; distal; dorsal; lateral; medial; posterior; proximal.)
V
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zo ne (see bioz one) .z
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(Upper Ordovician): Palaios 17(6): 545-555.
We bb y, Barry D. 2002. Patterns of Ordovician Reef Development, 129-179. In
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, Klorentin Paris, Mary L. Droser, and Ian G. Percival, eds. 2004. The Great
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, Robert J. Elias, Graham A. Young, et al . 2004. Corals , 124-146. In Barry D.
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We is s, Malcolm P. 1961. The American Upper Ordovician Standard. V. A Critical
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Expl anat ion of the Holes in Ord ovi cia n Shells An aly zed by Kaplan and
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INDEX
Page numbers in italics indicate illustrations.
I .ike most indexes, this one is nowhere near as thorough as it mighthave been; the tremendous number of items that might have beenincluded would have made the index completely unwieldy.
Certain words and concepts are not separately listed in theindex, because they appear far too many times in the book. Ex amples include Cincinnatian, Ordovician, and Ohio River.
Although places in the states of Indiana, Kentucky, and Ohioare included in the index, every appearance of the name of the
whole state is not.This volume is not intended as a comprehensive taxonomicwork. Hence, we have not endeavored to put every genus, species,and so on, in its complete Linnaean Hierarchy in the index.R h h d h f l l h
Chlorophyta ("green algae"), 67, 68
Dasyeladaceae, 67, 68
Anomaloides: A. reticulatus, 68Cyclocrinites: C. darwini, 66, 67 -68
Ischadites: I. circularis, 68Lepidolites: L. dickhauti, 66, 67 , 68
Cyanophyta (called "blue-green algae," but not actually
algae) . See Cyanobacter ia
Cylindrocoelia: C. covingtonensis, 68Faberia: F. anomala, 68
Girvanella. See Cyanobacter iaoncolites, 67
Phaeophyta ("brown algae"), 67
Pyrrophyta: dinoflagellates 69 239 248 283
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Rather, we have used the names of upper-level taxa that appear inthe text. However, we have departed from this policy in some cases,with the hope of greater clarity and usefulness for the reader.
Al h h ll d " " hi lik "b i l
Pyrrophyta: dinoflagellates, 69, 239, 248, 283
Rhodophyta ("red algae"), 67, 68
Solenopora: S. richmondensis, 68stromatolites. See Cyanobacter ia
Nereigenys: N. alata, PL 5 Annual Reviews, 4
anomala, Faberia. See Algae
Anomalocrinus. See Echinodermata: Crinoidea: Disparida Anomalodonta. See Mollusca: Pelecypoda
Anomaloides. See Algae: Chlorophyta: Dasycladaceae
An tho ny, Jo hn Gould, 259-260 , 261, 26 9, 27 6
An tho zoa, anthozoans . Se e Cnidar ia
An ti co st i Isl an d, Ca nad a , 72 -73
Aphelognathus. See c o n o d o n t s
Ap la eo pl io ra . See Mollusca
Appalachian Mounta ins , bas in , 3, 5, 8, 10, 53 , 61 , 21 6, 220,231, 233, pl. 12
approximate, Palaeasterina. See Echinodermata: Asteroidea
Arachnida. Se e Ar thro pod a: Che l i cerata
aragonite, aragonitic. See calcium carbonate
area, Archinacetla. See Mol lusca: Monoplacophora
Ar chae oga stropod a. See Mollusca: Gastropoda
Archinacella. See Mol lusca: Monoplacophora
Arnhe im Fo rm ati on, 44 , 48 , 53 ,68 , 105, 107, 10 8, 123, 151,153, 164, 176, 180, 196, pl. 8
Oregonia Member, 44, 53, 56, 63, 223. See also Oregonia
Formarion
Eoleperditia, 163Lepcrditicopida, 164
Palaeocopida, 163
Podocopida, 163 Quadrijugator: Q. regularis, 160, 161
Diplopoda: mill ipedes, 147
Insecta, 105, 119, 147, 162, 282
scorpions. See Chelicerata: Arachnida
spiders. See Chelicerata
ticks. See Chelicerata: Arachnida
Tri lo bi ta, 6, 7, 27 , 31 ,40 , 56 , 6 0 , 9 2 , 9 3 , 117, 137, 14
147-157, 1 60- 161 ,16 2,1 63, 202 , 203, 204207, 210, 212, 219, 220, 223, 224, 225, 22
229 , 231 , 232 , 233 , 235 , 238 ,240 , 2 41 , 244
247, 251, 252, 253, 260, 268, 275, 286
Acidaspis. See O d o n t o p l c u r i d a
Calymene , 149, 150, 296, 305
C. meeki. See Flexicalymene meekiC. meeki-retrorsa. See Flexicalymene retrorsa
C. senaria. See Flexicalymene senariaCalymenida, calymenids, 149, 151, 226, 241
Flexicalymene, 40, 148, 150, 151, 152, 153, 154,207, 210, 224, 225, 226, 229, 232, 251, 253
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Sunset Member , 44, 53, 56, 63, 164, 224. Se e also Sunset
Formation
Arnheimograptus. See Hemichordata: Graptoloidea
298, 317
F. granulosa, 149, 150, 151,309
F. meeki, 148, 149,150, 151,207, 228, 229, 2
Platycoryphe: P. christyi, 157Tricopelta: T. breviceps, 157
Platycoryphe. See P h a c o p i d a
Primaspis. See O d o n t o p l e u r i d aProetus parviusculus. See Decoroproetus parviusculustrace fossils, 151-153. See also taphonomy: trace fossils
Rusophycus. See taphonomy: trace fossils
Triarthrus. See Olenida
Ar ti cu la ta . See Brachiopoda
Asaphoidichnus. See taphonomy: trace fossils
As co ce rid a. See Mollusca: Cephalopoda: Naut i loidea
Ashermann, George (? ), 27 3, 27 5 As hgi l l ian Stage, pl. 2
associations. See chapters on individu al groups of ani mal s;
ecologic associations
Asteriacites. See taphonomy: trace fossils
As te ro id ea . See Echinodermata
Astraspis. See Chordata: Vertebrata: Agnatha
auhurnensis, Platystrophia ponderosa. See Brachiopoda:
Platystrophia ponderosa Au du bo n, Jo hn James , 16, 27 7
Aulacera. See Porifera: Stromatoporoidea
Au si ch , W i l l i a m I„ 186
Bellevue Limestone, Jl, 12, 53, 55, 56, 63, 68, 85, 89, 95, 99,
125, 174,214, 223, 232, 2 5 1 . See also McMil lan
Eormation
Beloitoceras. See Mol lusca : Cephalopodabenthic, benthonic, benthos, 3, 4, 7, 50, 102, 145, 163, 195,
204, 205, 218, 230, 234, 237, 239, 280, 288, 290
bentonite. See K-bentonite
Berdan, Jean, 163, 164, 320
Bergman, Claes F., 143, 304
Bergstrom, Jan, 153, 318
Bergstrom, Stig M., 2,51, 62, 196, 306, 308, 319, 321, 331
Best, Robert, 260, 276bibliographies, compilation of, 15, 33, 35
biclavata, Diplocraterion. See taphonomy: trace fossils
bilix lata, Cyclonema. See Mollusca: Gastropoda
bilix, Cyclonema. See Mollusca: Gastropoda
binomen. See taxonomy: nomenclature: species name
Binomial System of Nomen clature . See taxonomy:
nomenclature
bioclaustration, 156, 210, 243, 281. See also taphonomy:bioimmurat ion
biocoenose, biocoenosis . See life assemblage
biofacies. See facies
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Au sti n, George M. , 26 0
Austinella. See Brachiopoda: Articulata
Au str al ia , 199 , 215 , 24 1, 25 1. See also Great Barrier Reef
bioh erm, 71, 72, 78 -8 1, 85, 91, 131, 242, 281
"bryoherm," 91
Great Barrier Reef, Australia, 91, 241
92,93,98-115, 119, 123, 124, 125, 127, 128,
129, 130, 137, 144, 162, 172, 178, 179, 182, 188,
189, 206, 219, 220, 221, 223, 224, 225, 231, 232,
233, 235, 237, 240, 241, 242, 243, 244, 245, 248,250, 251, 252, 253, 260, 261, 272, 274, 280, 281,
283, 285, 287, 291, 293
Articulata , 4 0 , 93, 100, 101, 102, 103, 105, 112, 130, 261,
283 ,291 ,293
Austinella, 112 A. scovillei, 273
Catazyga, 112
C. schuchertana, 111Dalmanella, 57, 113, 114, 225, 232, 233, 240, 253D. emacerata, 106
Glyptorthis, 112, 220, 224, 233G. insculpta, 107
Hebertel la, 100, 104, 105, 113, 223, 233, 240, 251H. occidentalis, 107
Hiscobeccus, 112, 220, 224, 240
H. capax, 111Holtedahlina, 112 ,220 ,240
Lepidocyclus, 75, 220Leptaena, 40 , 112, 220, 224, 233, 240
L i h d i 40
Ungulate brachiopods, lingulides, 103, 105, 247,
Petrocrania: P. scahiosa, 40, 104, 105Philhedra: P. laelia, 104, 105, 108
Pseudolingula, 104, 105, 225. See also LingulaSchizomania: S. filosa, 104, 105Trematis, 128
T. millepunctata, 40, 104, 105Brachiospongia. See Porifera
brachycephalus, lsotelus. See A r t h r o p o d a : T r i l o b i t a
Bracken County, Kentucky, 178, 209
Bradshaw, Lael E., 22, 24, 25
Brandt, Danita S., 151, 153Branson, C. C, 196
Branson, E. B., 196, pl . 5
Branstrator, Jon W., 182, 184, 189
Braun, E. Lucy, 260
Braun, Frederick, 260 -2 61 , 270
Bretsky, Peter, 231
Brett, Carlton E., 60, 61, 236, 295, 297, 310, 311, 318
breviceps, Tricopelta. See Arthropoda: Trilobita: PhacoBridge, Josiah, 261
British Museum (Natural History). See Natural Histor
seum, London
b i l S E hi d O hi id
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L. richmondensis, 40Onniel la , 112, 124
O . meeki , 60O hi O ill i (S A i ll ill i)
brittle stars. See Echinodermata: Ophiuroidea
Broaddus, Bill ie, 29
Brongniart, Alexandre, 61
60,67, 71, 72, 79, 13?, 144, 145, 169, 188, 206,
239, 280, 281,289,292
calcite, calcitic, 6, 7,74, 89, 101, 117, 120, J24, 130, 137,
139, 163, 167, 168, 188, 280, 281, 283, 290calcium phosphate, 6, 81, 102, 182, 196, 198, 219, 223
Calloway Creek Formation, 214
Calymene meeki. See Arthropoda: Trilobita: Flexicalymenemeeki
Calymenida, calymenids. Sec Arthropoda: Trilobita
Calvptomatida. See hyolithids; Mollusca
Cambrian, xix, 2,4,6, 30, 82,99, 139, 147, 152, 160, 163,
167, 168, 182, 186, 191, 195, 199, 237, 249, 280,282, 289
"Cambrian Explosion," 2
"Cambrian Fauna," 2, 237
Camerata . See Echinodermata: Crinoidea
Cameroceras. See Mol lusca: Cephalopoda: Endoceratoidea
camouflage, 96, 280
campanulaeformis, Cyathochitina. See chitinozoans
Campbell County, Kentucky, 59, 109, 203, 209Campbell, Kenton S. W., 155, 160
Canada , 3 ,215,220, 233,249,253
Anti cost i Is land , 72 -73
Charactoceras. See Mollusca: Cephalopoda
Chee tham.A l anH. , 185 ,298
Chei lostomata. See Ectoprocta
Cheirocystis. See Echinodermata: Rhombifera
Chelicerata. See Arthropoda
Chicago, University of. See University of Chi cag o
Chilopoda. See Arthropoda
chitin, chitinous, 6,69, 82, 119, 147, 148, 161, 181, 282. See
also eonchiolin; pseudochitin
chitinozoans, 52, 68, 69, 247, 282. See also Algae
Conochitina: C. hirsuta, 66, 67
Cyathochitina, 66, 67; C. campanulaeformis, 66, 67Hercochitina: H. turnbulli, 66, 67
chitons. See Mollusca: Polyplacophora
Chlorophyta. See Algae
Chondrites. See taphonomy: trace fossils
Chordata, 38 , 169, 196, 198, 282. See also conodonts
Urochordata: tunicates, 210, 281. See also Catellocaula Ve rt eb ra ta
Agnatha (" jawl es s fi sh ") , 199, 200 Astraspis: A. desiderata, 199gnathostome fish ("jawed fish"), 200
Osteichthyes ("bony fish"), 198
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British Columbia, 6
Canadian Shield, 3, 216
Manitoba, 153
Reptilia: Virginia, 41
Christy, David, 262
christyi, Platycoryphe. See Arthropoda: 'Trilobita: Phacopid
Gincinnaticrinus. See Echinodermata: Crinoidea: Disparida
Cincinnatidiscus. See Echinodermata: Edrioasteroidea
cincinnatiensis, Acidaspis. See Arthropoda: Trilobita:
Odontoplcuridacincinnatiensis, Diplocraterion. See taphonomy: trace fossils
cincinnatiensis, Isorophus. See Echinodermata:
Edrioasteroidea
circular is, Ischadites. See Algae: Chlorophvta: Dasvcladaceae
Cladida . See Echinodermata: Crinoidea
c h i l l i s . See Mollusca: Pclccvpoda
Clark, Thomas H., 3
Clarkson, Euan N. K., 160, 299Clarksvi l le Member. See Waynesvil le Formation
class. See taxonomy: l . innaean Hierarchy
c l . i s s i h i ation. Sec lavonoim
clastic ratio, 49. See also shale-to-limestone ratio
clavacoidea: Leptotrypa. See Ectoprocta: Trepostomata
Clays Ferry Formation, 163, 214
Clermont County, Ohio, 111, 126, 128, 153, 158, 163, 170,
186, 203, 207, 208, 209, 261Stonelick Creek, 11,254
Climacograptus. See Hcmichordata: Graptoloidea
Clinton County, Ohio, 60, 72, 75, 128, 136, 142
Wi l i 260 27 3 27 6
polyps, 74, 77,79, 182, 206, 235
reefs. See biohcrms
Scyphozoa, 74, 82, 182. See also Byroniida
Conulariida, 74, 81-82, 241, 247; but may not be cndarians, 74, 81-82, 241, 247
ConulariaC. formosa, 80, 82 , 267, 269C. miseneri, 270
sea anemones. See Anthozoa
sea fans, 74
sea whips, 74
coarsening-upward cycle. See cycles, cyclicitycoccinea, luhastraea. See Cnidaria: Anthozoa: Scleractin
cockles. See Mollusca: Pelecypoda
Code of Stratigraphic Nomenclature. See stratigraphy
(loclenterata. Sec I Inidaria
co il in g of shell , 117, 119, 120, 132 , 134, 139
planispiral coiling, 119, 120, 139, 289
Colbath, G. Kent, 67, 69
Cold Spring, Kentucky, 174Goleolus. See hyolitbids
collagen, 195, 196
colonial organisms, colonies, 189, 281, 290. See Cnidaria
E l li
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Wi lmington, 260, 27 3, 27 6
cluster analysis. See techniques
C M C . See Cincinnati , Ohio: Cincinnati Museum Center
Ectoprocta, graplolites
color patterns, 96, 134, 135, 137, 252, pl. 6
Colorado, 199, 216
Cooper, Edward M, 263
coprolites. See feces
coprophagy. See food
copulation, 162
coquina, 99 , 283. See also shell pavements; shingled beds;
Waynesv i l le Form atio n: Marb le Hi ll be d
coral. See Cnidar ia
coral reef. See bioherm
Corallidomus. See Mollusca: Pelecypoda
coralline sponges. See Porifera: sclerosponges
Cornulites, Sec: Annelida
correlation, 52. See also time
Corryville Formation, Member, 12, 21, 53, 55, 60, 63, 68, 80,
90, 105, 110, 111, 114, 120, 126, 128, 142, 144,
151, 153, 158, 163, 170, 174, 175, 178, 180, 186,
203, 207, 208, 209, 211,214, 224, 251. See also:
Grant Lake Limestone; also: McMillan
Formation
cotypes. See taxonomy: type specimens
Covington Croup, 48
Covington, Kentucky, 19, 28, 127, 142, 256, 263, 266, 274
covmgroiiensis, Cylindrocoelia. See Algae
Cowen. Richard, 199
eustatic sea-level change, 55, 285
fining-upward cycle, fining-upward packet, 59
megacycle, 55, 59, 288
meter-scale cycles, 55, 59, 61
Milankovitch Cycles , 64, 288
shoaling-upward cycles, 53
storm cycles, 55, 56, 292
Cycloconcha. See Mollusca: Pelecypoda
Cyclocrinites, See: Algae : Chlorophyta : Dasycladaceae
Cyclocystoidea. See Echinodermata
Cyclonema. See Mollusca: Gastropoda
Cyclora. See Mollusca: Gastropoda
Cyclostomata. See Ectoprocta
Cylindrocoelia. See Algae
Cymaronora. See Mollusca: Pelecypoda
Cyntluana Formation, 123
Cyrroceras . See Mollusca: Cephalopoda
Cyrtodontula. See Mollusca: Pelecypoda
Cyrtolites. See Mollusca: Monoplacophora
Cystaster. See Echinodermata: Edrioasteroidea( A s t o i d c a . Sec Echinodermata
cystoids. See Echinodermata
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crabs. See Arthropoda: Crustacea: Malacostraca
crateriformis, Lichenocrinus. See Echinodermata: Crinoidea:
Disparida
Dalmanella. See Brachiopoda: Articulata
Dalve, Elizabeth A., 48, 230, 238, 263
damage (in life). See injuries
desiderata, Astraspis. See Chordata: Vertebrata: Agnatha
detrended correspondence analysis (DCA). See techniques
Devonian, xix, 4, 9, 62, 72, 79, 144, 151, 186, 191, 239, 251
diagnosis . See taxonomy: Linnaean HierarchyDickhaut, H. E., 263
dickhauti, Lepidolites. S e e A l g a e : C h l o r o p b y t a :
Dasycladaceae
Diekmeyer, Sharon C. St. Louis, 232, 307
Diestoceras. See M o l l u s c a : C e p h a l o p o d a
Dill Robert F, 54
Dillsboro Formation, 44, 51
dimorphism, sexual, 153, 283, 290dinoflagellates. See Algae: Pyrrophyta
"dioramas," pl. 13
diplobathrid camerates. See Echinodermata: Crinoidea:
Camerata
Diplocraterion. See taphonomy: trace fossils
Diplopoda. See Arthropoda
disarticulation of hard parts. See taphonomy
Disparida. See Echinodermata: Crinoideadivaricans, Streptelasma. See Cnidaria: Anthozoa: Rugosa
diversity, species diversity, taxonomic diversity, 2, 3, 4, 57, 61,
69, 103, 113, 115, 118, 120, 130, 131, 137, 147,
161 163 167 204 232 234 239 241 283
Dystactophycus. See incertae sedis
e a r t h q u a k e s , 7 , 6 1 , 2 9 1 . See also s e i s m i t e s
earwigs. See Arthropoda: Insecta
eatoni, Triarthrus. See Arthropoda: Trilobita
eccentr icity (in Earth's orbit). See cycles: Milankovitch Cy
ecdysis, 6,92,93, 117, 147, 148
Echinodermata, 24, 26, 144, 145, 166-192,219,223, 233
2 3 7 , 2 4 1 , 2 4 5 , 2 4 6 , 2 6 0 , 2 6 9
As te ro id ea , 24 , 167, 168, 169, 182,183, 184, 185, 186,
189-190, 208, 210, 235, 241, 247, 248, 259,
265, 266, 274 Asteriacites (See taphonomy: trace fossils)
Fromia: F. nodosa, pl. 9Goniasteridae, 189
Hudsonaster, 189 ; H. s imp/ex, 185, 274
Lanthanaster, 189; L. intermedius, 184Ophidiasteridae, 189
Palaeaster: P. simplex (See Hudsonaster simplex)
Palaeasterina: P. approximata, 259, 26 5; P. speciosa260 ,266
Petraster: P. speciosus, 184, 189Promopalaeast er , 189, 235
P dyeri 184
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161, 163 ,167 , 204, 232, 234, 239, 241, 283
d'Orbigny, Alcide, 85, 90,95, 271
dolomite, 3, 53, 217, 221, 222, 283
P. dyeri, 184P. magnificus, 182, 183P. speciosus, 182, 183
Cincinnaticrinus, 113, 178, 182, 186, 226, 232,240,
241,253
C. pentagonus, 173C. varihrachialus, 172, 173
Dystactocrinus: D. constrictus, 173, 183 Ectenocrinus, 113, 178, 226, 232,240, 241, 253
E. geniculatus, 173 E. simplex, 172, 173, PL 10
locr inus, 178 , 224/. oehanus, 271/. subcrassus, 110, 142, 170, 172, 174, PL 10
Lichenocrinus, 178L. crateriformis, 172L. dubius, 186L. milleri, 172
Metacrinus, 182 Neocrinus: N. decorus, pl. 9Pontiometra: P. andersoni, PL 9
Cyclocystoidea, 169, 190-191, 247
7.ygocycloides: Z. magnus, 185cystoids, 237. See Rhombifera
Echinoidea, 167, 238, 248
sand dollars, 167
facultative, 285, 288
inqui l ine , inqui l inism, 286. See also incolent
mutualism, 234, 236, 288, 290, 292
obligate, 123, 285, 288
parasite, parasitism, 86, 94, 123, 182, 223, 229, 234, 236,
243, 246, 280, 281, 282, 285, 286, 288, 289,
290, 292
predation, 7,92,93, 95, 123, 124, 125, 131, 132, 137, 143,
147, 151, 153, 154, 155, 161, 162, 182, 183, 186,
189,190, 205, 210, 229, 234, 235-236, 241, 282
283 , 284 , 285 , 289 , 290 , 291 , 293 . See alsoboring
durophagy, 131, 284
symbiosis, 210, 235, 282, 284, 288, 289, 290, 292
endosymbiont, 210, 284
ecologic succession, 56, 57, 115, 281, 292
ecology, ecologic needs. See also chapters on individual
groups of ani mal s; ecologic associations;
environment
autecology, 229, 280, 284, 292
ecosystems, x, 1, 67, 143, 222, 233 -24 8, 284
communities, 4, 5,56, 57, 169, 182,218, 230, 231, 237,
241
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sea urchins, 167, 168, 169, 237
Strongylocentrotus: S. franciscanus, PL 9Edrioasteroidea 56 60 114 169 182 186 188 189 190
consumers, 234, 282, 285, 290
food chain, food web, 67, 68, 234, 282, 285, 290
ild 237 239 247 248 286
M . mam mula ta , 84 , 85 , 90 , 271Parvohallopora, 111, 252, 265, 270
P. onealli, 271P . ramosa, 84, 85 , 90 , 271P. rugosa (now included in P. ramosa), 265, 270
Spatiopora, 87 , 88 , 94Eden Park, Cincinnati, Ohio, 29, 263
Eden Park Reservoir, 29
Eden Shale, Eden Formation, Eden Group, 47, 48, 51, 52,
296, 301. See also Kope Formation; Middle
(Eden) Shales
edenense, Veryhachium. See acritarchs
Edenian Age, Edenian Stage, 11, 44, 46, 47, 51, 55, 63, 67,
81, 130, 138, 139, 143, 144, 151, 155, 203, 208,
209, 220, 233, 284, 291, 302, 311, 320
Edrioasteroidea. See Echinodermata
elegans, Rhodesognathus. See conodonts
Elias, Robert J., 81, 206, 235, 316, 321
Elkhorn Formation, 44, 56, 63, 68, 73, 75, 79, 158, 162, 172,
178, 186,214
emacerata, Dalmanella. See Brachiopoda: Art iculata
Embree, Jesse, 264
encrustation, 56, 60, 64, 72, 74, 77, 79, 87, 88, 89, 90, 93, 94,
eustatic sea-level change. See cycles, cyclicity
event beds, event horizons. 59-61. 224. 225, 285
evolution. See organic evolution
evolutionary faunas, sens!/ Sepkoski, 2, 237; "Modern
Fauna," 2. See also Cambrian , "Cambrian
Fauna"; Paleozoic, "Paleozoic Fauna"
Excello Member. See Brookville Formation
excrement. See feces
exoskeleton, 6, 92, 93, 117, 119, 147, 148, 151, 155, 161,2
285
extinction: mass extinction, 2, 240, 287
Faber, Charles, 14, 24, 27-28, 32, 144, 172, 264, 273, 27
275, 276
faberi, Technophorus. See Mollusca: Rostroconchia
Faberia. See Algae
facies. See also environment
biofacies, 59-61,281, 285, 287
lithofacies, 59,281,285, 287
factor analysis. See techniques
facultative association. See ecologic associations
fair-weather wave-base. See wave base: normal wave-bas
Fairmount Member. See Fairview Formation
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105, 107, 110, 114, 115, 120,122, 123, 128,137,
139, 144, 178, 182, 196, 219, 223, 224, 231, 232,
236 237 240 242 243 244 245 251 286 See
Fairview Formation, 12, 21, 48, 51, 53, 55, 56, 58, 59, 60
63, 68, 87, 106, 110, 120, 123, 127, 173, 175
180 196 214 224 225 232 251 252 pl. 5
florida, Constellaria. See Ectoprocta: Cvstoporata
Flower, Rousseau H., 43, 133, 134, 138, 295
floweri, Fascifodina. See taphonomy: trace fossils
floweri, Palaeoscia. See Cnidaria: Hydrozoa: Siphonophor-
ida: Porpitidae; taphonomy: trace fossils
FMNH. See Field Mu se um of Natural History
fodiniclmia. See taphonomy: trace fossils: behavior
categories
Foerste, August F., 18, 29,48, 77,78, 109, 149, 150, 151,153,
162, 207, 229, 264, 270, 271
Foerstephyllum. See Cnidaria: Anthozoa: Tabulata
food supply. See foodfood chain, food web. See ecology: ecosystems
food, 4,67,68,71, 74, 85, 87,92, 101, 123, 151, 155, 160, 168,
170, 172, 180, 188, 191-192, 198, 211, 229, 234,
235, 237-238, 239, 253, 280, 282, 283, 285, 286,
287 , 290 . See also chapters on individual
groups of an ima ls ; ecolo gic associations: co m-
mensalism; ecologic associations: predation;
ecology: ecosystemscoprophagy, 121, 123,282
deposit feeding, 123, 131, 140, 163, 190, 204, 205,211,224,
283, 285, 292
fil f d fil f di 71 72 87 101 127 131 144
Caurocrinus. See Echinodermata : Cr inoidea : Camerata
Gee, Henry, 191
gener ic name. See taxonomy: nomenc lature : Binomia l Sys
tem of Nomenclature
genkulatus, Ectenocrinus. See Echinodermata: Crinoidea:
Disparida
Ceniculograptus. See Hemichordata: Graptoloidea
genus, genera. See taxonomy: I.innaean Hierarchy
geologic t ime-scale, xix
geologic t ime-units , 45-46
Ge olo gic al So cie ty of Am er ic a, 5, 11, 30, 32, 36, 54, 59, 133
257, 271,274Penrose Medal, 30, 32
geopetal structures, 124
George Washington University (formerly Columbian Unive
sity), 22, 35
Gibson, Bruce, 105, 176
Gibson, Charlotte, 105, 176
gigantea, Anomalodonta. See Mollusca: Pelecypoda
gigas, Isotelus. See Arthropoda: TrilobitaGilbert Formation, 214
Girvanella. See Cyanobacteria
Clyptocrinus. See Echinodermata: Crinoidea: Camerata
Glyptorthis See B hi d A i l
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filter feeder, filter feeding, 71, 72, 87, 101, 127, 131, 144,
155, 168, 172, 188,283,285,292
scavenging, 7, 151, 205, 223, 224, 225
Glyptorthis. See Brachiopoda: Articulata
Goldman, Daniel, 195
Gondwana . See paleogeography
111. 120, 12", 134, 137, 142, 145, 155, 157, 172,
173, 175, 176, 180, 183, 195, 207, 210, 229, 262,
2 6 5 - 2 6 6 , 2 7 2 , 2 7 6 , 2 7 7
Hall , John W., Jr ., 259 ,26 5,2 661 1 . i l l .m i . A n t h o n y 3 , 4
halli, Alloliehas. See Arthropoda: Trilobita: Lichida
halli, Amphilichas. See Arthropoda: Trilobita: Lichida: Alloli- chas halli
Hallopora. See Ectop rocta: Trepostomata: Parvohallopora1 lamilton Comity, Ohio, 10, 89,90, 106, 142, 151, 155, 156,
173, 180, 186, 277, pl. 10
'Trammel Eossil Park, Sharonville, 12, 124, 257I lammond, W. E., 266
Hand, Greg, 306
hardgrounds, 60,64, 105, 124, 125, 178, 188, 203, 206, 211,
219,250, 251,286
I larding Sandstone, 199
Harper, George W., 17, 18, 26, 27, 32, 33-34
Harris, Frank W., 56,57, 114, 115, 231
Harris, G. D., 25Harris, I., 259, 266
I larvard University, xv, 18, 23, 32, 260, 273, 274
Mu se um of Comp ara tive /ool ogy (MC Z) , xv, 18, 23, 32,
2 6 0 , 2 7 3 , 2 7 4 , pl . 13
Holbrook, R. H., 267
boles (in shells). See taphonomy: trace fossils: borings
Holland, Steven M., 53, 55, 59,63, 113, 120, 130, 204,
215-226, 229, 231, 232, 233, 241, 255, 312, 12
I lolothuroidca. See Echinodermata
holotype. See taxonomy: type specimens
I loltedahlina. See Brachiopoda: Articulata
homeomorphy. See organic evolution: convergent evolut
horseshoe crabs. See Arthropoda: Chelicerata: Xiphosuri
Hosea, L. M., 24, 267
host, 77, 86, 87, 88, 93,94, 103, 123, 144, 182, 189, 210, 2235, 236, 242, 243, 244, 245, 246, 282, 285,
288, 289
Hovey Museum, Wabash College. See Crawfordsville,
Indiana
Howe, A. J . , 267
I [udson River Croup, 266, 269, 277
I ludsonaster. See Echinodermata: Asteroidea
I luffman Dam. near Davton, ()hio, 153Hughes, Nigel C, 82, 153
humerosum. See M o l l u s c a : G a s t r o p o d a : Cyclonema
humerosum, Cyclonema. See Mollusca: Gastropoda
Hunda, Brenda, 151
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6 0 , 7 3 , 7 4 , p 3
Harvey, J. W„ 260, 266
Haucr, Kendall, 311
Hunda, Brenda, 151
Hunter, W. H. H., 267
huronensis, Labechia. See Porifera
Dearborn County, pl. 3, pl. 5
Franklin County, 105, 107, 157, 184, 196, pl. 6, pl. 10
Brookville, 184, 196
Indiana Geological Survey, 51, 214, 255, 263
Jefferson County , 111
Mad i son , 7 7 , 7 9 , 2 2 1 , 2 6 6 , 2 6 9
Richmond, 47, 120, 265, 270, 272
Wayne County, 67 , 75 , 228
infauna, infaunal, 7, 131, 139, 153, 204, 205, 237, 238, 239,
247, 248, 284, 286
injuries, damage (in life), 137, 183, 186, 235, 236. See also
ecologic associations: predationinquil ine, inquil inism. See ecologic associations
insculpta, Glyptorthis. See Brachiopoda: Art iculata
Insecta. See Arthropoda
intermedins, Lanthanaster. See Echinodermata: Asteroidea
International Code of Zoologi cal Nom encl atu re (ICZ N), xi
International Geological Correlation Programme, 63, 255
International Stratigraphic Code. See stratigraphy
interspecific interactions. See ecologic associationslocrinus. See Echinodermata: Crinoidea: Disparida
Ischadites. See Algae: Chlorophyta: Dasycladaceae
Ischyrodonta. See Mollusca: Pelecypoda
isochronous surfaces See t ime
Campbell County, 59, 109, 203, 209, pl . 5
Cold Spring, 174
Covington, 19, 28, 127, 142, 256, 263, 266, 274
Fleming County, 72
Kenton County, 67, 87, 90, 110, 123, 209
Kentucky Geological Survey, 10, 51, 214, 256
Lexington, 40, 67, 123, 272
Transylva nia Co l lege , Un iv er si ty , 40 , 27 2
Madison County, 77
Marion County, 120
Maysvil le , 11, 59,79,91, 180
Nelson County, 78 Tr imble County , 124, 129
Bedford, 129
kentuckyensis, Rhytiodentalium. See Mollusca: Scaphopoda
Kesling, Robert V., 178
Kiessling, Wolfgang, pl. 2
k ingdom. See taxonomy: Linnaean Hierarchy
Kjellesvig-Waering, Erik N., 158, 159, 162, 300
Knell, Simon J., 265Kope Formation, II, 19, 21, 44, 49, 51, 52, 53, 55, 56, 57, 59,
60,61,67,68,69, 106, 113, 120, 123, 127, 130,
142, 144, 149, 151, 155, 156, 160, 173, 176, 178
180 186 190 196 203 208 209 211 214 219
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isochronous surfaces. See t ime
Isopoda. See Arthropoda: Crustacea
Isorophus. See Echinodermata: Edrioasteroidea
180, 186, 190, 196, 203, 208, 209, 211,214, 219
225, 231, 232, 253, pl. 5. See also Eden Shale;
Latonia Formation
Leptotrypa. See Ectoprocta: Trepostomata
l.esquerenx, Leo, 266, 268
Letton, Ralph. See Letton's Museum
I.etton's Museum, 268Lexington Limestone, 52, 62, 123, 140, 163, 199, 214, 220
Lexington Platform, 216, pl . 12
Lexington, Kentucky, 40, 67, 123, 272
Liberty Eormation, Member, 44, 48, 53, 56, 63, 68, 72, 77,
78, 79, 105, 107, 108, 111, 134, 162, 176, 180,
211 , 214 , 224 , Pi 1 1 . See also B rookv i l l e
Eormation
Lichenocrinus. See Echinodermata: Crinoidea: DisparidaLicrophycus. See taphonomy: trace fossils
l ife assemblage , 8, 230, 283, 287. See also ecology: ecosys
tems: communit ies
life habits. See chapters on indi vidual grou ps of ani mal s
l imestone
grainstone, 50, 55, 59, 114, 286, 289
packstone, 50, 55, 59, 114, 286, 289
wacke sto ne, 50Limper Geologica l Mus eu m See: Miam i University
l impets. See Mollusca: Gastropoda: Archaeogastropoda
Limulus. See Arthropoda: Chelicerata: Xiphosurida
Lindahl, Josua, 33, 268
Malays ia , pl . 3
mammulata, Monticulipora. See Ec to p roc ta : T repos tom
Manitoba, Canada, 153
Manitoulinoceras. See M o l l u s c a : C e p h a l o p o d aMaquoketa Group, 81
Marble Hill bed. See Waynesville Eormation
March and , Pau l, pl . 13
Marion County, Kentucky, 120
marker beds, 59, 60, 287
Martin, Wayne D., 49, 50, 56,99, 114, 115, 218, 256, 298
307, 315
martini, Neostrabops. See Arthropoda, AglaspidaMason County, Kentucky: Maysville, 11, 59, 79, 91, 180
mass ext inct ion. See extinction: mass extinction
Mastigograptus. See Hemichordata: Dendroidea
Mather, William W., 268, 269, 277
Maysville, Mason County, Kentucky, 11, 59, 79, 91, 180
Maysv i l l i an S tage , 11 , 21 ,44 ,46 , 51 , 55 ,63 ,67 ,68 ,72 , 79
80, 81, 89, 109, 123, 125, 130, 139, 144, 145
151, 153, 155, 158, 160, 163, 182, 184, 191, 207 , 208 , 209 , 219 , 220 , 223 , 233 , 287 . See Lorraine Group
McGraw-Hil l Companies, 238
McKinney, Frank K., 94
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J
Lingula. See Brachiopoda: LingulaI .innaean I licrarchy. See taxonomj
McMick en 1 hill, 21. See also University of C inc innat i
McMicken Member . See Latonia Formation
Miamitown Shale, 12,21, 53, 60, 124, 125, 129
Michigan, University of. See University of Michigan
Mickleborough, John, 26, 27, 153, 155, 262, 276, 313
micraulaxum, Multiplicisphaeridium, See: acritarchs
microscopy. See techniques: microscopy
infrastructure, 6, 68,72, 144, 166, 168, 192, 292
Middle (Eden) Shales, 47
Milankov itch Cycles. See cycles, cyclicity
Millbrig K-bentonite bed. See K-bentonite, K-bentonite beds,
K-bentonite "zones"
millepunctata, Trematis. See B r a c h i o p o d a : I n a r t i c u l a t a
Miller, Arnold I., 113, 231, 232, 305, 308, 312
Miller, C. A., 269
Miller, S. A., 14, 17, 23, 24-25, 28, 29,42, 68, 80, 82, 126,
127, 134, 137, 142, 144, 157, 160, 161, 176, 182,
184, 185, 203, 204, 205, 20 6, 20 8, 209 , 210, 259,
260, 261, 265, 266, 267, 270, 272, 274, 275
milleranus, Ceraurus. See Arthr opod a: Trilobita: Phacop ida
milleri, Eichenocrinus. See Echinodermata: Crinoidea:
Disparida
milleri, Technophorus. See Mollusca: Rostroconchia
mill ipede s. See Arthropoda: Diplopoda
Milne-Edwards, Henri, 265, 270
i i t C i S E hi d H l h id
Endoceras, 134Corbyoceras, 138
G. curvatum, 137, 138G. duncanae, 134, 135
lsorthoceras, 137 Manitoulinoceras
M. tenuiseptum, 137M. williamsae, 137
Narthecoceras: N. dunni, 138nautiloid cephalopods, 60, 87, 88, 89,90, 94, 132-139,
205,228, 229, 235, 236, 238, 241, 244, 245, 249,
251,252, 264, pl. 8 (includes
Ac ti no ce ra to id ea, End oce rat oi de a, and
Nautiloidea)
Nautiloidea, 245
As coc er id a, 137
N au ti lu s, 9 5 , 1 1 7 , 1 3 1 , 1 3 2 , 1 3 4 , 1 3 7 , 1 3 9 , 2 4 1 . p l . 4Orthocerida, 137
octopus, 117, 131, 139
Oncoceras, 137O. delicatum, 137
Orthonyhyoceras, 43. See also TreptocerasSchuchertoceras
O b 137 138
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miniata, Cucumaria. See Echinodermata: Holothuroidea
Minnesota Geological Survey, 31
Mi t L i h W 178 310
O.obscurum, 137, 138squid, 117, 118, 131, 137, 139,252
t f il 138 S l t h t f il
platyceratids, 120
Ptcropoda, 145
Salpingostoma: S. richmondensh, 120, 12 Jslugs, sea slugs, 117, 119
Monoplacophora, 93, 119, 120, 125, 139, 223, 244, 247
Archinacella: A. area, 120, 121
Cyrto l i t es , 125, 139C. ornatus, 120, 121
Helcionopsis: H. striata, 120 , 121 Neopilina, 139Sinuites, 225
S. cancellatus, 120, 121Pelecypoda, 7, 92, 93, 98, 99, 100, 101, 112, 113,117, 118,
119, 123,125, 128-131, 139, 144, 157, 168, 189,
206, 210, 223, 225, 231, 235, 237, 238, 240, 241,
242, 244, 245, 251, 269, 276, 280, 281, 283, 284,
291,293
Am bo ny chi a, 88 , 89, 126, 129, 130, 131, 223, 225, 240,241, 252
A. cultrata, 126 Anomalodonta: A. gigantea, 126Caritodens, 128, 130, 223, 241, 251
C. demissa, 126cockles 129
monobathrid camcratcs. Sec Echinodennata: Crinoidea
Camerata
Monoplacophora. See Mollusca
Montgomery County, Ohio, 72, 157
monticule . See Ectoprocta: anatomy and morphology
Monticulipora. See Ectoprocta: Trepostomata
Moore, Richard B., 17, 270, 279
Moore, Tim, 38
moorei, Lepadocystis. See Echinodermata: Rhombifera
moorei, Paupospira. See Mollusca: Castropoda
Morris, Robert W., 123
Morrow, Warren County. Ohio, 175
Mount Auburn Member, Formation, 21, 44, 53, 55, 59,
68, 160,214, 223, 250. See also McMillan
formation
Mount I lope Member. See Fairview Formation
mountain building, orogeny. See tectonic activity
mudstone, 50, 217, 218, 219, 222, 224, 225
Ml 'CM. Sec Miami I University, ( )hio, ( hlnrd
multipartitum, Petalichnus. See taphonomy: trace f o ss i l s
Multiplicisphaeridium. See acritarchs
murexes, murices. See Mollusca: Castropoda:
Neogastropoda
mussels See Mollusca: Pelecypoda
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cockles, 129
Corallidomus: C. scobina, 128, 206Ctenodonta 129 131
mussels. See Mollusca: Pelecypoda
mutua l i sm. See ecologic associations
mvti l ids See Mollusca: Pelecypoda
nicholsoni, Ceramopora. See Ectoprocta: Cystoporata
Nickles, John M., 18,19,20, 27, 30, 32-33, 34,47, 52, 230,
259, 260, 261, 263, 265, 266, 267, 269, 270, 272,
275, 277Nitecki, Matthew H„ 68
nodosa, Fromia. See Echino dermata: Asteroidea
nomen duhium (pl., nomina dubia), 42, 288. See taxonomy:
nomenclature
nomen nudum (pl., nomina nuda), 42, 288. See taxonomy:
nomenclature
nomenclature. See taxonomy
normal wave-base, fair-weather wave-base. See wave baseNorth Atlantic Province. See faunal provinces
North Bend Tongue. See Fairview Formation
numerosum, Trachomatichnus. See taphonomy: trace fossils
nutrit ion. See food
Nyctopora. See Cnidaria: Anthozoa: Tabulata
O'NeallJ. K., 271
Obata, Tadahiro, 43,317obligate association. See ecologic associations
obrution deposits, 60, 288
obscurum, Schuchertoceras. See Mol lusca : Cephalopoda
occidentalis, Hebertella. See Brachiopoda: Articulata
Ohio State University, Columbus, Ohio, xv, 2, 51, 256, 271
Orton GeologicalMuseum (OSU), xv, 67, 128, 134, 229
ohioensis, Megalograptus. See A r t h r o p o d a : C h e l i c e r a t a :
Eurypterida
Oldroyd, John David, 231
Oligocene, xix, 4
Oncoceras. See Mollusca: Cephalopoda
oncolites. See Algae
onealli, Acidaspis. See Arthropoda: Trilobita:
Odontopleurida
onealli, Parvohallopora. See Ectoprocta: Trepostomata
Ophidiasteridae. See Echinodermata: AsteroideaOphiothrix. See Echinodermata: Ophiuroidea
Opisthoptera. See Mollusca: Pelecypoda
Orbigny, Alcide d', 85, 90, 95, 271
orbital parameters. See cycles: Milankovitch Cycles
order. See taxonomy: Linnaean Hierarchy
Ordovician: "Ordovician Biodiversification Event," 2, 289;
"Ordovician Radiation," 1, 2, 289
ordovicicus, Amorphognathus. See conodontsOrdovicium. See acritarchs
Oregonia Formation, Member, 44, 53, 56, 63,214, 223. See
also Arnheim Formation
oregonia, Oulodus. See conodonts
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occidentalis, Hebertella. See Brachiopoda: Articulata
octopus. See Mollusca: Cephalopoda
Odontopleurida odontopleurids See Arthropoda: Trilobita
oregonia, Oulodus. See conodonts
organic evolution, 1, 2, 39, 61, 62, 67, 82, 105, 115, 118, 131,
134 139 163 167 195 196 229 233 237 267
Palaeasterina. See Echinodermata: Asteroidea
Palacocopida. See Arthropoda: Crustacea: Ostracoda
Palaeophycus. See taphonomy: trace fossils
Palaeoscia. See Cnidaria: Hydrozoa: Siphonophorida: Por-pitidae; laphonomv: trace lossils
Palau, I'alau islands, pl. 3, pl. 9 '
paleo bathy metry , 61, 113, 289
Palcoccne, Palaeocene, xix, 4
Paleodictyon. See taphonomy: trace fossils
paleoecology. See ecology
paleoenvironmental interpretation. See environment
paleogcographvBal t i ca , 62 ,215,216
Gondwana, 215, 216
Iapetus Ocean, Sea, 3, 62, 216. 286, 292
Laurentia, 62, 215, 216, 238, /'/. J
paleogeographic maps, 30, 32, pl . 1, pl . 2, pl . 12
Paleotcthys Ocean. 216
Panthalassic Ocean, 216
Queenst on Delta, 216, 220, PL 12Rodinia, 216
Paleontological Research Institution, 75, 87, 158, 159, 173,
186, 203, 209, 257
Paleontological Society, 2, 30, 32, 36, 172, 252, 257, 274
pentagonus, Cincinnaticrinus. See Echinodermata: Cri
noidea: Disparida
pera, Petroxestes. See taphonomy: trace fossils: borings
Pericharax. See Poriferaperiwinkl es. See Mollu sca: Gastropoda: Archaeogastrop
Permian, xix, 4, 74, 77, 121, 145, 182, 186
Petalichnus. See taphonomy: trace fossils
Peter, Mark, 151, 157
Petraster. See Echinodermata: Asteroidea
Petrocrania. See Brachiopoda: luarticulata
Perroxesfes. See taphonomv: trace fossils: borings
Phac opid a, phacop ids. See Arthrop oda: TrilobitaPhaeophyta. See Algae
Phanerozoic Eon, 2,4,46, 62, 239, 284, 289
Philhedra. See Brachiopoda: luarticulata
Pholadomorpha. See Mollusca: Pelecypoda
Phosphannulus, 182. See also byroniidsphosphatization. See taphonomv: phosphatization
Phragmodus. See conodonts
phytogeny, phylogenetic, 198. See also organic evolutionphvlum, phvla. See taxouomv: l.innacan Hierarchy
pilea, Carneyella. See Echinodermata: Edrioasteroidea
pillbug. See Arthropoda: Crustacea: Isopoda
Plaesiomys. See Brachiopoda: Articulata
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Schuchcrt Medal, 32
The Paleontologist, 17, 22
planis piral coili ng. See coi ling of shell
plankton, planktonic, 7, 52, 68, 69, 74, 145, 172, 195, 234
Brachiospongia: B. tuberculata, 72, 73Labechia: L. huronensis, 72 , 73Pattersonia: P. tuberosa, 72 , 73
Pericharax, pl. 3sclerosponges, coralline sponges, 71, 72
Acanthochaetetes: A. wellsi, pl. 3Stromatoporoidea, 68, 72, 128, 206, 220, 221, 233, 242,
243, 245, 247, 292
Aulacera, 72 A . undulata, 72 , 73
Portsmouth, Ohio, 10
Portuguese Man-of-War. See Cnidaria: Hydrozoa:
Siphonophorida
post-mortem transportation, 114, 137
Potter, Paul Edwin, 8, 11, 59, 106
Preachersvil le Member. See Drakes Formation
Preble County, Ohio, 77, 105, 107, 108, 109, 111, 123, 126,
137, 178, 184
Prec ambr ian, xix, 67, 71 , 151
predation. See ecologic associationspredator-prey interactions. See ecologic associations
preservation. See taphonomy
primary producers. See ecology: ecosystems
primary productivity, primary production, 238-239, 282. See
reef. See bioherm
regeneration, 173, 182, 183, 186, 198, 236
regularis, Quadrijugator. See Arthropoda: Crustacea:
Ostracoda
Reinhardt, E., 272
relative age, relative dating, 46
repichnia. See taphonomy: trace fossils: behavior categories
reticularis, Anomaloides. See Algae: C h l o r o p h y t a :
Dasycladaceae
retrorsa, Calymene. See Arthropoda: Trilobita: Flexicalymenretrorsa
Retrorsirostra. See Brachiopoda: Articulatarex, lsotelus. See Arth ropoda: Trilobita
Rbahdopleura. See Hemichordata: Pterobrancbia
Rhodes, F. H. T, 196
Rhodesognathus. See conodonts
Rhodophyta. See Algae
Rhombifera, rhombiferan "cystoids." See Echinodermata
Rhynchotrema. See Brachiopoda: Articulata
Rhytimya. See Mollusca: PelecypodaRhytiodentalium. See Mollusca: Scaphopoda
Richards, R. Peter, 102
Richardson, J. M., 273
richardsoni, Clyptocrinus. See Echinodermata: Crinoidea:
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also ecology: ecosystems
Primaspis. See Arthropoda: Trilobita: Odontopleurida
, ypCamerata
Richmond Group Richmondian Stage 40 44 46 47 48
russelli, 7,itteIloceras. See M o l l u s c a : C e p h a l o p o d a
Sacco, Wil l iam K., pl . 3
salinity. See environmentSalpingostoma. See Mollusca: Gastropoda
Salteraster. See Echinodermata: Asteroidea
Saluda Formation, Member, 44, 48, 49, 53, 79, 81, 138, 162,
214 , 221 . See also W h i t e w a t e r F o r m a t i o n
Sanctum. See taphonomy: trace fossils: borings
sand dollars. See Fchinodermata: Echinoidea
Sanner, JoAnn, 19, 21
Sansom, Ivan )., 199scabiosa, Petrocrania. See B r a c h i o p o d a : I n a r t i c u l a t a
Scaphopoda. See Mollusca
scavenging. See food
Schafer, Wilhelm, 204
Schizomania. See B r a c h i o p o d a : I n a r t i c u l a t a
Schlemmer, Charles, 273, 275
Schmidt, David A., 186, 303
Schuchert, Charles, 20, 28, 30, 31-32, 33, 35, 184, 264, 273,274, 275
schuchertana, Catazyga. See Brachiopoda: Articulata
Schuchertoceras. See Mollusca: Cephalopoda
Schumacher, Gregory A., 44, 178, 312, 318
serpulid worms. See Annelida: Polychaeta
sexual dimorphism. See d imorphism, sexual
Seychelles, Indian Ocean, pl . 9
shalc-to-lhnestone ratio, 232. See also elastic ratioShaler, Nathaniel Southgate, 23, 273-274
shedding. See ecdysis
shell coiling. See coil ing of shell
shell pavements, shingled beds, 59, 60, 99, 110, 114, 251
also B r a c h i o p o d a : A r t i c u l a t a : Rafinesquina
Shideler, William Henry, 28, 262, 263, 265, 268, 274
shideleri, Megalograptus. See A r t h r o p o d a : C h e l i c e r a t a :
Fury pteridaShimizu, Saburo, 43
shingled beds. See shell pavements
Shirle y, J . , 150
Shrake, Douglas L., 92,93, 317
shrimps. See Arthropoda: Crustacea: Malacostraca
Sigma Gamma Fpsilon, 91
siliciclastics, 3, 220, 291
silicification. See taphonomysiliquaria, Lockeia. See Mollusca: Pelecypoda
Silurian, x ix, 2 ,4 ,9 , 35, 56,72 ,79 ,16 1,1 95, 217, 237, 240
simplex, Ectenocrinus. See Echinodermata: Crinoidea:
Disparida
l H d S
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Schweizcrbarfschc Science Publishers, 67
scientific name. See taxonomy: nomenclature: species name
simplex, Hudsonaster. See Echinodermata: Asteroidea
simplex, Palaeaster. See E chinoder mata: Asteroidea: Hu
Sprinkle, James, 168,185,191, 318
squid. See Mollusca: Cephalopoda
Stanley, George D., 212
starfish. See Echinodermata: Asteroidea
"stateline boundaries," "stateline stratigraphic divisions," 51,
214
Station Hollow Member. See Brookville Formation
Stearn, Colin W., 3, 301
Steinkern. See taphonomy: molds: internal molds
stellata, Cyathophylloides. See C n i d a r i a : A n t h o z o a : R u g o s a
stellatus, Cystaster. See E c h i n o d e r m a t a : E d r i o a s t e r o i d e a
stelliforme, Asteriacites. See taphonomy: trace fossils
sterlingensis, Tentaculites. See Tentaculitoid eaStevens, W. J., 274
Stingy Creek Formation, 214
Stokes, William Fee, 1
Stonelick Creek, Clermont County, Ohio, II, 254
storm wave-base. See wave base
storms, 7, 56,57, 59,60,63,64, 77, 81,92, 113, 114, 115, 151,
169, 178, 180, 210, 215, 217, 218, 219, 220, 221,223, 224, 225, 251, 253, 285, 292, 293
hurricanes, 56, 217, 219, 222, 223, 225, 253
storm cycle. See cycles, cyclicity
storm wave-base, 56, 221, 240. See also wave base
synecology. See ecology
synonym, synonymy. See taxonomy: nomenclature
syntypes. See taxonomy: type specimens: cotypes
Systema Naturae. See taxonomy: Linnaeus
systemat ics. See taxonomy
Tabulata . See Cnidaria: Anthozoa
Taconian . See Taconic Orogeny
laconic Orogeny, Mountains, 3, 5, 8, 62, 216, 220, 233,
292-293 , pl. 12
Taeniaster. See Echinodermata: Ophiuroidea
tangent ial thin-sect ions. See techniques: thin-sect ions
Tanners Creek Formati on, 44
Ta pani la, Leif, 210
taphonomy, 1,4-8, 27,46, 52, 56-57, 60, 79,92, 101, 102,
118, 119, 120, 125, 127, 129-130, 132, 133,
134-135, 137, 147, 148, 151, 153, 155, 163, 169,
1 8 8 , 2 2 4 , 2 3 1 , 2 8 2 , 2 8 9 , 2 9 0 , 2 9 3 . See also c h a p
ters on indiv idual gr oups of ani ma ls ; death
assemblage
alignment, 77, 142, 145, 178, 196, pl. 8, pl. 10
b i o i m m u r a t i o n , 2 0 9 , 2 3 6 , 2 4 2 , 2 4 3 , 2 4 4 , 2 8 1 , 3 2 2 . See also
Catellocaulabody fossils, 6, 143, 203, 204, 212, 281, 293
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stratigraphic code. See stratigraphy: Co de of Strat igra phic
N l
y , , , , , , ,
casts, 6, 210, 282, 288, 292
d f i ( ) 8 130
153, 155, 189, 190, 203-212, 217, 218, 221, 222,
223, 224, 225, 226, 235, 238, 293
Chondrites, 56, 205, 211, 224, 226
Chondrites type-A, 205, 208Chondrites type-B, 205, 208
Chondrites type-C, 205, 208
Cruziana, 202, 203,211
Diplocraterion, 60 , 206 , 211 , 224 , 226 , 247
D. biclavata, 2 0 6 , 2 0 9
D. cincinnatiensis, 206 ,208 , 209
D . luni formis, 2 0 6
Dystactophycus. See incertae sedisepircliefs, 204, 284
Fascifodina: F. floweri, 138fucoids, 203, 211, 285
holes (in shells). See borings
hyporel iefs, 204,207,212, 286
ichnofacies, 210-211,286
ichnogenus, ichnogenera, 204, 211
ichnospecies, 204, 206,210,211Licrophycus: L. flahellum, 261l.ockeia: L. siliquaria, 209, 210. See also Mollusca:
Pelecypoda
Palaeophycus, 204, 207, 224, 226P l i P fl S l
subspecies, 39, 285, 292, 294
taxon (pl., taxa), xi, 38, 39, 42, 43, 293
type-species, 40, 42, 43
va ri et y. See subspeciesLinnaeus, Karl, 38; Systema Naturae, 38
nomenclature
Bino mial Sys tem of Nome ncla ture , 38, 204
binomen. See species name
generic name, 37, 41, 42, 43, 286, 291
scientific name. See species name
species name, xi, 37, 41,43, 198, 203, 281, 286,
specific name, 37, 39, 43, 286, 291, 293trivial name. See specific name
der iv a t ion o l nanus , 40
Internatio nal Cod e of Zoologica l Nome ncla ture
( ICZN),x i
priority, 43, 290
pronun ciatio n of scientific nam es , 41
synonym, synonymy, 151; junior synonym, 153, 1
See also prioritytype specimens, 22, 25,42, 158, 260, 266, 272, 274,
cotypes, 42, 283, 286, 289, 292
holotype, 42, 72, 158, 173, 178, 182, 184, 186, 209
286, 289
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Palaeoscia: P. floweri, 209, 212. See also Cnidar ia :
Hydrozoa: Siphonophorida: Porpitidae
paratype, 42, 158, 159, 186, 283, 286, 289
syntypes See cotypes
Thompson, Es ther H., 120
ticks. See Arthropoda: Chelicerata: Arachnida
tiering. See ecology: ecosystems
time. See also geologic time-units
absolute age, absolute dating, 46, 61, 62, 63, 279, 290
radiometric dat ing, 1,46,62,63, 279, 290
isochronous surfaces, 46
relative age, relative dating, 45, 46, 61, 62, 195, 281, 290.
See also biotal succession
superposition, 46
time-averaged accum ulat ions , 7, 230, 231, 234, 293
time-environment diagram, 240time-scale, geologic. See geologic time-scale
time-stratigraphic units, 46
To bin, Ri ck C, 4 4 , 5 3 , 54, 55, 56 , 57, 312
Tow ns en d, M . ) . , 212, 297
trace fossils. See taphonomy
Trachomatichnus. See taphonomy: trace fossils
Trammel , R. L. , 12
transverse thin-sections. See techniques: thin-sections Tr an sy lv an ia Col lege , Un iversi ty . See Kentucky: Lexington
Treatise on Invertebrate Paleontology, xi , 35, 144, 145, 255,297, 300, 301, 304, 307, 308, 319, 322
Trematis. See Brachiopoda: luarticulata
undatus, Phragmodus. See conodonts
undulata, Aulaeera. See Porifera: S t r o m a t o p o r o i d e a
United States Depart ment of Agricu lture , 22
United States Geological Survey, 20, 22, 29, 30, 32, 33, 35,
51, 261,266
United States National Museum, 22, 30, 32, 35, 260, 261. Se
also National Museum of National History;
Smithsonian Institution
University of Chicago, 22, 25, 28, 262
Wa lk er Museum, 2^
Universit y of Ci nc in na ti , 14, 16, 18, 20, 21, 22, 25, 26, 27, 28
29, 32, 33, 34, 35,48, 80,90,93, 110, 138, 142,151, 155, 156, 170, 180, 203, 209, 225, 256, 260
261 , 263 , 264 , 272 , 275 , 298 . See also M c -Micken Hall
University of Mi ch ig an , xiii, 178, 273
Utica Group, Utica Shale, 47,48, 160
vacuum, Foerstephyllum. See Cnidaria: Anthozoa: Tabulata
Va il , Peter R., 4, 320 Va l l and igham, Ge or ge L. , 259, 275
vallandighami, Cyrtoceras. See Mol lusca: Cephalopoda
vallata, Catellocaula. See Catellocaula Va n Cleve , J . W. , 275
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Tr ep os to ma ta . See Ectoprocta
Treptoceras See Mollusca: Cephalopoda: Actinoceratoidea
Vanuxemia. See Mollusca, Pelecypoda
varihrachialus Cincinnaticrinus See E c h i n o d e r m a t a : Cri
Wavncsvi l lc Format ion, 48, 53, 56, 60, 63, 75, 79, 87, 107,
111, 115. 120, 124, 126, 128, 129, 130, 136, 138,
142, 149, 150, 151, 153, 157, 160, 162, 172, 176,
180, 184, 196,2J4, 225, 229, 307, 314, 317, pl . 6, pl. 10. See also Brookville Formation: Waj nes-
x ilk- Member
Blanchester Member, 44
Clarksvil le Member, 44
Fort Ancient Member, 44
Marble Hill bed, 124, 125, 127, 129
"Treptoceras duseri shale ," 60, 136, 137, 138
Waynesvi l le , Ohio, 120, 182, 184, 185, 256, 259, 266. See Warren County, Ohio
waynesvillensis. Vanuxemia. See Mollusca. Pelecypoda
We aver, T h o m a s , 75 , PI S
Weedon , M. J . , 82, 309
Weichold , Char l es , 93, 275
"Weichold doughnut ." See Ectoprocta
"Weichold ring." See Ectoprocta: "Weichold doughnut"
Welch, James R., 182 Welch. I. B . 276
welchi, Megalograptus. See Arthropoda: Chelicerat a:
Furypterida
wellsi, Acanthochaetetes. See Porifcra: sclcrosponges, coral
l
"Lower Member," 44, 2/4
"Upper Member," 44, 214
Saluda Member. See Saluda Formation, Member
Whitf ie ld , R. P., 134, 137, 142, 145, 229, 276-277 Whit t ington , Ha rr y B. , 160
Whitt lesey, Char l e s , 269, 277
Wi l le t , 268, 277
\\ illiains. I Icnn Shaler. 23
williamsae, Manitoulinoceras. See Mollusca: Cephalopo
williamsae, Megalograptus. See Arthropoda: Chelicerata
Furypterida
williamsae, Zittelloceras. See Mollusca: Cephalopoda Wi lm ing ton , Ohio, 260, 273, 276
Wi lson , Mark A ., 124, 209, 210, 281, 296, 298, 311, 315
Wi l son , W i l l i am W. , 277
wilsoni. Polygnathus, 277 Woodward High School, C inc inna t i , Ohio, 18, 26, 32, 3
Woodward, Hen ry, 26
worms and "wo rm s, " 5, 6, 77, 86, 103,142-145, 152, 153,
182, 196, 206, 210, 222, 224, 241, 242, 244,282. See also Annelida
Worthen , A . H. , 47, 126, 137, 170, 172, 174, 266, 268, 269
277,311
X i S h d d
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line sponges
Wcsschnan Tongue See Kope Formation
Xenocrinus. See Fchinodcrmata: Crinoidea: Camerata
Xiphosurida See Arthropoda: Chelicerata
ABOUT THE AUTHORS
DAVID L. MEYER i s Professor of Ge ol og y at the Un iversi ty of Ci nc in na ti . His
research interests are chiefly in the field of inverteb rate pal eon tol ogy , and
they extend to studies of l iv i ng and fossi l reefs , pale oe co lo gy , and
tap hon omy.
RICHARD ARNOLD DAVIS is Professor of B i o log y an d G e ol og y a t the Col le ge
of Mount St. Joseph in Cincinnati. His research interests focus on fossiland l iv ing cephalopods, symbioses and similar re lationships in the fossi l
record, and the history o f the geolog ical sc ie nces.
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STEVEN M HOLLAND i P f f G l t th U i it f G i
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