A Sea Without Fish

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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



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

http://indiana.edu/

mailto:[email protected]



http://indiana.edu/

http://indiana.edu/



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!





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



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?



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



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



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



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



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 )



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



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 ,





A SEA WITHOUT FISH





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



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



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



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.



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.



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



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 .



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



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.













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 .



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



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









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



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



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



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



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.



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



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

http://ickles/

http://ickles/

http://ickles/

http://ickles/



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



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,



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





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



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



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 )



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



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



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



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



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



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



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



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



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





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 (



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



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



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



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.



f raught with dif f icult ies Eve n the international ly rec ogn iz ed Cin ci nn at ia n



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



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



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



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



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



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



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


the Society for Sedimen

tary Geology.



scale of cycl ic ity the megacycle, and described it as a "f ining-upward" Figure 4.9. Kope cyclicity



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



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



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



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),



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



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





ALGAE: THE BASE OF



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



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 ) .



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-



PORIFERANS AND CNIDARIANS:



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



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



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



corallum


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





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


colonial corals. A -E .



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-



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)



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



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



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



(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 .





BRYOZOANS: "TWIGS"

AND "BONES"



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-



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


bryozoans. A. Colonies

of the bryozoan genus



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.



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



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




tion of the lower part of

the zoarium of Het-



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


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



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)



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



, 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-



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



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



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 .



BRACHIOPODS: THE OTHER BIVALVES



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



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



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



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



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



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



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













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



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,



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



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



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



MOLLUSCS: HARD, BUT

WITH A SOFT CENTER



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



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



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


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.



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





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


gastropods. A. Cy

clonema varicosum

Hall, CMC IP 51118, Cyn-

thiana Formation, Lexing



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



part filled with sediment,

upper part filled with

calcite crystals). B. Mar

ble Hill Bed, Waynesville

Formation, Trimble Co.,

Kentucky. Scale in mm.











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).



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



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



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



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



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





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



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)



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



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,



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.).





ANNELIDS AND WORM-LIKE FOSSILS

10



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



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



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



ARTHROPODS: TRILOBITES AND

OTHER LEGGED CREATURES 11



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



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



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.



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



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.



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



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,


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



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



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.











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) .

http://www.scotese.com/

http://www.scotese.com/







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



undatus Branson and

Mehl, CMC IP 50705,

Kope Formation, Campbell

Co., Kentucky, x 227.









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,



height about 15 cm. A. by Charles G. Messing, all others by D. L. Meyer.











Plate 14. The Cincinnati an, by John Agn ew , 2007.



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



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



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.



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) .



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



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.



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.



g

Comactinia sp., Carib

bean. Scanning electron

micrograph, x 79.

ECHINODERMS: A WORLD

UNTO THEMSELVES 12



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



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



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,



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



right. Drawing by John

Agnew.


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



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











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;



, 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



Figure 12.12. Edrioaster-

oid Isorophus cincinna-

tiensis (Roemer), recon

structed as in life, with

food grooves open for

feeding. Drawing by John

Agnew.




edrioasteroids A,B. Isorophus cincinna-











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



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



(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



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 ,



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



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



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).







GRAPTOLITES AND CONODONTS:

OUR CLOSEST RELATIVES?

Graptol ites are among the most dist inct ive foss i ls found in Cincinnatian Graptolites



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


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 ,

http://lthough/

http://lthough/



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



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



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


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.



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.



their eve ntu al discover) shoul d not be over loo ked.





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



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.



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



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



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,


Formation, Clermont Co.,

Ohio, x 0.6. Note impres

sions of cephalic margin,

genal spines, pleurae,

pygidial margin, as well

as coxae (medial paired



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





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



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



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



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



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



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



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



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



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



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.



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) ,



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



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-



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



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



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



, 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 .







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



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



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



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



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



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



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



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



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





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



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



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



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



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]



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)



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)


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



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)


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



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



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



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



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



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



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



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



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!



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



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


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).

http://www.es.mq.edu.au/MUCEP/igcp410/

http://wwvv.uky.edu/KGS/

http://www.uky.edu/OtherOrgs/KPS/

http://www.ohiodnr.com/gcosurvcv/

http://www.uc.edu/

http://www.uc.edu/

http://www.ohiodnr.com/gcosurvcv/

http://www.uky.edu/OtherOrgs/KPS/

http://wwvv.uky.edu/KGS/




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

http://www/

http://cincinnati-oh.gov/crc/pages/-5708-/

http://www.sharonville.org/

http://www.sharonville.org/

http://cincinnati-oh.gov/crc/pages/-5708-/

http://www/



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


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

http://paleosoc.org/

http://paleosoc.org/



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,



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).



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.



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



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.



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



(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)



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



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



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



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



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 (



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



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



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



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)



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.



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)



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



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



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



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



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



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



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.



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



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



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



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



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



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



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



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



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



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



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



zo ne (see bioz one) .z

REFERENCES CITED

Ag er , De re k V. 1963. Principles of Paleoecology. New York: McGra w-H ill . Al be rs ta dt , Le ona rd P., Kenneth. R. Wal ke r, and Ron al d P. Zu raw ski . 1974. Patch

Reefs in the Carters Limestone (Middle Ordovician) in Tennessee, and Verti

cal Zonation in Ordovician Reefs. Geological Society of America Bulletin

85(7): 1171-118.

Al dr id ge , Ri ch ar d ]. , an d Mark A. Pur nel l. 1996. The Conodont Controversies.

Trends in Ecology and Evolution 11(11): 463-468. Al dr id ge , Ri ch ar d J., De re k E. G. Bri ggs, M. P. Smith, et al . 1993. The Anatomy of

Conodonts. Philosophical Transactions: Biological Sciences 340: 405-421.

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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



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



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



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



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



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



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



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



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



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



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



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



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



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



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



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



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



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



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:



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



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



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



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



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



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.



STEVEN M HOLLAND i P f f G l t th U i it f G i





A Sea Without Fish

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