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

Indiana University Press

Bloomington C? Indianapolis
This book is a publication of

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2009 by Richard Arnold Davis and David Lachlan Meyer

Except chapter 15 2008 by Steven M. Holland
All rights reserved

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Library of Congress Cataloging-in-Publication Data

Meyer, David L.
A sea without fish: life in the Ordovician sea of the Cincinnati region / Da-
vid L. Meyer and Richard Arnold Davis; with a chapter by Steven M. Holland.
p. cm. (Life of the past)
Includes bibliographical references and index.
ISBN 978-0-253-35198-2 (cloth: alk. paper) 1. PaleontologyOrdovician.
2. FossilsOhioCincinnati Region. I. Davis, R. A. (Richard Arnold), date-
II. Title.
QE726. 2. M49 2008
560'. 17310977178dc22
2008020036

1 2 3 4 5 14 13 12 11 10 09
The w o r l d w i d e f a m e o f t h e fossils a n d r o c k s o f t h e C i n c i n n a t i , O h i o , re-
g i o n g r e w o u t o f the labors o f m y r i a d a m a t e u r fossil c o l l e c t o r s . T h e c u r r e n t
e m b o d i m e n t of those folk is t h e " D r y D r e d g e r s , " a g r o u p f o u n d e d in C i n -
c i n n a t i in 1942 a n d , to this day, d e d i c a t e d to c o l l e c t i n g a n d u n d e r s t a n d i n g
t h o s e fossils.

W e d e d i c a t e this v o l u m e t o t h e " D r y D r e d g e r s " a n d t o t h e h o s t o f fossil

collectors they represent. Vos salukimus!
 CONTENTS

ix PREFACE

xiil ACKNOWLEDGMENTS

xv REPOSITORIES OF FOSSILS ILLUSTRATED IN THIS BOOK

1 Introduction
1 <

2 Science in the Hinterland

15 THE C I N C I N N A T I S C H O O L OF P A L E O N T O L O G Y

3 Naming and Classifying Organisms

37 <

4 Rocks, Fossils, and Time

45 <

5 Algae
67 THE BASE OF THE F O O D C H A I N

6 Poriferans and Cnidarians

71 S P O N G E S , C O R A L S , A N D JELLYFISH

7 Bryozoans
85 "TWIGS" AND "BONES"

8 Brachiopods
99 THE OTHER BIVALVES

VII
 9 Molluscs
117 HARD, BUT WITH A SOFT CENTER

10 Annelids and Worm-Like Fossils

143 <

11 Arthropods
147 TRILOBITES AND OTHER LEGGED CREATURES

12 Echinoderms
167 A WORLD UNTO THEMSELVES

13 Graptolites and Conodonts

195 OUR CLOSEST RELATIVES?

14 Type-Cincinnatian Trace Fossils

203 TRACKS, TRAILS, AND BURROWS

15 Paleogeography and Paleoenvironment

215 BY STEVEN M HOLLAND

16 Life in the Cincinnatian Sea

229 <

Epilogue
249 DIVING IN THE CINCINNATIAN SEA

255 APPENDIX 1. RESOURCES: WHERE TO GO FOR MORE INFORMATION

259 APPENDIX 2. INDIVIDUALS AND INSTITUTIONS

ASSOCIATED WITH THE TYPE-CINCINNATIAN

279 GLOSSARY

295 REFERENCES CITED

323 INDEX

viii Contents
PREFACE

Two principal goals motivated us to write this b o o k . First, k n o w l e d g e of the

E a r t h ' s a n c i e n t history from g e o l o g y p r o v i d e s a p o w e r f u l lesson a b o u t t h e
e v e r - c h a n g i n g n a t u r e o f the p l a n e t , a n d t h e a n c i e n t history o f one's h o m e
region c a n b e particularly m e a n i n g f u l . The p r e s e n t n a t u r e o f t h e l a n d s c a p e
i n the C i n c i n n a t i region (southwestern O h i o , n o r t h e r n K e n t u c k y , a n d s o u t h -
eastern Indiana) is t h e p r o d u c t of its m o s t r e c e n t g e o l o g i c history, the Pleisto-
c e n e i c e A g e , w h e n c o n t i n e n t a l ice s h e e t s r e p e a t e d l y forced their w a y a s far
south a s the O h i o River. A s r e c e n t l y a s 20, 000 years a g o , m u c h o f southwest-
ern O h i o was c o v e r e d with an ice sheet m u c h as Greenland is today. As the
glaciers r e c e d e d , m e l t waters c a r v e d the present valleys a n d left a m a n t l e of
debris that d e t e r m i n e d the t o p o g r a p h y , d r a i n a g e , soils, a n d v e g e t a t i o n o f the
region. A m a g n i f i c e n t lee A g e exhibit a t the C i n c i n n a t i M u s e u m C e n t e r
e n h a n c e s p u b l i c a w a r e n e s s o f the p r o f o u n d e n v i r o n m e n t a l c h a n g e s that took
p l a c e across the region in t h e short t i m e span in w h i c h h u m a n s i n h a b i t e d t h e
ice-free land. Three works also provide a c o n c i s e history of t h e e n v i r o n m e n t a l
c h a n g e s d u r i n g the Ice A g e : Richard H. D u r r e l l ' s A Recycled Landscape
(1977), R i c h a r d A r n o l d Davis's " L a n d Fit for a Q u e e n : T h e G e o l o g y o f C i n -
c i n n a t i " (1981), a n d the r e c e n t l y p u b l i s h e d Natural History of the Cincinnati
Region, by S t a n l e y H e d e e n (2006).
As impressive as the Ice A g e history of t h e r e g i o n is as e v i d e n c e of g e o -
logic a n d c l i m a t i c c h a n g e , t h e story that c a n b e told from the a n c i e n t b e d -
rock u n d e r l y i n g the P l e i s t o c e n e c o v e r e x t e n d s the record o f g l o b a l c h a n g e
into d e e p t i m e . The b e d r o c k e x p o s e d a t t h e s u r f a c e across s o u t h w e s t e r n
O h i o , n o r t h e r n K e n t u c k y , a n d s o u t h e a s t e r n I n d i a n a i s the r e c o r d o f t h e
O r d o v i c i a n sea o f s o m e 450, 0 0 0 , 000 years a g o , o n e o f t h e m o s t e x t e n s i v e
m a r i n e f l o o d i n g intervals o f t h e N o r t h A m e r i c a n c o n t i n e n t d u r i n g E a r t h
history. In stark contrast to the barren ice sheet of the P l e i s t o c e n e , t h e C i n -
cinnati s e a s c a p e o f t h e O r d o v i c i a n w a s water from h o r i z o n t o h o r i z o n n o t
a d e e p o c e a n blue, but p e r h a p s s h a d e s of a q u a m a r i n e like the waters over t h e
present-day s h a l l o w G r e a t B a h a m a B a n k . N o l a n d m a s s e s b r o k e t h e h o r i z o n ,
and no birds crossed the skies. All the a c t i o n was b e n e a t h t h e sea s u r f a c e ,
w h e r e life thrived in a b u n d a n c e . This profusion of life left a fossil record in
the rocks that f o r m e d from the b o t t o m s e d i m e n t s o f t h e C i n c i n n a t i a n sea
that is a m o n g the world's richest treasure troves of the past. For present-day
C i n c i n n a t i a n s , fossils in their b a c k y a r d s are a c o m m o n p l a c e , a n d m a n y n a -
tives grow up not r e a l i z i n g that m o s t of the rest of the world has n o t h i n g to
rival the fossil riches of their h o m e ! We seek to r e c o u n t the history of the
C i n c i n n a t i region in d e e p t i m e , its vastly different e n v i r o n m e n t a n d m a r i n e
life, for the general p u b l i c a n d for a m a t e u r g e o l o g i s t s .
 M a n y l o c a l residents w h o h a v e b e e n fascinated b y the fossils u n d e r f o o t
c o l l e c t e d a n d s t u d i e d t h e m a l m o s t s i n c e the earliest s e t t l e m e n t s o f the e i g h -
t e e n t h a n d n i n e t e e n t h c e n t u r i e s . G e n e r a t i o n s o f geologists and p a l e o n t o l o -
gists from abroad h a v e visited the r e g i o n a n d written of the a b u n d a n t fossils
a n d t h e strata, i n c l u d i n g the p i o n e e r i n g British g e o l o g i s t C h a r l e s Lyell i n
1842. B e c a u s e t h e C i n c i n n a t i r e g i o n has b e e n a f o c u s for g e o l o g i c a l research
by so many scientists over so m a i n years, there exists today a vast a m o u n t of
i n f o r m a t i o n a b o u t the fossils and rocks of the region. This information is
scattered in m a n y s o u r c e s , i n c l u d i n g the latest issues of s o m e of the world's
l e a d i n g i n t e r n a t i o n a l g e o l o g i c a l j o u r n a l s , Internet websites, a n d n u m e r o u s
t y p e s o f p u b l i c a t i o n s , s o m e w i d e l y available, s o m e o b s c u r e . M u c h o f the early
w o r k d e s c r i b i n g n e w s p e c i e s o f C i n c i n n a t i fossils dates t o the s e c o n d half o f
the n i n e t e e n t h c e n t u r y , and is found in p e r i o d i c a l s no longer published, such
as the Cincinnati Quarterly journal of Science, The Paleontologist, and the
Journal of the Cincinnati Society of Natural History. No single library houses
all o f t h e g e o l o g i c a l i n f o r m a t i o n p u b l i s h e d a b o u t t h e C i n c i n n a t i r e g i o n .
M o r e o v e r , m o s t studies d e a l w i t h o n l y a s m a l l fraction of the total fossil rich-
ness of t h e r e g i o n , a n d , m o s t i m p o r t a n t l y for u s , there has never b e e n a
synthesis of the vast r a n g e of fossil diversity a n d its g e o l o g i c a l context. In this
b o o k we p r e s e n t a synthesis that will r e c o n s t r u c t t h e life of the O r d o v i c i a n
sea i n order t o s h o w n o t o n l y w h a t o r g a n i s m s i n h a b i t e d this sea b u t also h o w
t h e y lived a n d interacted w i t h e a c h o t h e r to c o n s t i t u t e the variety of ecosys-
tems of t h e O r d o v i c i a n sea in the C i n c i n n a t i r e g i o n . The b o o k is not in-
t e n d e d as a t e x t b o o k of g e o l o g y or p a l e o n t o l o g y , but we present sufficient
b a c k g r o u n d i n f o r m a t i o n o n e a c h fossil g r o u p a n d the g e o l o g i c a l c o n t e x t for
readers u n f a m i l i a r w i t h fossils a n d g e o l o g y . W e e x p l a i n w h a t kind o f a n i m a l
e a c h fossil represents a n d h o w it lived a n d interacted with other o r g a n i s m s ,
t h e r e b y d e f i n i n g t h e role of e a c h g r o u p of a n i m a l s in its a n c i e n t e c o s y s t e m .
We h o p e that this a p p r o a c h will b e n e f i t readers with a b a c k g r o u n d in geol-
o g y as w e l l as t h o s e s e e k i n g an i n t r o d u c t i o n to t h e fossils a n d rocks of the
C i n c i n n a t i region.

Conventions I n s c i e n t i f i c p u b l i c a t i o n s , c e r t a i n c o n v e n t i o n s are u s e d t o save t i m e a n d

t r o u b l e . T h e s e are u n d e r s t o o d by the scientists w h o generally write and read
such publications. B e c a u s e this is a scientific work, we h a v e used s o m e of these
c o n v e n t i o n s . H o w e v e r , this b o o k is also i n t e n d e d for the general reader w h o
m i g h t not be familiar with s u c h c o n v e n t i o n s . I Here arc s o m e explanations:

Literature Citations in the Text

F o o t n o t e s o r e n d n o t e s are n o t o r d i n a r i l y u s e d i n scientific p u b l i c a t i o n s .
I n s t e a d , literature c i t a t i o n s arc inserted in t h e text. This c o m m o n l y is d o n e
w h e r e it is a p p r o p r i a t e in t h e c o n t e x t . At o t h e r t i m e s , e s p e c i a l l y in i n s t a n c e s
in w h i c h t h e r e a d e r is b e i n g referred to a n u m b e r of p u b l i c a t i o n s , the litera-
ture citation m a y be at the end of the appropriate sentence or paragraph.
T h o s e e n a m o r e d o f f o o t n o t e s o r e n d n o t e s m i g h t f i n d this p e c u l i a r , but the
idea is for t h e r e a d e r to be referred to o t h e r p u b l i c a t i o n s i m m e d i a t e l y , and

Preface
not h a v e t o s e a r c h a t t h e b o t t o m o f t h e p a g e o r t h e e n d o f t h e c h a p t e r , or,
e v e n , v o l u m e , for t h e p e r t i n e n t r e f e r e n c e .
T h u s , w h e n w e refer y o u t o a p u b l i c a t i o n , t h e literature c i t a t i o n w i l l
b e i n t h e f o l l o w i n g f o r m a t : " ( S . A . M i l l e r 1875). " T h i s m e a n s that y o u are
b e i n g referred to a p u b l i c a t i o n a u t h o r e d by S. A. M i l l e r a n d p u b l i s h e d in
1875; h e n c e , y o u know w h o said w h a t i s b e i n g cited a n d w h e n . I f y o u n e e d
the c o m p l e t e b i b l i o g r a p h i c i n f o r m a t i o n a b o u t that p u b l i c a t i o n , it is p r o -
v i d e d in t h e b i b l i o g r a p h y toward the e n d of the v o l u m e . In c a s e s in w h i c h
it is i m p o r t a n t for y o u to know t h e p a g e n u m b e r w i t h i n that p u b l i c a t i o n
w h e r e the i n f o r m a t i o n o r q u o t a t i o n i s f o u n d , t h e literature c i t a t i o n w i l l b e
in t h e form "(S. A. M i l l e r 1875, 87). "

Names of Organisms and Groups of Organisms

By international a g r e e m e n t of z o o l o g i s t s , the International Code of Zoo-

logical Nomenclature is t h e d o c u m e n t that s p e c i f i e s h o w the n a m e s of
s p e c i e s , g e n e r a , a n d o t h e r g r o u p s o f a n i m a l s are stated a n d u s e d i n s c i e n -
tific works ( I n t e r n a t i o n a l C o m m i s s i o n o n Z o o l o g i c a l N o m e n c l a t u r e 1999).
G e n e r a l r e c o m m e n d a t i o n B i o o f the C o d e e n c o u r a g e s that t h e a u t h o r a n d
date o f every taxon i n the s p e c i e s g r o u p , g e n u s g r o u p , o r f a m i l y g r o u p
m e n t i o n e d in a p u b l i c a t i o n be cited at least o n c e in that p u b l i c a t i o n , a n d
r e c o m m e n d a t i o n 5 1 G e n c o u r a g e s the f u l l c i t a t i o n o f o r i g i n a l a u t h o r s a n d
dates as w e l l as revisers a n d their dates. H o w e v e r , s u c h c i t a t i o n of a u t h o r s ,
dates, revisers, and dates o f revisions d o e s d e t r a c t f r o m t h e f l o w o f t h e
words. B e c a u s e o f t h e i n t e n d e d a u d i e n c e o f this v o l u m e , w e h a v e d e c i d e d
not to do s u c h d e t a i l e d c i t a t i o n s on a r o u t i n e basis, b u t , rather, o n l y w h e n
clarity d e m a n d s it. If you want to k n o w t h e n o m e n c l a t o r i a l history of a
p a r t i c u l a r g r o u p o f o r g a n i s m s , w e r e c o m m e n d that y o u c o n s u l t t h e s c i e n -
tific literature a b o u t the larger g r o u p o f o r g a n i s m s t o w h i c h t h o s e o r g a n -
isms b e l o n g . T h e b i b l i o g r a p h y of this v o l u m e is a g o o d p l a c e to start.
We d e b a t e d at s o m e l e n g t h as to w h e t h e r to g i v e a c o m p l e t e list of all
t h e s u b d i v i s i o n s for e a c h m a j o r g r o u p o f o r g a n i s m s d i s c u s s e d . W e r e c o g -
n i z e that s u c h listings m i g h t b e g e n u i n e l y u s e f u l for t h e r e a l l y s e r i o u s
fossil-collector. H o w e v e r , w e d e c i d e d that, f o r t h e i n t e n d e d a u d i e n c e o f this
v o l u m e , the n u m b e r o f p a g e s n e c e s s a r y w o u l d h a v e m a d e t h e b o o k too
long, and, h e n c e , inordinately expensive. Up-to-date classifications c a n be
f o u n d in t h e f o l l o w i n g r e f e r e n c e s : the m a n y v o l u m e s of t h e Treatise on
Invertebrate Paleontology (a m u l t i - a u t h o r e d , m u l t i - e d i t e d series of v o l u m e s
p u b l i s h e d b y the G e o l o g i c a l S o c i e t y o f A m e r i c a a n d t h e U n i v e r s i t y Press
of Kansas), the t e x t b o o k Fossil Invertebrates ( B o a r d m a n et al. 1987), or Fos-
sils of Ohio ( F e l d m a n n a n d H a c k a t h o r n 1996).

M a n v of the illustrations in this v o l u m e w e r e m a d e specifically for this work: Photographs,

however, s o m e w e r e m a d e by others a n d are used here w i t h p e r m i s s i o n , in Drawings, Maps
s o m e instances, after modification (for e x a m p l e , to r e m o v e labels not pertinent
to the present context). Unless o t h e r w i s e i n d i c a t e d , a given p h o t o g r a p h in this
v o l u m e was prepared e s p e c i a l l y for this work, primarily by o n e of us ( D L M ) .

Preface xi
Technical Terms S c i e n c e is r e p l e t e w i t h t e c h n i c a l t e r m s that d o n o t a p p e a r c o m m o n l y in
a n d the Glossary n o n - s c i e n t i f i c c o n t e x t s . T o m a k e m a t t e r s w o r s e , scientists often u s e c o m -
m o n , e v e r y d a y t e r m s i n w a y s t h a t are n o t their c o m m o n , e v e r y d a y u s a g e s .
T h u s , we felt it i m p o r t a n t to i n c l u d e a glossary; this is f o u n d n e a r t h e e n d
o f t h e v o l u m e . I n t h e interests o f s p a c e , h o w e v e r , w e h a v e not i n c l u d e d
e v e r y t e c h n i c a l t e r m in this b o o k in t h e glossary. For its first u s e , e a c h
t e c h n i c a l term is defined and is in b o l d f a c e t y p e . Those technical terms
that are u s e d in m o r e t h a n o n e c h a p t e r are listed in t h e glossary. A t e c h n i -
cal term that is used in only o n e chapter, such as the n a m e of an anatomi-
c a l f e a t u r e t h a t o c c u r s i n o n l y o n e m a j o r g r o u p o f o r g a n i s m s , i s d e f i n e d the
f i r s t t i m e i t i s u s e d i n t h e v o l u m e ; h o w e v e r , w e h a v e n o t listed s u c h terms
in t h e g l o s s a r y a g a i n , in t h e interests of s p a c e . S u c h w o r d s are listed in
the index to the v o l u m e .
S o w h a t d o y o u d o i f y o u f i n d a t e c h n i c a l t e r m that i s u n f a m i l i a r t o
y o u and the definition is not right there w h e r e you e n c o u n t e r the word?
First, go to t h e glossary. If t h e t e c h n i c a l t e r m is n o t in t h e glossary, or if,
G o d forbid!, t h e c o v e r a g e o f that t e r m i n t h e g l o s s a r y i s i n s u f f i c i e n t , t h e n
g o t o t h e i n d e x a n d t h e n t o t h e text o f t h e b o o k t o w h i c h y o u are referred.
( C o l l e g e professors, l i k e u s , s o m e t i m e s are a c c u s e d o f s t a t i n g t h e o b v i o u s .
G e n e r a l l y , t h i s i s d o n e i n a n a t t e m p t t o a n s w e r t h e q u e s t i o n s o f s o m e stu-
d e n t s in a g i v e n class b e f o r e t h e y are a s k e d . T h e r e is, of c o u r s e , a d a n g e r
o f o f f e n d i n g o t h e r s t u d e n t s i n t h e s a m e class w h o are m o r e a d e p t a t r e c o g -
n i z i n g t h e o b v i o u s . A n d so it is w i t h r e a d e r s as well!)
I n t h e glossary, a n d e l s e w h e r e , w e h a v e i n c l u d e d a d v i c e o n h o w t o
p r o n o u n c e terms. As you know, lexicographers have developed a scheme
o f s y m b o l s t o i n d i c a t e h o w t h e y feel p a r t i c u l a r letters, syllables, a n d w o r d s
s h o u l d b e p r o n o u n c e d . W e h a v e tried t o k e e p t h e u s e o f s u c h s y m b o l s t o a
m i n i m u m . W e h o p e t h a t , i n s o d o i n g , w e still h a v e m a n a g e d t o h e l p y o u
p r o n o u n c e words in a way useful to y o u .

XII Preface
ACKNOWLEDGMENTS

W e are v e r y g r a t e f u l t o t h e f o l l o w i n g c o l l e a g u e s w h o r e a d p r e l i m i n a r y
drafts o f v a r i o u s c h a p t e r s : L o r e n E . B a b c o c k ( T h e O h i o S t a t e U n i v e r s i t y ) ,
R i c h a r d B a m b a c h ( V i r g i n i a P o l y t e c h n i c Institute a n d S t a t e U n i v e r s i t y ) ,
S t e v e n H. Felton ( C i n c i n n a t i ) , R o b e r t J. E l i a s ( U n i v e r s i t y of M a n i t o b a ) , J.
M a r k E r i c k s o n (St. L a w r e n c e U n i v e r s i t y ) , S t e v e n M . H o l l a n d ( U n i v e r s i t y
of Georgia), Steven Leslie (University of Arkansas, Little Rock), James
S p r i n k l e (University o f T e x a s ) , a n d C o l i n S u m r a l l ( U n i v e r s i t y o f T e n n e s -
see). I n p a r t i c u l a r , w e t h a n k Professor H o l l a n d for c o n t r i b u t i n g t h e c h a p t e r
o n the C i n c i n n a t i a n p a l e o e n v i r o n m e n t .
W e t h a n k the f o l l o w i n g c o l l e a g u e s w h o k i n d l y p r o v i d e d illustrations for
our use or p e r m i t t e d us to r e p r o d u c e their illustrations: L o r e n E. B a b c o c k
( T h e O h i o State University), Stig B e r g s t r o m ( T h e O h i o State University), Jon
W . Branstrator ( E a r l h a m C o l l e g e ) , D e v i n B u i c k (University o f C i n c i n n a t i ) ,
G . K e n t C o l b a t h (Cerritos C o l l e g e ) , R o g e r J . C u f f e y ( P e n n s y l v a n i a State
University), Robert J. Elias (University of M a n i t o b a ) , J. M a r k E r i c k s o n (St.
L a w r e n c e University), D a n i e l G o l d m a n (University o f D a y t o n ) , K e v i n G r a c e
(Archives a n d Rare B o o k s Library, University o f C i n c i n n a t i ) , K e n d a l l H a u e r
( M i a m i University), S t e v e n M . H o l l a n d (University o f G e o r g i a ) , W o l f g a n g
Kiessling (Natural History M u s e u m , Berlin), W a y n e M a r t i n ( M i a m i U n i v e r -
sity), C h a r l e s G . M e s s i n g ( N o v a S o u t h e a s t e r n University), M e r r e l l M i l l e r ( B P
A m e r i c a ) , Robert A . P o h o w s k y ( M o r r o w , O h i o ) , John Pojeta, Jr. (U. S . G e o -
l o g i c a l Survey), Paul E . Potter (University o f C i n c i n n a t i ) , W i l l i a m K . S a c c o
(Vale P e a b o d y M u s e u m ) , J o A n n S a n n e r ( S m i t h s o n i a n Institution), C h r i s
S c o t e s e (University o f T e x a s , A r l i n g t o n ) , D o u g l a s L . S h r a k e ( O h i o D i v i s i o n
o f G e o l o g i c a l Survey), James S p r i n k l e (University o f T e x a s , A u s t i n ) , C o l i n
S u m r a l l (University o f T e n n e s s e e ) , R i c k C . T o b i n ( B P A m e r i c a ) , G r e g o r y P .
W a h l m a n ( B P A m e r i c a ) , S t e v e n M . W a r s h a u e r ( D o m i n i o n E x p l o r a t i o n and
P r o d u c t i o n ) , and D a v i d A . W a u g h (Kent State University).
W e t h a n k the f o l l o w i n g publishers and o r g a n i z a t i o n s w h o k i n d l y granted
us p e r m i s s i o n to r e p r o d u c e illustrations from their p u b l i c a t i o n s : American
Midland Naturalist, Annual Reviews, Blackwell Publishing, Cincinnati His-
torical Society, C o l u m b i a University Press, C o n n e c t i c u t A c a d e m y o f A r t s
and S c i e n c e s , E . S c h w e i z e r b a r t ' s c h e S c i e n c e Publishers, G e o l o g i c a l S o c i e t y
of A m e r i c a , journal of Geology, Kentucky G e o l o g i c a l Survey, M c G r a w - H i l l
C o m p a n i e s , M i d - A m e r i c a P a l e o n t o l o g y S o c i e t y , N e w York State M u s e u m ,
O h i o Division o f G e o l o g i c a l Survey, P a l e o n t o l o g i c a l R e s e a r c h Institution,
Paleontological S o c i e t y P e n n s y l v a n i a A c a d e m y o f S c i e n c e , President a n d
Fellows o f Harvard C o l l e g e , S i g m a G a m m a E p s i l o n , S o c i e t y for S e d i m e n -
tary G e o l o g y ( S E P M ) , University o f C h i c a g o Press, University o f C i n c i n n a t i ,
and University o f M i c h i g a n M u s e u m o f P a l e o n t o l o g y .
 W e arc p a r t i c u l a r l y grateful t o John A g n e w o f C i n c i n n a t i w h o painted
" T h e C i n c i n n a t i a n " for t h e c o v e r a n d c o l o r plate, and w h o also did new
d r a w i n g s of a s p o n g e , a s t r o m a t o p o r o i d , a c r i n o i d , a n d an edrioasteroid. T h e
illustrations c o u l d n o t h a v e b e e n c o m p l e t e d w i t h o u t the t e c h n i c a l and artis-
tic skills o f T i m o t h y Phillips ( D e p a r t m e n t o f G e o l o g y , University o f C i n c i n -
nati), E v e l y n M o h a l s k i (formerly o f t h e D e p a r t m e n t o f G e o l o g y , University
o f C i n c i n n a t i ) , a n d Jay Y o c i s ( P h o t o g r a p h i c S e r v i c e s , Universitv o f C i n c i n -
nati). Professor K e v i n a V u l i n e c ( D e p a r t m e n t o f A g r i c u l t u r e and Natural
R e s o u r c e s , D e l a w a r e State University, D o v e r ) kindly p e r m i t t e d us to repro-
d u c e h e r d r a w i n g s that w e r e o r i g i n a l l y m a d e for a n e x h i b i t a t the C i n c i n n a t i
M u s e u m o f N a t u r a l History.
M a n y c o l l e a g u e s and friends a l l o w e d u s t o p h o t o g r a p h s p e c i m e n s i n
their c o l l e c t i o n s : S t e v e B r o w n ( Z a n e s v i l l e , O h i o ) , Fred C o l l i e r (formerly o f
t h e M u s e u m o f C o m p a r a t i v e Z o o l o g y , H a r v a r d University), D a n C o o p e r
(Cincinnati), Steven H. Felton (Cincinnati), Ron Fine (Cincinnati), Bruce
and Charlotte G i b s o n (Cincinnati), Brenda Hunda (Cincinnati M u s e u m
Center), Kendall Haucr ( L i m p e r M u s e u m . M i a m i University), W i l l i a m
H e i m b r o c k ( C i n c i n n a t i ) , M a r k Peter ( C o l u m b u s , O h i o ) , and Janice T h o m p -
son ( N a t i o n a l M u s e u m o f N a t u r a l History, S m i t h s o n i a n Institution).
W i l l i a m B u t c h e r a n d D e n n i s Kytasaari o f t h e N o r t h A m e r i c a n Jules
V e r n e Society provided the accurate translation of and information about
t h e q u o t a t i o n f r o m Jules V e r n e ' s 1864 n o v e l , Voyage au centre de la terre.
A n g e l a G o o d e n ( G e o l o g y - M a t h - P h y s i c s Library, U n i v e r s i t y o f C i n c i n n a t i )
and M a g g i e Heran ( T h e Lloyd Library and M u s e u m , C i n c i n n a t i ) provided
help in finding references.
For m a n y s t i m u l a t i n g d i s c u s s i o n s a n d s u g g e s t i o n s , w e t h a n k Stig Berg-
strom ( T h e O h i o State U n i v e r s i t y ) , D a n i e l B . B l a k e ( U n i v e r s i t y o f Illinois),
Lael Bradshaw (Sinclair C o m m u n i t y C o l l e g e , Dayton, Ohio), Carlton
Brett ( U n i v e r s i t y o f C i n c i n n a t i ) , Billie B r o a d d u s (former h e a d o f the His-
tory o f t h e H e a l t h S c i e n c e s L i b r a r y a n d M u s e u m , U n i v e r s i t y o f C i n c i n -
nati), S t e v e B r o w n ( Z a n e s v i l l e , O h i o ) , t h e late K e n n e t h E. C a s t e r , J. M a r k
E r i c k s o n (St. L a w r e n c e U n i v e r s i t y ) , R o g e r J . Cuffey P e n n s y l v a n i a State-
University), S t e p h e n H. Felton ( C i n c i n n a t i , O h i o ) , Kevin G r a c e (Archives
a n d R a r e B o o k s L i b r a r y , U n i v e r s i t y o f C i n c i n n a t i ) , G r e g Hand ( O f f i c e o f
Public Information, University of C i n c i n n a t i ) , S i m o n J. Knell (University
o f L e i c e s t e r , E n g l a n d ) , G e n e Kritsky ( C o l l e g e o f M o u n t St. Joseph, C i n c i n -
n a t i , O h i o ) , F r a n k K . M c K i n n e y ( A p p a l a c h i a n State University), A r n o l d
M i l l e r ( U n i v e r s i t y o f C i n c i n n a t i ) , T i m M o o r e (University o f H o n g K o n g ) ,
P a u l F . Potter ( U n i v e r s i t y o f C i n c i n n a t i ) , D o l f S e i l a c h e r (Vale University),
a n d t h e late E l l i s Y o c h e l s o n ( U n i t e d States G e o l o g i c a l S u r v e y ) .
f o r their c o n t i n u i n g e n t h u s i a s m f o r this project, and m a n y helpful dis-
c u s s i o n s , we t h a n k R o b e r t J. S l o a n , Editorial D i r e c t o r at I n d i a n a University
Press a n d James O. Farlow, I n d i a n a U n i v e r s i t y - P u r d u e University at Fort
Wayne, a n d editor of t h e L i f e of t h e Past series for I n d i a n a University Press.
For moral support we thank M a r y L. Davis, Kani Meyer, and Univer-
sity o f C i n c i n n a t i g r a d u a t e s t u d e n t s D e v i n Buick, Katherine Bulinski,
Bradley D e l i n e , and Austin Hendy.

xiv Acknowledgments
REPOSITORIES OF FOSSILS
ILLUSTRATED IN THIS BOOK

BMNH N a t u r a l History M u s e u m , L o n d o n

C M C IP C i n c i n n a t i M u s e u m C e n t e r , Invertebrate Paleontology C o l l e c t i o n s

FMNH F i e l d M u s e u m o f N a t u r a l History, C h i c a g o , Illinois

M C Z Harvard University, Museum of Comparative Z o o l o g y

MUGM M i a m i University, 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

OSU 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 State U n i v e r s i t y , C o l u m b u s , O h i o

USNM N a t i o n a l M u s e u m o f N a t u r a l History, S m i t h s o n i a n I n s t i t u t i o n ,
Washington, D. C.

XV
A SEA WITHOUT FISH
Figure 1. 1. 7999 Geologic Time Scale. Reprinted by permission of the Geological Society of America. In
order to read this chart in stratigraphic order, read the columns from bottom to top, starting at the bottom
of the Precambrian column, adding the bottom of the Paleozoic column to the top of the Precambrian col-
umn, the bottom of the Mesozoic column to the top of the Paleozoic column, and the bottom of the Ceno-
zoic column to the top of the Mesozoic column.
INTRODUCTION

The vicinity o f C i n c i n n a t i , i n the O h i o River V a l l e y o f s o u t h w e s t e r n O h i o , Of the many prolific col-

i n c l u d i n g adjacent n o r t h e r n K e n t u c k y a n d s o u t h e a s t e r n I n d i a n a , i s a m o n g lecting grounds in the
the m o s t fossil-rich r e g i o n s i n N o r t h A m e r i c a , i f not t h e entire w o r l d . T h e continental interior, none

profusion of fossils in t h e local l i m e s t o n e a n d s h a l e attracted m a n y p i o n e e r - excels the Ohio river

bluffs at Cincinnati, Ohio.
i n g geologists and p a l e o n t o l o g i s t s o f the n i n e t e e n t h c e n t u r y , a n d m u c h f u n -
Here the Upper Ordovi-
d a m e n t a l work i n A m e r i c a n p a l e o n t o l o g y a n d s t r a t i g r a p h y w a s a c c o m -
cian rocks are almost
plished here. H u n d r e d s of fossil species w e r e first d i s c o v e r e d and n a m e d from
literally made of fossils;
these rocks. Early geologists g a v e the entire series of strata e x p o s e d h e r e t h e many are as perfectly
n a m e " C i n c i n n a t i a n , " a n d this n a m e w a s a p p l i e d t o strata o f s i m i l a r a g e preserved as fossils can
t h r o u g h o u t N o r t h A m e r i c a . C i n c i n n a t i a n fossils are d i s p l a y e d i n m u s e u m s be. The river banks,
all over the world. R e s e a r c h e r s , s t u d e n t s , a n d a m a t e u r fossil c o l l e c t o r s r e g u - road cuts, and even the
larly visit the C i n c i n n a t i r e g i o n t o c o l l e c t fossils. M a n y o f t h o s e w h o h a v e soil in the gardens are

g r o w n up in the region are a w a r e of the a b u n d a n c e of fossils, yet few a p p r e c i - replete with fossils more
common than pebbles.
ate the u n i q u e n e s s of this r i c h n e s s a n d its b r o a d e r s i g n i f i c a n c e to o u r u n d e r -
Almost every museum
s t a n d i n g of the Earth's past. The p u r p o s e of this b o o k is to e x p l o r e t h e rich-
in the world has speci-
ness of C i n c i n n a t i a n fossils a n d t h e stories t h e y tell a b o u t life over 450 m i l l i o n
mens from this locality.
years a g o , w h e n s h a l l o w seas i n u n d a t e d N o r t h A m e r i c a a n d the site o f C i n -
cinnati was in the S o u t h e r n H e m i s p h e r e . W i l l i a m Lee S t o k e s
1960, 1 8 8 - 1 8 9
W h y are fossils so a b u n d a n t in the rocks of C i n c i n n a t i ' s hills? B e y o n d
sheer a b u n d a n c e , w h a t is their s i g n i f i c a n c e for o u r k n o w l e d g e of the history
of life, evolution, and a n c i e n t e n v i r o n m e n t s ? T h e r e is no single a n s w e r to these
questions, but rather several answers c a n be given w h i c h collectively reveal the
significance o f C i n c i n n a t i a n fossils. T h e s e answers c a n b e f o u n d u n d e r four
categories: organic evolution, environment, preservation, and history.

Fossils f o u n d i n C i n c i n n a t i ' s l i m e s t o n e s a n d s h a l e s are t h e r e m a i n s o f a n i - Organic Evolution

m a l s that lived d u r i n g a n interval o f E a r t h history c a l l e d t h e O r d o v i c i a n
P e r i o d . T h e O r d o v i c i a n i s the s e c o n d oldest p e r i o d o f the larger t i m e interval
[The Ordovician radiation]
k n o w n a s the P a l e o z o i c E r a ( F i g u r e 1 . 1). The b e g i n n i n g o f t h e P a l e o z o i c E r a
represents one of the
( m e a n i n g " t i m e of a n c i e n t a n i m a l s " ) is m a r k e d by the oldest rocks c o n t a i n i n g
largest major turnovers
a b u n d a n t fossils o f m u l t i - c e l l e d a n i m a l s ( m e t a z o a n s ) . R a d i o m e t r i c d a t i n g in the history of life and
o f v o l c a n i c ash b e d s i n t e r b e d d e d with t h e s e fossiliferous r o c k s p l a c e s t h e marks the appearance
b e g i n n i n g o f the P a l e o z o i c a t a b o u t 543 m i l l i o n years a g o . S i m i l a r m e t h o d s of groups that came to
date the b e g i n n i n g o f the O r d o v i c i a n Period a t a b o u t 4 9 0 m i l l i o n years a g o dominate marine eco-
and its end at about 443 m i l l i o n years a g o . T h e s p a n of O r d o v i c i a n t i m e rep- systems for the next

resented by the C i n c i n n a t i a n strata a m o u n t e d to less t h a n 10 m i l l i o n y e a r s , 250 million years.

and tell a p p r o x i m a t e l y d u r i n g the latter part of the O r d o v i c i a n , t e r m e d t h e D r o s e r , Fortey, a n d

1, ate O r d o v i c i a n . In the C i n c i n n a t i r e g i o n , a total t h i c k n e s s of o v e r 250 m e - Li 1996, 122
Figure 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
ters (820 feet) of i n t e r b e d d e d l i m e s t o n e a n d s h a l e w a s d e p o s i t e d d u r i n g the
Fauna. From Sepkoski
(1981) and reprinted by L a t e O r d o v i c i a n , c o n s t i t u t i n g t h e C i n c i n n a t i a n a n d c o n t a i n i n g fossils
permission of The Pale- t h r o u g h o u t . F u r t h e r d i s c u s s i o n o f the n a t u r e a n d subdivisions o f C i n c i n n a -
ontological Society. tian r o c k s , a n d e s t i m a t e s o f their a g e , are t h e s u b j e c t o f c h a p t e r 4 .
Professor S t i g M . B e r g s t r o m o f t h e O h i o State University i s a m o n g the
world's m o s t k n o w l e d g e a b l e a n d w i d e l y - t r a v e l e d specialists o n O r d o v i c i a n
fossils a n d stratigraphy. He i n d i c a t e d to us that " t h e r e is n o t h i n g that c a n be
c o m p a r e d e l s e w h e r e in the w o r l d " to t h e d i v e r s i t y of shelly fossils in the
C i n c i n n a t i a n ( B e r g s t r o m , pers. c o m m . ). Metazoan m a r i n e life first b e g a n to
diversify d u r i n g the s o - c a l l e d C a m b r i a n e x p l o s i o n that m a r k e d the onset o f
the P a l e o z o i c , but a c c e l e r a t e d d u r i n g the Cambrian and Ordovician Periods
to r e a c h a p e a k late in t h e O r d o v i c i a n w h e n t h e C i n c i n n a t i a n strata w e r e
d e p o s i t e d . In fact t h e O r d o v i c i a n Period is r e c o g n i z e d as a u n i q u e t i m e of
e v o l u t i o n a r y diversification, t e r m e d t h e O r d o v i c i a n R a d i a t i o n ( D r o s e r e t al.
1996) o r t h e O r d o v i c i a n B i o d i v e r s i f i c a t i o n E v e n t ( W e b b y , Paris, D r o s e r ,
a n d Percival 2004). T h e O r d o v i c i a n m a r k e d a c o n v e r g e n c e o f w h a t Sepkoski
(1981) c a l l e d t h r e e " e v o l u t i o n a r y faunas": metazoan g r o u p s that first a p p e a r e d
d u r i n g t h e C a m b r i a n b u t persisted into the O r d o v i c i a n ( " C a m b r i a n F a u n a " ) ,
g r o u p s that b e g a n t o diversify d u r i n g t h e O r d o v i c i a n ( " P a l e o z o i c F a u n a " ) ,
a n d g r o u p s that first a p p e a r e d in t h e O r d o v i c i a n that diversified after the e n d
of t h e P a l e o z o i c ( " M o d e r n F a u n a " ) ( F i g u r e 1. 2). At t h e e n d of the O r d o v i c i a n
there o c c u r r e d a g l o b a l m a s s e x t i n c t i o n that e l i m i n a t e d s p e c i e s on a large
scale. T h u s t h e C i n c i n n a t i a n t i m e was significant in the history of life as a
G o l d e n A g e of e v o l u t i o n a r y diversification just before a major crisis of mass
e x t i n c t i o n . In m a n y w a y s t h e F a t e O r d o v i c i a n is c o m p a r a b l e to t h e L a t e
C r e t a c e o u s P e r i o d , a n o t h e r G o l d e n A g e p r e c e d i n g a crisis ( F i g u r e 1. 2; Sei-
l a c h e r 1998). F e w if a n y fossil s p e c i e s f o u n d in t h e C i n c i n n a t i a n strata sur-
v i v e d into t h e s u c c e e d i n g S i l u r i a n P e r i o d . C h a p t e r s 5-14 i n t r o d u c e e a c h o f
t h e m a j o r g r o u p s of o r g a n i s m s f o u n d as fossils in t h e C i n c i n n a t i a n .

A Sea without Fish

 The e n v i r o n m e n t o f L a t e O r d o v i c i a n t i m e i n the C i n c i n n a t i r e g i o n c o n t r i b - Environment
uted to the a b u n d a n c e a n d r i c h n e s s of fossils in several f u n d a m e n t a l ways.
C i n c i n n a t i a n fossils a n d rocks b e a r p r o f o u n d t e s t i m o n y t o t h e e x i s t e n c e o f
widespread s h a l l o w seas (called e p i c o n t i n e n t a l o r e p e i r i c seas) o v e r m o s t o f
the N o r t h A m e r i c a n c o n t i n e n t at this t i m e (Plate 1). U s i n g m a n y s o u r c e s of We are accustomed to
e v i d e n c e , g e o l o g i s t s h a v e c o m p i l e d a record of t h e rise a n d fall of sea level thinking of North Amer-
d u r i n g the past h a l f billion years of Earth history ( f i g u r e 1. 3). The L a t e O r - ica as terra firma, one
d o v i c i a n w a s o n e o f the t i m e s o f m a x i m u m rise o f sea level o v e r t h e entire of the large high and dry

g l o b e , rivaled o n l y by the L a t e C r e t a c e o u s ( a c c o r d i n g to t h e r e c o n s t r u c t i o n segments of the earth's

crust, and it is difficult for
by H a l l a m [1984]). The c a u s e of this f l o o d i n g has b e e n attributed to h i g h rates
us to imagine a time in
o f s e a floor s p r e a d i n g w h i c h s w e l l e d the m i d - o c e a n r i d g e s , d i s p l a c i n g i m -
the past when our conti-
m e n s e v o l u m e s o f seawater from the d e e p o c e a n basins o n t o t h e c o n t i n e n t a l
nent was so submerged
plates. The Atlantic O c e a n as we k n o w it did not exist, but instead, a n a r r o w e r beneath the sea that fish
o c e a n c a l l e d the I a p e t u s O c e a n separated N o r t h A m e r i c a f r o m c o n t i n e n t a l could have swum directly
plates later to c o n s t i t u t e E u r o p e a n d A f r i c a (Plate 1). T h e nearest l a n d m a s s e s from the Atlantic Ocean
t o the C i n c i n n a t i region w e r e the rising A p p a l a c h i a n m o u n t a i n c h a i n , a b o u t to the Pacific Ocean,
from Hudson Bay to
300 m i l e s to the east, a n d the l o w - l y i n g C a n a d i a n S h i e l d to t h e n o r t h . Just
the Gulf of Mexico. Yet
before and d u r i n g the Late Ordovician, a p h a s e of m a j o r t e c t o n i c ( m o u n t a i n -
such a time did exist 450
b u i l d i n g ) activity, the T a c o n i c O r o g e n y , resulted i n severe crustal d e f o r m a -
million years ago when
tion and uplift a l o n g t h e r e g i o n b o r d e r i n g N e w York a n d N e w E n g l a n d . Is- the epeiric sea spread
lands w e r e raised h i g h a b o v e sea level as lofty a n d j a g g e d m o u n t a i n c h a i n s from Arctic to Gulf, from
r e s e m b l i n g the m o d e r n A l p s o r H i m a l a y a s . W e a t h e r i n g a n d erosion a t t a c k e d Atlantic to Pacific*
these r a n g e s , a n d rivers c a r r i e d h u g e l o a d s o f fresh water, s e d i m e n t s , a n d
Clark a n d Stearn
nutrients into the s h a l l o w sea.
1960, 6 8
G r e a t v o l u m e s o f s e d i m e n t , c o n s i s t i n g o f c o a r s e g r a v e l s , s a n d s , silts,
a n d m u d s ( t e r m e d s i l i c i c l a s t i c s ) w e r e d e p o s i t e d a s river deltas a n d redis-
tributed b y o c e a n i c c u r r e n t s n e a r t h e c o a s t l i n e i n t h e A p p a l a c h i a n B a s i n .
T h e total t h i c k n e s s o f t h e Late O r d o v i c i a n strata i n t h e A p p a l a c h i a n B a s i n
i n V i r g i n i a r e a c h e s a b o u t 1000 m e t e r s (over 3000 feet) w h e r e a s t h e s a m e
t i m e interval is r e p r e s e n t e d in the C i n c i n n a t i r e g i o n by strata less t h a n 300
m e t e r s (less t h a n 1000 feet) t h i c k ( F i g u r e 1. 4; Kay 1951). O f f s h o r e , o n l y t h e
m u d d y c o m p o n e n t s o f this h e a v y s e d i m e n t i n p u t r e m a i n e d s u s p e n d e d a s
clay p a r t i c l e s , a n d w e r e c a r r i e d b y c u r r e n t s t o r e a c h t h e C i n c i n n a t i a r e a .
These m u d s were thus imports to the region that e v e n t u a l l y lithified
(turned to stone) to form shales. In the C i n c i n n a t i a r e a , s h a l e s are i n t e r b e d -
ded w i t h l i m e s t o n e s , w h i c h are c o m p o s e d o f c a l c a r e o u s shells a n d skele-
tons o f " n a t i v e " m a r i n e i n v e r t e b r a t e s . I n t h e w e s t e r n U n i t e d States a n d
C a n a d a , the Late O r d o v i c i a n c o n t a i n s m o s t l y l i m e s t o n e s s e c o n d a r i l y c o n -
verted t o d o l o m i t e s . T h u s , t h e C i n c i n n a t i r e g i o n r e p r e s e n t s a n i n t e r m e d i -
ate z o n e o f m i x e d s h a l e s a n d l i m e s t o n e s b e t w e e n t h e g r e a t t h i c k n e s s o f
siliciclastics t o t h e east a n d p u r e l i m e s t o n e s f a r t h e r west. B o t h s e d i m e n t s "Of course, true bony fish
i n t e r m i n g l e d i n t h e C i n c i n n a t i r e g i o n , p r o d u c i n g a v a r i e d a n d p a t c h y sea had not yet evolved in Late
floor that was m u d d y in p l a c e s a n d s h e l l y in o t h e r s . S u c h a v a r i e g a t e d bot- Ordovician time, and as we
will see, Cincinnatian
t o m e n v i r o n m e n t offered m o r e p o t e n t i a l t y p e s o f l i v i n g s p a c e s for b o t t o m -
rocks contain no fossil evi-
d w e l l i n g o r g a n i s m s (the b e n t h o s ) , a n d p r o v i d e s a f u r t h e r r e a s o n w h y h i g h
dence of the early, jawless
diversity d e v e l o p e d i n t h e r e g i o n . B e c a u s e t h e r e w a s v e r y little v e g e t a t i o n fish that are known from
o n l a n d d u r i n g the L a t e O r d o v i c i a n , e r o s i o n m a y h a v e c a r r i e d a h e a v i e r the Late Ordovician
load o f dissolved i n o r g a n i c n u t r i e n t s into t h e sea. These nutrients m a y have elsewhere.

Introduction 3
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 was
lower 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 and
extensive over North
America.

a c t e d as a f e r t i l i z e r to s t i m u l a t e t h e p r o d u c t i o n of b e n t h i c b i o m a s s . In ad-
d i t i o n , c l i m a t e , o c e a n o g r a p h i c c o n d i t i o n s , a n d a v a i l a b l e food supply m u s t
h a v e b e e n c r u c i a l t o s u p p o r t prolific m a r i n e life i n t h e C i n c i n n a t i a n sea;
t h e s e factors are e x p l o r e d in d e t a i l in c h a p t e r 15.

Preservation W h e n w e l o o k at r o c k layers as c r o w d e d w i t h w e l l - p r e s e r v e d fossils as t h o s e

o f t h e C i n c i n n a t i a n , w e t e n d t o t h i n k w e arc l o o k i n g a t a c o m p l e t e p i c t u r e
o f life o n t h e O r d o v i c i a n sea f l o o r a s n a p s h o t i n t e r m s o f b o t h the diver-
sity o f s p e c i e s p r e s e n t a n d t h e i r a b u n d a n c e . U n f o r t u n a t e l y , t h e c o r r e s p o n -
d e n c e b e t w e e n t h i s fossil a s s e m b l a g e a n d t h e o r i g i n a l l i v i n g c o m m u n i t y
f r o m w h i c h i t w a s d e r i v e d i s rarely that s i m p l e a n d d i r e c t . T h e fossil r e c o r d
p r o v i d e s a m e r e g l i m p s e of a n c i e n t life, o n e that is h e a v i l y b i a s e d by many
factors. I n o r d e r t o assess t h e i m p a c t o f t h e s e factors o n t h e q u a l i t y o f t h e
fossil s a m p l e , p a l e o n t o l o g i s t s h a v e d e v o t e d a n e n t i r e s u b d i s c i p l i n e , c a l l e d
t a p h o n o m y , to the investigation of processes affecting organic remains

4 A Sea without Fish

 Figure 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.

The organic remains

here are remarkably well
preserved for so ancient
a rock, especially those
occurring in a compact
argillaceous blue lime-
stone, not unlike the lias
of Europe. Its deposition
from death to ultimate fossilization. T a p h o n o m y e m p h a s i z e s the w i d e appears to have gone on

variation i n t h e p r e s e r v a t i o n p o t e n t i a l o f o r g a n i s m s . A n a p p r e c i a t i o n o f t h e very tranquilly, as the Lin-

gula has been met with
significance of variable preservation can be gained by c o n s i d e r i n g aspects
in its natural and erect
o f life, d e a t h , a n d p o s t - m o r t e m history that e n t e r e d into t h e c o m p l e x e q u a -
position, as if enclosed in
tion that d e t e r m i n e d the u l t i m a t e fossil r e c o r d o f t h e O r d o v i c i a n sea.
mud when alive, or still
standing on its peduncle.

Nature of the laving O r g a n i s m C h a r l e s Lyell 1845, 49

B i o l o g i c a l factors a f f e c t i n g p r e s e r v a t i o n p o t e n t i a l include presence of

" h a r d parts, " their chemistry, m i n e r a l o g y , and c o n s t r u c t i o n , and the m o d e
of life of the o r g a n i s m . By far t h e m o s t i m p o r t a n t r e q u i r e m e n t for fossiliza-
tion is possession of m i n e r a l i z e d hard parts s u c h as shells or s k e l e t o n s . S o f t
b o d y parts i n c l u d i n g s k i n , m u s c l e , hair, a n d i n t e r n a l o r g a n s a l m o s t a l w a y s
d e c a y rapidly f o l l o w i n g d e a t h . M a n y c o m m o n m a r i n e i n v e r t e b r a t e s like
w o r m s lack hard parts a l t o g e t h e r o r h a v e o n l y h a r d e n e d j a w s t r u c t u r e s . I n
s o m e m a r i n e e n v i r o n m e n t s , a n i m a l c o m m u n i t i e s are d o m i n a t e d i n n u m -
bers of s p e c i e s or i n d i v i d u a l s by s u c h s o f t - b o d i e d s p e c i e s w i t h little or no
fossilization p o t e n t i a l . O n e o f t h e b e s t - k n o w n e x c e p t i o n s t o t h e d o m i n a n t

Introduction 5
 p r e s e r v a t i o n o f hard parts i s t h e C a m b r i a n B u r g e s s S h a l e o f B r i t i s h C o l u m -
b i a , w i t h its a m a z i n g w e a l t h o f s o f t - b o d i e d w o r m s , a r t h r o p o d s , a n d o t h e r
i n v e r t e b r a t e s , a l o n g w i t h s h e l l - b e a r i n g f o r m s ( C o u l d 1989). I n t h e C i n c i n -
n a t i a n , t h e r e i s v i r t u a l l y n o p r e s e r v a t i o n o f s o f t - b o d i e d s p e c i e s o r soft parts
o f s h e l l - o r s k e l e t o n - b e a r i n g s p e c i e s . T h e o n l y r e c o r d s k n o w n t o u s o f soft-
b o d y p r e s e r v a t i o n i n t h e C i n c i n n a t i a n are a w o r m d e s c r i b e d b y U l r i c h
(1878) a n d t h e r e c e n t d i s c o v e r y of f o s s i l i z e d " t u b e f e e t " in a brittle star
( G l a s s 2006). O u r k n o w l e d g e o f t h e C i n c i n n a t i a n biota i s thus h e a v i l y bi-
a s e d i n favor o f s p e c i e s w i t h h a r d p a r t s , the shells a n d s k e l e t o n s , c o m p l e t e
or p a r t i a l , k n o w n as b o d y f o s s i l s . F o r t u n a t e l y , this is offset to s o m e d e g r e e
b y e v i d e n c e o f t h e a c t i v i t y o f s o f t - b o d i e d s p e c i e s f r o m t r a c e fossils (bur-
r o w s , tracks, a n d t r a i l s t h e subject of c h a p t e r 14). H o w e v e r , it m u s t be kept
i n m i n d that p o t e n t i a l l y g r e a t n u m b e r s o f s p e c i e s i n t h e biota w i l l n e v e r b e
k n o w n b e c a u s e t h e y left n o fossil r e c o r d w h a t s o e v e r .
S h e l l s a n d s k e l e t o n s p r e s e r v e d i n C i n c i n n a t i a n strata are p r e d o m i -
n a n t l y c o m p o s e d o f c a l c i u m c a r b o n a t e ( C a C O 3 ) i n t h e m i n e r a l form c a l -
c i t e . S o m e shells o f b r a c h i o p o d s (see c h a p t e r 8 ) a n d the microfossils k n o w n
as c o n o d o n t s (see c h a p t e r 13) are p r e s e r v e d as c a l c i u m p h o s p h a t e . D e s p i t e
t h e a b u n d a n c e o f c a l c i u m c a r b o n a t e i n C i n c i n n a t i a n fossils, not all shells
h a v i n g this c h e m i c a l c o m p o s i t i o n are e q u a l l y well p r e s e r v e d . T h e reason
for this i s t h a t s o m e o r g a n i s m s f o r m c a l c i u m c a r b o n a t e shells o r s k e l e t o n s
n o t as c a l c i t e b u t as a d i f f e r e n t m i n e r a l c a l l e d a r a g o n i t e . A r a g o n i t e , w i t h a
d i f f e r e n t c r y s t a l l o g r a p h i c s t r u c t u r e t h a n c a l c i t e , b e c o m e s u n s t a b l e i n sea-
w a t e r after d e a t h of t h e o r g a n i s m a n d r e c r y s t a l l i z e s as calcite. In s o m e c a s e s
this t r a n s f o r m a t i o n o c c u r s as a solid-state r e p l a c e m e n t of a r a g o n i t e by cal-
cite, a l t e r i n g t h e m i c r o s t r u c t u r e b u t r e t a i n i n g t h e m a c r o s c o p i c s t r u c t u r e
o f a s h e l l . A r a g o n i t i c shells c a n a l s o b e lost e n t i r e l y b y d i s s o l u t i o n e v e n
b e f o r e b u r i a l i n s e d i m e n t . I n o t h e r c a s e s , a shell m a y b e c o m e b u r i e d , a n d
a s t h e i n t e r n a l soft parts d e c a y , s e d i m e n t s e e p s into t h e shells, r e p l a c i n g the
soft parts a n d f o r m i n g a p e r f e c t m o l d of t h e interior. A f t e r the a r a g o n i t i c
shell d i s s o l v e s , t h e s e d i m e n t i n f i l l i n g r e m a i n s a n d c a n b e lithified b y c a l -
cific c e m e n t . I n t h i s m a n n e r a n i n t e r n a l m o l d o r s t e i n k e r n i s f o r m e d
w h i c h p e r f e c t l y p r e s e r v e s t h e i n t e r n a l s p a c e s o f a s h e l l , often m o l d i n g fea-
t u r e s o f t h e i n n e r shell s u r f a c e l i k e m u s c l e scars, e v e n t h o u g h t h e a c t u a l
o r i g i n a l a r a g o n i t i c shell d i s a p p e a r s . I n o t h e r c a s e s t h e shell m a y n o t b e
i n f i l l e d , a n d o n c e t h e shell d i s s o l v e s , a void r e m a i n s a s a n e x t e r n a l m o l d
o f t h e o u t e r s u r f a c e o f t h e s h e l l , o r t h e e x t e r n a l m o l d c a n b e infilled w i t h
s e d i m e n t to form a c a s t . T h e s e arc o f t e n t h e o n l y w a y s a record of an ara-
g o n i t i c s h e l l i s p r e s e r v e d , a n d w e h a v e n o w a y o f g a u g i n g how m a n y ara-
g o n i t i c s h e l l s d i s s o l v e d l e a v i n g no t r a c e w h a t s o e v e r . T h u s it is very difficult
t o e s t i m a t e t h e o r i g i n a l a b u n d a n c e o f s p e c i e s f o r m i n g a r a g o n i t i c shells.
E v e n a m o n g s p e c i e s f o r m i n g c a l c i t i c shells, p r e s e r v a t i o n c a n b e h i g h l y
selective. T h i n n e r , m o r e d e l i c a t e shells are m o r e likely t o b e d e s t r o y e d
b e f o r e t h e y c a n b e b u r i e d . I n g r o u p s like trilobites (see c h a p t e r 11), t h e
exoskeleton is c o m p o s e d of the protein chitin, with varying a m o u n t s of
c a l c i u m c a r b o n a t e . J u v e n i l e , o r n e w l y m o l t e d , trilobites h a d w e a k l y c a l c i -
f i e d e x o s k e l e t o n s , a n d w e r e t h u s less p r e s e r v a b l e t h a n m o r e h e a v i l y c a l c i -
fied individuals. T h u s , w i t h i n a single species, preservational potential is

6 A Sea without Fish

u n e q u a l . S p e c i e s h a v i n g shells f o r m e d o f o n e o r t w o v a l v e s (snails, c l a m s ,
or brachiopods) have a h i g h e r preservation potential than species with
m u l t i - p a r t e d s k e l e t o n s s u c h as c r i n o i d s or trilobites. M u l t i - p a r t e d s k e l e t o n s
are held t o g e t h e r with c o n n e c t i v e tissue, w h i c h is s u s c e p t i b l e to s c a v e n g i n g
and d e c a y , c a u s i n g t h e s k e l e t o n t o b e c o m e d i s a r t i c u l a t e d a n d s c a t t e r e d b y
currents. The c o n s e q u e n c e o f all t h e s e v a r i a b l e factors o f shell c o m p o s i t i o n
a n d s t r u c t u r e is that all o r g a n i s m s p r o d u c i n g a c a l c i t i c shell c a p a b l e of
p r e s e r v a t i o n d o n o t h a v e a n e q u a l p o t e n t i a l for a c t u a l p r e s e r v a t i o n . Pres-
e r v a t i o n i s h i g h l y s e l e c t i v e e v e n a m o n g shells c h e m i c a l l y a n d m i n e r a l o g i -
c a l l y stable e n o u g h t o survive post m o r t e m .
T h e m o d e o f life o f o r g a n i s m s d e t e r m i n e s p r e s e r v a t i o n p o t e n t i a l e v e n
before a n i m a l s die. For a q u a t i c s p e c i e s , b o t t o m - d w e l l e r s ( b e n t h o s ) h a v e a
higher likelihood of preservation than s w i m m i n g (nektonic) or floating
( p l a n k t o n i c ) s p e c i e s . A m o n g t h e b e n t h o s , s p e c i e s that burrow into t h e
s e d i m e n t for a l i v i n g ( i n f a u n a ) o b v i o u s l y h a v e a m u c h h i g h e r p o t e n t i a l for
preservation than do surface dwellers (epifauna). A m o n g the epifauna,
species living p e r m a n e n t l y attached to the b o t t o m often have a h i g h e r
p o t e n t i a l for p r e s e r v a t i o n t h a n f r e e - l i v i n g , m o b i l e s p e c i e s , s i m p l y b e c a u s e
t h e y are u n a b l e t o e s c a p e s u d d e n burial b y s e d i m e n t .

Processes of Mortality

F o s s i l i z a t i o n is a rare e v e n t , n o t a p r o c e s s h a p p e n i n g e v e n day. M o s t a n i - The paleoecologist must

m a l s that survive t h r o u g h old a g e a n d d i e o f " n a t u r a l c a u s e s " s u c h a s p r e d a - never forget that he is
tion o r d i s e a s e will n o t b e c o m e f o s s i l i z e d . U n b u r i e d c a r c a s s e s are torn studying not the living

apart b y predators a n d s c a v e n g e r s o r d e s t r o y e d b y d e c a y a n d e x p o s u r e t o inhabitants of the vil-

lage but only the bodies
the e l e m e n t s . F o s s i l i z a t i o n very often d e p e n d s on a rare, c a t a s t r o p h i c e v e n t
in the churchyard, and
that b u r i e s a n e n t i r e a s s e m b l a g e o f l i v i n g o r g a n i s m s , m u c h i n w a y the
then only after many
e r u p t i o n of Mt. V e s u v i u s b u r i e d P o m p e i i in AD 7 9 , p r e s e r v i n g i n c r e d i b l e
visits by grave robbers.
details o f R o m a n life. Thus, processes of mortality are of f u n d a m e n t a l
Derek V. A g e r
i m p o r t a n c e i n d e t e r m i n i n g how o r g a n i s m s are p r e s e r v e d . W h e n w e see a
1963, 184
fossil, t h e first q u e s t i o n s h o u l d b e : " W h a t h a p p e n e d ? " T h e a n s w e r m a y tell
u s m o r e a b o u t the n a t u r e o f rare e v e n t s , s u c h a s s t o r m s , e a r t h q u a k e s , o r
volcanic eruptions than about day-to-day processes.
I n t h e C i n c i n n a t i a n , t h e b e s t - p r e s e r v e d fossils, s u c h a s c o m p l e t e trilo-
bites or c r i n o i d s , p r o b a b l y resulted from s u d d e n b u r i a l of a sea floor p o p u l a -
tion b y m u d d y s e d i m e n t . G r e a t s t o r m s are c a p a b l e o f s h i f t i n g m a s s e s o f
s e d i m e n t a r o u n d on the sea floor or s t i r r i n g it into s u s p e n s i o n , o n l y to settle-
out as a b l a n k e t over the b o t t o m w h e n t h e s t o r m s u b s i d e s (see c h a p t e r 4).
Organisms were smothered by these events and protected from the n o r m a l
cycle of scavenging, decay, and destruction. T h e s e c a s e s offer t h e b e s t o p -
p o r t u n i t y t o s e e a s n a p s h o t o f O r d o v i c i a n m a r i n e life. B u t e v e n h e r e w e
should b e c a u t i o u s , b e c a u s e s u c h s m o t h e r i n g e v e n t s c a n p r e s e r v e n o t o n l y
o r g a n i s m s l i v i n g a t the t u n c , but also r e m a i n s l o n g d e a d a n d a c c u m u l a t e d
over t i m e . I n d e e d , m a n y h i g h l y fossiliferous l i m e s t o n e b e d s i n t h e C i n c i n -
n a t i a n represent l o n g - t e r m ( t i m e - a v e r a g e d ) a c c u m u l a t i o n s o f s h e l l y m a t e -
rial a l o n g w i t h better-preserved s p e c i m e n s that w e r e b u r i e d a l i v e i n a n
instantaneous event. W i t h care, these c o m p o n e n t s can be r e c o g n i z e d , so

Introduction 7
 t h a t w e c a n assess w h a t s p e c i e s m a d e u p the l i f e a s s e m b l a g e . T h e d e a t h
a s s e m b l a g e o f r e m a i n s a l r e a d y d e a d a t t h e t i m e o f b u r i a l i s also i n f o r m a -
tive, b e c a u s e , like a g r a v e y a r d , it c a n record m u l t i p l e g e n e r a t i o n s a n d o c -
c u r r e n c e of rare s p e c i e s . T a b l e 1 lists s o m e of the most useful c h a r a c t e r i s t i c s
to look for in d i s t i n g u i s h i n g fossils b u r i e d w h i l e l i v i n g from those a c c u m u -
lated g r a d u a l l y a s d e a d r e m a i n s .

Table 1. Characteristics of Life a s s e m b l a g e Death assemblage

Life Assemblages and
Articulation good disarticulated
Death Assemblages
Breakage rare common

Abrasion rare common

Preserved in life position maybe not often

Size-sorting uncommon possible

History If, in light of t h e f o r e g o i n g d i s c u s s i o n , t h e reader is not fully c o n v i n c e d of

the e x t r e m e rarity o f f o s s i l i z a t i o n and t h e u n i q u e n e s s o f the f o s s i l r i c h n e s s

. . . our search for a o f t h e C i n c i n n a t i a n strata, t h e f o l l o w i n g s e c t i o n s h o u l d p r o v i d e a d d i t i o n a l

mechanism forces us to f o o d for t h o u g h t . S u b s e q u e n t t o life a n d d e a t h d u r i n g the L a t e O r d o v i c i a n

range far beyond the P e r i o d , 450 m i l l i o n years a g o , the r e m a i n s o f m a r i n e a n i m a l s w e r e b u r i e d
Cincinnati region and in s e d i m e n t . W o r l d w i d e , a great many fossils from very a n c i e n t P a l e o z o i c
consider the geologic his- strata are p o o r l y p r e s e r v e d b e c a u s e t h e y h a v e suffered g r e a t l y f r o m t h e
tory of much of eastern
ravages of t i m e c h e m i c a l and physical modifications o c c u r r i n g d u r i n g
North America, especially
and alter burial. T h e s e c h a n g e s , technically k n o w n as d i a g e n e s i s , include
the continental collisions
d i s s o l u t i o n o f o r i g i n a l shell m a t e r i a l , w i t h o r w i t h o u t r e p l a c e m e n t b y o t h e r
... referred to as the
Taconic, Acadian, and m i n e r a l s , p e r v a s i v e r e c r y s t a l l i z a t i o n of the r o c k , w i t h partial or c o m p l e t e

Alleghenian orogenies. obliteration of fossil contents, or c r u s h i n g and deformation of fossils during

c o m p a c t i o n o f t h e e n c l o s i n g s e d i m e n t s , f o s s i l s that s u r v i v e d i a g e n e t i c
Paul E. P o t t e r 1996, 71
d a m a g e a t low t e m p e r a t u r e s m a y b e later d e s t r o y e d b y a c t u a l m e t a m o r -
p h i s m , i n w h i c h d e e p l y b u r i e d strata are h e a t e d , r e c r y s t a l l i z e d , a n d d e -
formed to varying degrees. M e t a m o r p h i c processes transform primary
s e d i m e n t a r y r o c k s s u c h as s h a l e s into slates, l i m e s t o n e s into m a r b l e , and
s a n d s t o n e s into q u a r t z i t e , a c c o m p a n i e d b y n e a r l y total obliteration o f fos-
sils a n d o t h e r p r i m a r y f e a t u r e s of t h e s e d i m e n t . M e t a m o r p h i s m is associ-
ated with d e e p b u r i a l by o v e r l y i n g strata, or m o u n t a i n - b u i l d i n g p r o c e s s e s
o f t e c t o n i c s , i n c l u d i n g f o l d i n g , f a u l t i n g , s h e a r i n g , a n d v o l c a n i c activity.
T h e pristine quality of m a n y C i n c i n n a t i a n fossils is clear e v i d e n c e that
they h a v e u n d e r g o n e very little d i a g e n e t i c alteration and no m e t a m o r p h i c
c h a n g e over their l o n g burial since the O r d o v i c i a n . How c o u l d these fossils
have survived with so little alteration over s u c h a vast span of time? T h e a n -
swer is " l o c a t i o n - l o c a t i o n - l o c a t i o n " and the history of the C i n c i n n a t i A r c h .
B e c a u s e t h e C i n c i n n a t i region is located inland from the c o n t i n e n t a l
m a r g i n s , it is far distant from regions that h a v e u n d e r g o n e intense d e f o r m a -
tion and m e t a m o r p h i s m over the c o u r s e of t i m e . The closest d e f o r m e d strata
of the A p p a l a c h i a n t e c t o n i c z o n e lie a b o u t 200 m i l e s to the southeast (Pine
M o u n t a i n , T e n n e s s e e ) , a n d m e t a m o r p h o s e d rocks are e v e n farther (the

8 A Sea without Fish

 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 contact
that defines the Findlay
and Kankakee branches.

G r e a t S m o k i e s and Blue R i d g e o f T e n n e s s e e ) . F u r t h e r m o r e , the U p p e r O r -

d o v i c i a n strata o f the C i n c i n n a t i r e g i o n w e r e n e v e r b u r i e d d e e p l y b e n e a t h
y o u n g e r s e d i m e n t a r y strata. A t t h e close o f O r d o v i c i a n t i m e i n t h e r e g i o n ,
there is e v i d e n c e that the seas b e c a m e very shallow, the sea floor p e r h a p s
e x p o s e d subacrially (above sea level), p r o d u c i n g a g a p in t h e stratal record
k n o w n a s a n u n c o n f o r m i t y . S u b s e q u e n t s e d i m e n t a t i o n from the S i l u r i a n
through P e n n s v l v a n i a n Periods (a span of 150 m i l l i o n years) was again sub-
m a r i n e , but mostly of very shallow water o r i g i n s . A f t e r t h e P e n n s y l v a n i a n
Period (about 290 m i l l i o n years ago), there is no record of f u r t h e r m a r i n e
s e d i m e n t a t i o n in all of O h i o . A l t o g e t h e r , the total t h i c k n e s s of strata d e p o s -
ited over the O r d o v i c i a n m a y h a v e b e e n 3 0 0 - 6 0 0 m e t e r s (1000-2000 feet) a t
m o s t (Potter 2007). I n d i c a t i o n s of s h a l l o w i n g , retreating seas s u g g e s t that the
C i n c i n n a t i region was u n d e r g o i n g r e g i o n a l uplift, b e c a u s e m a r i n e deposits
are thicker t o the east and west t h a n those closer t o C i n c i n n a t i . O n t h e c o n -
tinental scale as w e l l , e p i c o n t i n e n t a l seas retreated by the e n d of t h e P a l e o -
zoic, o n l y t o return d u r i n g the M e s o z o i c , but farther west t h a n O h i o .
T h e r e g i o n a l uplilt that a f f e c t e d the C i n c i n n a t i r e g i o n w a s p a r t o f a
broad z o n e c a l l e d the C i n c i n n a t i A r c h , t r e n d i n g n o r t h - s o u t h a n d s p l i t t i n g
just n o r t h of C i n c i n n a t i into the n o r t h e a s t e r l y - t r e n d i n g F i n d l a y A r c h a n d
t h e n o r t h w e s t e r l y - t r e n d i n g K a n k a k e e A r c h ( F i g u r e 1. 5). The term " a r c h "
i m p l i e s a n u p w a r p i n g o f the E a r t h ' s c r u s t , b u t a n y o n e w h o h a s n o t i c e d t h e

Introduction 9
Figure 1. 6. Diagrammatic strata o f l i m e s t o n e a n d s h a l e e x p o s e d i n r o a d c u t s a r o u n d C i n c i n n a t i has
east-west cross section s e e n e s s e n t i a l l y h o r i z o n t a l layers. It is o n l y w h e n we travel e a s t w a r d or
from near Bedford in w e s t w a r d f r o m C i n c i n n a t i that w e e n c o u n t e r g e o l o g i c a l l y y o u n g e r strata
south-central Indiana
o v e r l y i n g t h e O r d o v i c i a n strata a l o n g t h e a x i s o f t h e A r c h , a n d t h e c h a r a c -
across the Cincinnati
ter o f t h e A r c h a s a v e r y g e n t l e , b r o a d u p w a r p i n g b e c o m e s a p p a r e n t . T h e
Arch to near Portsmouth,
tilt or d i p of the strata across the A r c h is u s u a l l y less t h a n o n e d e g r e e , or
Ohio. From Potter (1996)
and reprinted by permis- f o u r to s e v e n feet p e r m i l e , in H a m i l t o n C o u n t y (Potter 1996). O v e r a dis-

sion of the Kentucky t a n c e o f a b o u t 8 0 k m (50 m i l e s ) cast a n d w e s t o f C i n c i n n a t i , a cross-section

Geological Survey. t h r o u g h t h e b e d r o c k s h o w s t h e s t r u c t u r e o f t h e A r c h clearly ( f i g u r e 1 . 6).
E v e n t h o u g h t h e C i n c i n n a t i r e g i o n i s distant f r o m t h e d e f o r m e d rocks o f
the A p p a l a c h i a n m o u n t a i n b e l t , t h e C i n c i n n a t i A r c h m a y h a v e resulted
f r o m t h e s a m e l a r g e - s c a l e t e c t o n i c p r o c e s s e s that u p l i f t e d t h e A p p a l a -
c h i a n s , b e g i n n i n g i n O r d o v i c i a n t i m e . T h e r e a d e r i s referred t o Potter
(2007) for f u r t h e r d i s c u s s i o n o f t h e o r i g i n o f t h e C i n c i n n a t i A r c h .
B e c a u s e uplift c o n t i n u e d a l o n g the axis of the C i n c i n n a t i A r c h , it was
c o n t i n u a l l y e r o d e d , stripping a w a y strata l y i n g a b o v e the O r d o v i c i a n , pre-
v e n t i n g their d e e p b u r i a l . D u r i n g t h e past t w o m i l l i o n years, the glaciers o f
the P l e i s t o c e n e E p o c h c o v e r e d m o s t o f O h i o a n d t h e ice sheet scraped a w a y
r e m a i n i n g o v e r b u r d e n or w a s h e d it away- as the ice m e l t e d , c o m p l e t i n g the
e x p o s u r e of O r d o v i c i a n strata at the surface. By these processes, b e g u n virtu-
ally a t the s a m e t i m e t h e O r d o v i c i a n seas c o v e r e d the C i n c i n n a t i r e g i o n ,
s e d i m e n t s a n d fossils deposited t h e n w e r e n e v e r d e e p l y b u r i e d and d e f o r m e d ,
a n d b e c a m e e x p o s e d over a broad belt t h r o u g h the region. The significant
c o n s e q u e n c e is that the entire C i n c i n n a t i A r c h region has o n e of the most
e x t e n s i v e s u r f a c e e x p o s u r e s o f U p p e r O r d o v i c i a n strata i n N o r t h A m e r i c a , i f
n o t t h e entire world. N a t u r a l e x p o s u r e s ( o u t c r o p s ) in s t r e a m b e d s and the
sides of valleys ( F i g u r e 1. 7 A ) and h u m a n - m a d e e x p o s u r e s , m a i n l y as roadcuts

10 A Sea without Fish

Figure 1. 7. A. Natural exposure of Cincinnatian strata in the bed of Stonelick Creek, Clermont County, Ohio.
Streambed outcrops have wide surfaces of fossiliferous beds (bedding planes) that provide information
about fossil distribution and orientation. Here, the fossil-rich Bellevue Limestone is exposed and examined
by participants in a 1981 Geological Society of America field trip. B. One of the most extensive roadcut
exposures of Cincinnatian strata, 1. 3 km (0. 8 miles) in length and about 75 m (250 ft) high, along Kentucky
Route 3071, leading to the Ohio River, west of Maysville, Mason County, Kentucky. The lowest strata ex-
posed are the lower Cincinnatian (Edenian Stage) Kope Formation, and the highest strata are the Bellevue
Limestone (Maysvillian Stage). Photo by Paul E. Potter.

Introduction 11
Figure 1. 8. Trammel Fos- ( F i g u r e s 1. 7 B , 1. 8) a n d q u a r r i e s , p r o v i d e a c c e s s to the O r d o v i c i a n b e d r o c k
sil Park, Sharonville, t h r o u g h o u t the r e g i o n . In the C i n c i n n a t i A r c h region we have a truly u n i q u e
Hamilton County, Ohio. w i n d o w to t h e p a s t e a s y a c c e s s to a n c i e n t strata a n d fossils that e l s e w h e r e
This is a ten acre hillslope
lie b u r i e d u n d e r t h o u s a n d s o f m e t e r s o f rock.
where construction ex-
T h e foregoing overview s h o w s that t h e a b u n d a n c e o f fossils i n the
posed four fossiliferous
O r d o v i c i a n r o c k s o f t h e C i n c i n n a t i r e g i o n i s t h e result o f m a n y i n t e r a c t i n g
formations: the Fairview
factors. B e c a u s e o f t h i s u n i q u e a n d f o r t u n a t e c o m b i n a t i o n o f factors, t h e
Formation, Miamitown
Shale, Bellevue Lime- C i n c i n n a t i r e g i o n b e c a m e o n e o f t h e earliest c e n t e r s o f intense interest and

stone, and Corryville For- s t u d y o f fossils i n N o r t h A m e r i c a . S c o r e s o f O r d o v i c i a n fossils w e r e first

mation. The developer d i s c o v e r e d a n d d e s c r i b e d f r o m this r e g i o n , a n d m a i n p r a c t i c e s and c o n -
R. L. Trammel donated cepts of paleontology and geology originated from research on C i n c i n n a -
the site to the City of tian fossils a n d r o c k s . B e c a u s e t h e C i n c i n n a t i r e g i o n w a s o n e o f t h e b i r t h -
Sharonville as an educa- p l a c e s o f m o d e r n g e o l o g i c a l s c i e n c e , w e w i l l e x p l o r e the early history o f
tional, geological park
s t u d y o f t h e fossils a n d r o c k s h e r e i n t h e f o l l o w i n g c h a p t e r .
where visitors can learn
about the Ordovician
geology and paleontol-
ogy. The park has easy
access, parking space,
and includes interpretive
signage and a protected,
in situ fossil shell pave-
ment. Because of the
abundance of fossils, sur-
face collecting of small
specimens is permitted.

12 A Sea without Fish

Figure 2. 1. Members of the Cincinnati School of Paleontology who were amateur paleontologists: A. U. P.
James, publisher and owner of the James Book Store. B. S. A. Miller, attorney. C. Charles Faber, real-
for. D. C. B. Dyer; who, after he retired as a maker of soap and candles, devoted himself to fossil collect-
ing. Photograph of Dyer from an old album in the possession of Richard Arnold Davis ( Richard Arnold
Davis); all others from the Department of Geology, University of Cincinnati.

14 A Sea without Fish

SCIENCE IN THE HINTERLAND:
THE CINCINNATI SCHOOL
OF PALEONTOLOGY

T h e rocks b e n e a t h and a r o u n d C i n c i n n a t i w e r e d e p o s i t e d i n a n interval o f

t i m e universally c a l l e d the O r d o v i c i a n Period. This t i m e u n i t w a s p r o p o s e d
f o r m a l l y in 1879. In the s e c o n d h a l f of the n i n e t e e n t h c e n t u r y , b e g i n n i n g
e v e n before the O r d o v i c i a n Period w a s n a m e d , t h e r e w a s i n the r e g i o n o f
C i n c i n n a t i , Ohio, a g r o u p o f p a l e o n t o l o g i s t s w h o h a v e b e e n c a l l e d t h e " C i n -
cinnati S c h o o l of P a l e o n t o l o g y . " T h e r e is no s i n g l e , definitive list of the m e m -
bers o f the C i n c i n n a t i S c h o o l , a n d different authors h a v e i n c l u d e d different
p e o p l e as m e m b e r s , d e p e n d i n g on the p u r p o s e s of their c o m p i l a t i o n s . N o r is
there a definitive list of iron-clad criteria as to w h o s h o u l d be c o n s i d e r e d a
m e m b e r a n d w h o should not. N o n e t h e l e s s , the i n d i v i d u a l s i n c l u d e d i n the
bod) of this chapter have a n u m b e r of characteristics in c o m m o n .
First, t h e y w e r e all s e r i o u s c o l l e c t o r s o f l o c a l fossils. B u t t h e y w e n t
b e y o n d that. T h e y did n o t just a m a s s h o r d e s o f fossils. T h e y also assidu-
ously s t u d i e d their f i n d s a n d w h e r e they f o u n d t h e m . But t h e y w e n t b e y o n d
that, too. T h e y s h a r e d their finds w i t h o n e a n o t h e r , a n d t h e y s h a r e d their
i n f o r m a t i o n a b o u t fossils a n d their t h i n k i n g a b o u t fossils n o t o n l y w i t h i n
the l o c a l f o s s i l - c o l l e c t i n g c o m m u n i t y , b u t w i t h t h e w o r l d a s a w h o l e ,
through publication.
A significant n u m b e r o f the m e m b e r s o f the C i n c i n n a t i S c h o o l p r o -
d u c e d lists o f fossils, i n d i c e s , b i b l i o g r a p h i e s , a n d o t h e r c o m p i l a t o r y w o r k s .
But these are just o n e a s p e c t of an e s s e n t i a l c r i t e r i o n for i n c l u s i o n in t h e
Cincinnati School, namely, publication.
S e c o n d , there is a g e o g r a p h i c c o m p o n e n t . W h e t h e r b o r n in C i n c i n n a t i
or not, individuals spent a significant p o r t i o n of their lives, e s p e c i a l l y their
formative years, in the t y p e - C i n c i n n a t i a n o u t c r o p area. M o r e o v e r , all or m o s t
of their p u b l i s h e d work was p u b l i s h e d l o c a l l y i n scientific j o u r n a l s or in
books or other p u b l i c a t i o n s that w e r e printed in the C i n c i n n a t i area.
T h i r d , they all w e r e a m a t e u r s , i n t h e s e n s e that f i n d i n g a n d p u b l i s h i n g
a b o u t fossils w a s not how t h e y m a d e their l i v i n g s . T h i s c r i t e r i o n is a bit
difficult t o a p p l y c o n s i s t e n t l y , h o w e v e r , b e c a u s e s o m e d i d sell fossils a n d
s o m e did sell b o o k s a n d o t h e r p u b l i s h e d m a t t e r . M o r e o v e r , a l u c k y f e w
w e n t from h u m b l e , a m a t e u r b e g i n n i n g s i n the C i n c i n n a t i area t o b e c o m e
respected m e m b e r s of t h e g e o l o g i c a n d p a l e o n t o l o g i c profession as a w h o l e .
B u t e v e n they b e g a n as l o c a l a m a t e u r s .
F o u r t h , there i s a t i m e - a n d - p l a c e c o m p o n e n t . T h e C i n c i n n a t i S c h o o l
o f Paleontology w a s e s s e n t i a l l y a p h e n o m e n o n o f t h e p e r i o d b e t w e e n the
A m e r i c a n C i v i l W a r a n d shortly after t h e s u c c e e d i n g t u r n o f t h e c e n t u r y .
All o f the m e m b e r s w e r e a s s o c i a t e d w i t h t h e C i n c i n n a t i S o c i e t y o f N a t u r a l
History d u r i n g that p e r i o d (and s o m e , w i t h t h e W e s t e r n A c a d e m y o f N a t u -

15
 ral S c i e n c e s that p r e c e d e d t h e s o c i e t y ) . I n p r e s e n t - d a y " b u z z - w o r d " t e r m i -
nology, they comprised a " l e a r n i n g c o m m u n i t y . " They worked together;
t h e y s h a r e d r e s o u r c e s ; t h e y c o m m u n i c a t e d with o n e a n o t h e r ; t h e y e n c o u r -
a g e d o n e a n o t h e r ; t h e y c o m p e t e d a g a i n s t o n e a n o t h e r . A b o v e all, t h e y
s t i m u l a t e d o n e a n o t h e r to p e r f o r i n at a h i g h e r level t h a n t h e y o t h e r w i s e
m i g h t h a v e d o n e . T h e w h o l e w a s m o r e t h a n t h e s u m o f its parts. T h e r e w a s
true synergism in the C i n c i n n a t i S c h o o l of Paleontology.
A l t h o u g h c a l l e d a s c h o o l , t h e C i n c i n n a t i S c h o o l w a s n o t o n e , nor did
i t h a v e a n y f o r m a l r e l a t i o n s h i p w i t h a n y c o l l e g e o r university. ( T h e U n i v e r -
sity o f C i n c i n n a t i , a s s u c h , w a s n o t f o u n d e d u n t i l 1870, a n d t h e r e w a s n o
D e p a r t m e n t o f G e o l o g y t h e r e u n t i l t h e first d e c a d e o f t h e t w e n t i e t h c e n -
tury, w h e n t h e D e p a r t m e n t o f G e o l o g y a n d G e o g r a p h y w a s initiated. )
But w e n e e d t o p u t the C i n c i n n a t i S c h o o l into m o r e o f a n historical
p e r s p e c t i v e . I n the s e c o n d d e c a d e o f t h e n i n e t e e n t h c e n t u r y , C i n c i n n a t i was
t h e largest city w e s t of t h e A l l e g h e n i e s , a n d a l o c a l p h y s i c i a n , D a n i e l D r a k e ,
figured that the city n e e d e d a f i r s t - c l a s s m u s e u m . H e n c e , h e s p e a r h e a d e d
t h e e s t a b l i s h m e n t o f t h e W e s t e r n M u s e u m . A s part o f t h e preparations for
t h e o p e n i n g o f t h e n e w m u s e u m , a t a x i d e r m i s t a n d artist n a m e d John James
A u d u b o n w a s h i r e d a n d w o r k e d for t h e o r g a n i z a t i o n for a b o u t a year, before
m o v i n g o n e v e n t u a l l y t o b e c o m e t h e m o s t f a m o u s bird artist the U n i t e d
States has p r o d u c e d . In a n y c a s e , t h e W e s t e r n M u s e u m o p e n e d in 1820.
U n f o r t u n a t e l y , t h e r e w a s a d e p r e s s i o n in t h e 1820s, a n d t h e W e s t e r n
M u s e u m fell u p o n h a r d t i m e s . T o m a k e m a t t e r s w o r s e , D r . D r a k e h a d left
t h e a r e a . A l t h o u g h a b l e t o c o n t i n u e its o p e r a t i o n s , t h e W e s t e r n M u s e u m
s a n k t o b e i n g little m o r e t h a n a c h a m b e r o f horrors.
D a n i e l D r a k e r e t u r n e d t o t h e a r e a i n t h e 1830s. H e still f i g u r e d that
t h e city n e e d e d a f i r s t - c l a s s m u s e u m , s o h e s p e a r h e a d e d t h e e s t a b l i s h m e n t
o f t h e W e s t e r n A c a d e m y o f N a t u r a l S c i e n c e s . B y the t i m e the ink w a s dry
o n t h e d o c u m e n t s i g n e d a t A p p o m a t t o x C o u r t H o u s e that e n d e d t h e A m e r -
ican C i v i l War, the W e s t e r n A c a d e m y of Natural S c i e n c e s was m o r i b u n d ,
a n d t h e W e s t e r n M u s e u m w a s little m o r e t h a n a c o l l e c t i o n o f c u r i o s i t i e s
a n d a c h a m b e r of horrors.
In the late 1860s the c u l t u r a l a n d civic leaders of C i n c i n n a t i figured that
the city n e e d e d a f i r s t - c l a s s m u s e u m , a n d t h e C i n c i n n a t i S o c i e t y o f N a t u r a l
History w a s established in 1870. A b o u t a year after the f o u n d i n g of the C i n c i n -
nati S o c i e t y o f N a t u r a l History, the h a n d f u l o f r e m a i n i n g m e m b e r s o f the old
W e s t e r n A c a d e m y of N a t u r a l S c i e n c e s d e c i d e d to transfer all the assets of the
a c a d e m y to t h e n e w society, i n c l u d i n g its m o n e y , s p e c i m e n s , and library. In
r e t u r n , t h e m e m b e r s o f the a c a d e m y w e r e t o b e m e m b e r s o f t h e society for
life. T h u s c a m e into b e i n g the C i n c i n n a t i S o c i e t y o f N a t u r a l History that was
a part of t h e lives of all t h e m e m b e r s of the C i n c i n n a t i S c h o o l .
H e r e f o l l o w s a n a c c o u n t o f e a c h o f t h o s e m e m b e r s . (In a p p e n d i x 2 are
briefer e n t r i e s for o t h e r i n d i v i d u a l s w h o h a d c o n n e c t i o n s w i t h t h e t y p e -
C i n c i n n a t i a n a r e a , w i t h its r o c k s a n d fossils, o r w i t h b o t h . S o m e o f t h e s e
p e o p l e o c c a s i o n a l l y h a v e b e e n referred t o a s m e m b e r s o f t h e C i n c i n n a t i
School.)

16 A Sea without Fish

 Figure 2. 2 A. C o v e r of
an 7 8 4 9 publication of
the Western Academy of
Natural Sciences pub-
lished 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
of the Cincinnati School
of Paleontology.

U r i a h Pierson James ( F i g u r e 2.1 A) w a s born in t h e state o f N e w Y o r k in 1811, U. P. James

the son of a c a r p e n t e r . In 1831 he a n d his b r o t h e r , J o s e p h , t r a v e l e d to C i n -
c i n n a t i , w h e r e U. P. w o r k e d as a printer. By t h e e n d of t h e 1840s he w a s a
publisher and the proprietor of t h e J a m e s B o o k Store. In the s h o p he always
stocked the latest i n g e o l o g i c a l b o o k s , a n d h e d i s p l a y e d fossils i n t h e
windows.
U. P. James was very a c t i v e in the i n t e l l e c t u a l life of C i n c i n n a t i . He
served a t e r m as p r e s i d e n t a n d w a s l o n g - t i m e t r e a s u r e r of t h e W e s t e r n
A c a d e m y o f N a t u r a l S c i e n c e s . W h e n C h a r l e s L y e l l , p r o b a b l y t h e foremost

Science in the Hinterland 17

Figure 2.3. A. Albert Gallatin Wetherby was professor of natural history at the University of Cincinnati be-
fore he relocated to Harvard University and malacology. B. George W. Harper, long-time principal of
Woodward High School, where he facilitated the start of the careers of students Ray S. Bassler and John
M. Nickles. C. "Friendly enemies": left to right, August F. Foerste, Amadeus W. Grabau, and Edward O.
Ulrich, during the International Geological Congress in Washington, D.C, 1934 (picture taken by Ray S.
Bassler). Photograph of Wetherby courtesy of the Museum of Comparative Zoology, Harvard University (
President and Fellows of Harvard College); that of Harper, courtesy of the Archives and Rare Books Library,
University of Cincinnati; "Friendly enemies," from the Department of Geology, University of Cincinnati.

78 A Sea without Fish

Figure 2.4. Field work in the early days. A. John M. Nickles (left) and Ray S. Bassler, collecting from an
exposure of the Kope Formation, Cincinnati, Ohio, 1900. B. E. O. Ulrich at the contact between the Kope
and Point Pleasant Formations on the banks of the Ohio River below Covington, Kentucky, 1901. These
exposures are now underwater. (A and B from Bassler Archive, Department of Paleobiology, National Mu-
seum of Natural History, Smithsonian Institution, Washington, D.C, courtesy of JoAnn Sanner.)

Science in the Hinterland 19

Figure 2.5. Four Cincinnatians who became leading professional paleontologists. A. Charles Schuchert,
Professor, Yale University. B. John M. Nickles, U. S. Geological Survey, 1942. C. Ray S. Bassler, U. S.
National Museum. D. E. O. Ulrich, U. S. Geological Survey. (All photographs from the Department of Ge-
ology, University of Cincinnati.)

20 A Sea without Fish

 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 desig-
nated as the type section
of the Fairview Formation
by Ford (1967). (Image
courtesy of the Cincinnati
Historical Society Library,
Cincinnati Museum Cen-
ter.) B. Exposure of
Maysvillian strata (prob-
ably Corryville and Mt.
Auburn Formations) at
corner of Clifton Avenue
and Calhoun Street
(right), Cincinnati, 1900.
Note, at left, trolley car
and McMicken Hall of
the University of Cincin-
nati. This exposure has
been leveled and is pres-
ently the site of the Uni-
versity of Cincinnati Col-
lege of Law. (From the
Bassler Archive, Depart-
ment of Paleobiology,
g e o l o g i s t in the w o r l d , visited C i n c i n n a t i in t h e 1840s, J a m e s w a s o n e of his National Museum of Nat-
hosts. A b o u t the s a m e t i m e , U . P . J a m e s b e c a m e o n e o f t h e c h a r t e r m e m - ural History, Smithsonian
bers of t h e C i n c i n n a t i A s t r o n o m i c a l S o c i e t y ( a c c o r d i n g to a list at t h e Institution, Washington,
C i n c i n n a t i O b s e r v a t o r y , F e b r u a r y 11, 2007). H e w a s o n e o f t h e s u r v i v i n g D.C, courtesy of JoAnn
Sanner.)
m e m b e r s o f the W e s t e r n A c a d e m y w h e n i t w a s d i s s o l v e d a n d its assets w e r e
d o n a t e d t o the C i n c i n n a t i Society- o f N a t u r a l History i n 1872, w h e r e u p o n
he b e c a m e a lite m e m b e r in the society.
U. P. James's fossil c o l l e c t i o n w a s widely r e n o w n e d . L o u i s A g a s s i z , o n e
o f the foremost p a l e o n t o l o g i s t s i n this c o u n t r y , visited C i n c i n n a t i , a n d ,
after s e e i n g James's c o l l e c t i o n , p r o c l a i m e d it o n e of the finest he had ever
s e e n . James's favorite f o s s i l s s e e m t o h a v e b e e n b r y o z o a n s , w h i c h h e c o n -

Science in the Hinterland 21

 sidered t o b e c o r a l s . M a n y o f t h e t y p e - s p e c i m e n s i n his c o l l e c t i o n e n d e d
u p a t t h e U n i t e d States N a t i o n a l M u s e u m ; o t h e r m a t e r i a l w e n t t o t h e U n i -
versity o f C h i c a g o a n d t o t h e U n i v e r s i t y o f C i n c i n n a t i .
N o t o n l y w a s J a m e s t h e a u t h o r o f m a n y p a p e r s a b o u t l o c a l fossils, b u t
h e w a s t h e p u b l i s h e r o f m a n y o t h e r s . I n d e e d , J a m e s w a s the p u b l i s h e r o f
t h e j o u r n a l The Paleontologist ( F i g u r e 2.2), w h i c h ran for s e v e n n u m b e r s ,
b e g i n n i n g i n 1879. H e a l s o p u b l i s h e d a c a t a l o g u e o f C i n c i n n a t i freshwater
mussels and another of local plants.
U. P. J a m e s retired f r o m t h e b o o k s t o r e b u s i n e s s in 1886. He d i e d in
1889 a n d w a s b u r i e d i n C i n c i n n a t i ' s b e a u t i f u l S p r i n g G r o v e C e m e t e r y .
( B e c k e r 1938; B r a d s h a w , p e r s . c o m m . ; C a s t e r 1 9 5 1 , 1 9 8 1 , 1 9 8 2 ; C r o n e i s 1963;
C u f f e y , D a v i s , a n d U t g a a r d 2002; H e n d r i c k s o n 1947; H o w e , Fisher, a n d
K e e k e l e r 1889; J. F. J a m e s 1889; S h i d e l e r [1952] 2002; A n o n . 1849,1878.)

Joseph F. James Joseph F r a n c i s J a m e s a l m o s t c e r t a i n l y w a s i n d u c t e d into the w o n d e r s o f

fossil c o l l e c t i n g by his father, U. P. J a m e s (above). However, Joseph's inter-
ests i n n a t u r a l history w e r e b r o a d e r t h a n w e r e his father's; t h e son p u b -
l i s h e d n o t o n l y a b o u t fossils, b u t a b o u t p h y s i c a l g e o l o g y , b o t a n y , a n d o t h e r
subjects.
Joseph F. J a m e s b e g a n as a clerk in h i s father's b o o k s t o r e , but he be-
c a m e t h e f i r s t o f t h e C i n c i n n a t i S c h o o l t o g a i n professional status. H e w a s
e l e c t e d to m e m b e r s h i p in t h e C i n c i n n a t i S o c i e t y of N a t u r a l History in 1876
( C i n c i n n a t i S o c i e t y o f N a t u r a l History, 94), a n d h e l o n g was associated
w i t h that i n s t i t u t i o n a s a m e m b e r , officer, staff m e m b e r , a n d a u t h o r o f
p a p e r s in its j o u r n a l . A f t e r a t w o - y e a r stint in b u s i n e s s pursuits in C a l i f o r n i a
a n d a d j a c e n t states, h e w a s e l e c t e d c u s t o d i a n o f t h e society i n 1881 a n d held
that p o s i t i o n for six y e a r s . T h e p o s i t i o n o f c u s t o d i a n i n v o l v e d a g o o d d e a l
m o r e t h a n janitorial w o r k ; i t w o u l d a p p e a r that h e w a s i n c h a r g e o f day-to-
d a y o p e r a t i o n s o f t h e s o c i e t y ' s b u i l d i n g . M e a n w h i l e , h e was also professor
o f m e d i c a l botany a t t h e C i n c i n n a t i C o l l e g e o f P h a r m a c y .
I n 1886 J a m e s w a s e l e c t e d t o t h e c h a i r o f B o t a n y a n d G e o l o g y a t M i -
a m i U n i v e r s i t y i n O x f o r d , O h i o , " b u t this p o s i t i o n w a s lost t w o years later
t h r o u g h t h e d i s r u p t i o n o f t h e f a c u l t y a r i s i n g from r e l i g i o u s p r e j u d i c e s "
( G i l b e r t 1898, 2). " W h e n r e l i g i o u s b e l i e f s w e r e u n d e r f i r e a t O x f o r d , profes-
sor J a m e s w a s a c c u s e d o f b e i n g a n a g n o s t i c a n d d e f e n d e d a s b e i n g e s s e n -
tially a U n i t a r i a n . So far as I k n e w it, his r e l i g i o n w a s an u n s w e r v i n g d e v o -
tion to s c i e n c e " ( G i l b e r t 1898, 3).
F o r o n e y e a r , h e w a s professor o f n a t u r a l history a t t h e A g r i c u l t u r a l
C o l l e g e o f M a r y l a n d , d u r i n g w h i c h t i m e h e also did w o r k for the U n i t e d
States G e o l o g i c a l S u r v e y . T h e n , i n 1889 J a m e s w a s a p p o i n t e d assistant
p a l e o n t o l o g i s t w i t h t h e U n i t e d States G e o l o g i c a l S u r v e y i n t h e D i v i s i o n o f
P a l e o z o i c P a l e o n t o l o g y , i n W a s h i n g t o n , D . C . T w o years later, h e b e c a m e
assistant v e g e t a b l e p a t h o l o g i s t w i t h t h e U . S . D e p a r t m e n t o f A g r i c u l t u r e ,
a l s o i n W a s h i n g t o n , D . C , a n d served i n that p o s i t i o n for four years. D u r i n g
t h o s e four y e a r s , J a m e s d e v o t e d his e v e n i n g s t o the study o f m e d i c i n e a n d
graduated with a m e d i c a l d e g r e e from C o l u m b i a n University (now G e o r g e
W a s h i n g t o n U n i v e r s i t y ) i n 1895. H e s p e n t t h e w i n t e r o f 1895-1896 i n N e w

22 A Sea without Fish

York and L o n d o n d o i n g hospital w o r k a n d b a c t e r i o l o g i c a l study, after w h i c h
h e set u p i n m e d i c a l p r a c t i c e i n H i n g h a m , M a s s a c h u s e t t s .
Joseph K James w a s a prolific a u t h o r , n o t o n l y in p a l e o n t o l o g y , b u t a l s o
i n g e o l o g y a n d botany. N o t c o u n t i n g m a n y items i n n e w s p a p e r s a n d m a g a -
z i n e s , his o u t p u t a m o u n t e d t o w e l l o v e r o n e h u n d r e d s c i e n t i f i c p a p e r s
a b o u t e q u a l l y spread a m o n g t h o s e t h r e e a r e a s , a l o n g w i t h a n u m b e r o f
others o n m i s c e l l a n e o u s s u b j e c t s . S o m e o f his p a l e o n t o l o g i c p a p e r s w e r e
c o - a u t h o r e d with his lather, U.P. James. The y o u n g e r J a m e s w a s t h e a u t h o r
or c o - a u t h o r of a n u m b e r of taxa in the t y p e - C i n c i n n a t i a n , a n d at least o n e
was n a m e d after h i m .
Joseph F . James died o n M a r c h 2 9 , 1 8 9 7 , i n H i n g h a m , M a s s a c h u s e t t s ,
and his ashes w e r e b u r i e d i n 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 . ( B e c k e r
1938; Caster 19S2; C r o n e i s 1963; C u f f e y , D a v i s , a n d U t g a a r d 2002; G i l b e r t
1898; S h i d e l e r [1952] 2002; A n o n . 1 8 7 9 , 1 8 8 2 , 1 8 8 5 b , 1886a.)

C h a r l e s Brian D y e r ( F i g u r e 2.1D) w a s b o r n on April 1, 1806, n e a r D u d l e y C. B. Dyer

Castle, Worcestershire, England. H a v i n g h a d t o s u p p o r t h i m s e l f a n d his
m o t h e r , h e h a d little f o r m a l e d u c a t i o n , i f any. H e c a m e t o C i n c i n n a t i i n
1828 and set up as a m a n u f a c t u r e r of s o a p a n d c a n d l e s . A r o u n d 1850, h a v i n g
m a d e w h a t h e c o n s i d e r e d t o b e a s u f f i c i e n t s u m , h e retired a n d d e v o t e d
h i m s e l f t o fossil c o l l e t i n g .
D y e r was one of the original m e m b e r s of the C i n c i n n a t i Society of
N a t u r a l History, and he c o - a u t h o r e d p a p e r s w i t h S. A. M i l l e r (see b e l o w ) .
O n e o f t h e s e w a s t h e r e p o r t o f a c o m m i t t e e o n the g e o l o g i c a l n o m e n c l a -
ture o f the t y p e - C i n c i n n a t i a n a p p o i n t e d b y t h e s o c i e t y (S. A . M i l l e r e t al.
1879); o f t h e ten m e m b e r s o f that c o m m i t t e e , six o f t h e i n d i v i d u a l s are
g e n e r a l l y r e c o g n i z e d a s m e m b e r s o f t h e C i n c i n n a t i S c h o o l , a n d all o f t h e m
have b e e n listed in o n e p l a c e or a n o t h e r as c o l l e c t o r s of l o c a l fossils.
However, it is for his a v i d c o l l e c t i n g of fossils that C. B. D y e r is best
remembered. As a young m a n , he enjoyed hunting, but upon retirement
h e a b a n d o n e d the g u n i n favor o f t h e h a m m e r a n d live g a m e i n favor o f
l o n g - d e a d fossils. He was a w e l l - k n o w n c o l l e c t o r of fossils d u r i n g his life-
t i m e , to the extent that, in the 1870s, fossils from his c o l l e c t i o n w e r e figured
i n p u b l i c a t i o n s o f the O h i o a n d N e w York G e o l o g i c a l S u r v e y s a n d e l s e -
w h e r e (Hall 1872a, b; M e e k 1872a, b, 1873). In 1880 his p e r s o n a l c o l l e c t i o n ,
w h i c h w e i g h e d m o r e t h a n 17,000 p o u n d s (!), w a s sold to Harvard U n i v e r s i t y
for its M u s e u m o f C o m p a r a t i v e Z o o l o g y . " T h e a r r a n g e m e n t s for this for-
tunate d i s p o s i t i o n o f i m p o r t a n t s c i e n t i f i c m a t e r i a l a p p a r e n t l y w e r e m a d e
possible by N a t h a n i a l S o u t h g a t e S h a l e r " ( C r o n e i s 1963, 82).
However, C. B. D y e r is not o n l y k n o w n for his c o l l e c t i n g a c t i v i t i e s in
the local area. A r o u n d 1857 he b e c a m e interested in t h e b e d s at C r a w f o r d s -
ville, I n d i a n a , f a m o u s for C a r b o n i f e r o u s c r i n o i d s , a n d h e m a d e e x t e n s i v e
e x c a v a t i o n s there. H e also t r a d e d O r d o v i c i a n fossils f r o m t h e C i n c i n n a t i
region to the Hovey M u s e u m at W a b a s h C o l l e g e , in Crawfordsville. T h e
c r i n o i d c o l l e c t i o n t h e r e b y a s s e m b l e d b y D y e r w a s sold b y h i m t o t h e British
M u s e u m o f N a t u r a l History, a n d i t w a s t h e first large g r o u p o f s p e c i m e n s
f r o m C r a w f o r d s v i l l e t o b e sent a b r o a d ( V a n S a n t a n d L a n e 1964).

Science in the Hinterland 23

 T h r o u g h his w o r k w i t h S . A . M i l l e r , C . B . D y e r was i n v o l v e d i n the
n a m i n g o f m a n y taxa o f l o c a l fossils, i n c l u d i n g a n n e l i d w o r m s , b r y o z o a n s ,
s n a i l s , s p o n g e s , starfish a n d o t h e r e c h i n o d e r m s , trace fossils, a n d o t h e r s .
M o r e o v e r , at least o n e g e n u s and t w e l v e s p e c i e s of fossils w e r e n a m e d after
h i m , including the w e l l - k n o w n species of crinoids originally designated
Glyptocrinus dyeri M e e k , 1872, now a s s i g n e d to Pycnocrinus.
A c c o r d i n g to records at C i n c i n n a t i ' s Spring G r o v e C e m e t e r y , C. B.
D y e r d i e d o n July 11, 1883, i n H a r r i s o n , O h i o , n e a r C i n c i n n a t i . ( B e c k e r
1938; B y r n e s et al. 1883; C a s t e r 1982; C r o n e i s 1963; S. A. M i l l e r and D y e r
1878a, 1878b; R a y m o n d 1936; S h e r b o r n 1940; S h i d e l e r [1952] 2002.)

S. A. Miller S a m u e l A l m o n d M i l l e r ( F i g u r e 2.1B) i s c e r t a i n l y the most i m p o r t a n t o f the

" a m a t e u r s " o f t h e C i n c i n n a t i S c h o o l . H e was b o r n n e a r A t h e n s , O h i o , i n
1837. By profession he w a s a l a w y e r ; he h a d s t u d i e d at the C i n c i n n a t i L a w
C o l l e g e a n d w a s a d m i t t e d to t h e bar in 1860.
S . A . M i l l e r w a s a l s o i n v o l v e d i n p u b l i s h i n g . I n 1861-1862 h e p u b l i s h e d
t h e M a r i e t t a , O h i o , Republican, w h i c h , i n t e r e s t i n g l y e n o u g h , w a s a D e m o -
c r a t i c n e w s p a p e r . In 1874 a n d 1875, he w a s t h e p r o p r i e t o r of the Cincinnati
Quarterly journal of Science; many important papers on Cincinnatian fossils
were published in that journal. After two years or so, Miller (and L. M.
Hosea, w h o had b e c o m e co-proprietor) ceased production of the journal
alter eight n u m b e r s had a p p e a r e d . W h e n the C i n c i n n a t i Society o f N a t u -
ral History c o m m e n c e d its o w n j o u r n a l in 1878, it was rather s i m i l a r to
M i l l e r ' s d e f u n c t o n e . T h i s i s hardly s u r p r i s i n g g i v e n that M i l l e r had b e e n
a f o u n d i n g m e m b e r o f the society a n d had b e e n c a m p a i g n i n g f o r the soci-
ety t o p u b l i s h its o w n j o u r n a l ( A n o n . 1875). A s a n a c t i v e m e m b e r o f the
society, he s e r v e d at v a r i o u s t i m e s as v i c e p r e s i d e n t , p r e s i d e n t , c u r a t o r of
p a l e o n t o l o g y , a n d an e d i t o r of their j o u r n a l , in a d d i t i o n to p r e s e n t i n g pa-
p e r s a t t h e i r m e e t i n g s . S . A . M i l l e r w a s o n e o f a c o m m i t t e e o f ten e s t a b -
l i s h e d b y the society t o c o n s i d e r t h e g e o l o g i c a l n o m e n c l a t u r e o f the t y p e -
C i n c i n n a t i a n , a n d he w a s t h e first a u t h o r of their report (S. A. M i l l e r et al.
1879). A s i n d i c a t e d i n t h e s e c t i o n a b o u t C . B . D y e r a b o v e , six o f the indi-
v i d u a l s o n that c o m m i t t e e are g e n e r a l l y r e c o g n i z e d a s m e m b e r s o f t h e
C i n c i n n a t i S c h o o l , a n d all of t h e m h a v e b e e n listed in o n e p l a c e or a n o t h e r
as c o l l e c t o r s of l o c a l fossils.
M i l l e r was a c t i v e e l s e w h e r e i n t h e c o m m u n i t y , too. H e s e r v e d o n t h e
l o c a l s c h o o l b o a r d , ran for t h e U. S. S e n a t e , a n d also for c i r c u i t c o u r t j u d g e .
He did not w i n his e l e c t i o n s , h o w e v e r ; it s e e m s that he refused to take a n y
contributions.
In 1882 M i l l e r w a s c o n s i d e r e d for t h e p o s i t i o n of O h i o state g e o l o g i s t ,
to s u c c e e d John S t r o n g N e w b e r r y . Edward O r i o n actually got the job.)
M i l l e r w a s a w a r d e d a n honorary d o c t o r a t e b y O h i o U n i v e r s i t y , w h e r e , years
b e f o r e , he h a d b e e n a s t u d e n t for o n e year.
M i l l e r p r o d u c e d a great many p u b l i c a t i o n s d e v o t e d to fossils, often
co-authored with other Cincinnati-area collectors, such as C. B. Dyer and
C h a r l e s Faber. L a e l Bradshaw c o n c l u d e d that M i l l e r n a m e d oyer 1000 taxa
( B r a d s h a w , pers. c o m m . ) . But M i l l e r is p e r h a p s best k n o w n for his c o m p i l a -

24 A Sea without Fish

tions of k n o w l e d g e : 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 s u p p l e m e n t s in 1892 and 1897). ' T h e last v o l u m e listed,
according to K e n n e t h Caster, is probably the most used v o l u m e about
A m e r i c a n p a l e o n t o l o g y e v e r c o m p i l e d a n d c e r t a i n l y w a s the m o s t a m b i -
tious private p u b l i c a t i o n in p a l e o n t o l o g y ever.
M i l l e r ' s c o m p i l a t o r y works w e r e l o o k e d d o w n u p o n b y m o s t profession-
als, b u t w e r e used by t h e m n o n e t h e l e s s . C a s t e r r e c o u n t e d a story a b o u t his
professor, G . D . H a r r i s , t o t h e e f f e c t that Harris's o w n professor, H e n r y
S h a l e r W i l l i a m s , was d i s d a i n f u l o f M i l l e r ' s w o r k s . I n C a s t e r ' s w o r d s : " ' Y e t , '
said Harris, ' M i l l e r ' s g r e a t North American Geology and Paleontology w a s
a l w a y s o n W i l l i a m s ' desk, a n d o n t h e desk o f e v e r y o t h e r p a l e o n t o l o g i s t o f
the l a n d ! ' " ( C a s t e r 1982, 24).
N o r did S . A . M i l l e r c o n f i n e his w o r k t o fossils f r o m t h e C i n c i n n a t i
region. H e also w o r k e d o n t h o s e o f Illinois, M i s s o u r i , a n d W i s c o n s i n . M i l l -
er's fossil c o l l e c t i o n m u s t h a v e b e e n fantastic: o n e n e w s p a p e r a c c o u n t re-
ported that it c o n t a i n e d o v e r a m i l l i o n s p e c i m e n s ! A c c o r d i n g to B r a d s h a w ,
he rose early a n d w o r k e d on fossils u n t i l 10 AM, t h e n w e n t to his law office
until s u p p e r ; after s u p p e r he w o r k e d a c o u p l e m o r e h o u r s on fossils.
T h i s s c h e d u l e must have taken its toll, for C a s t e r r e c o r d e d that M i l l e r
was addicted to drink. A c c o r d i n g to the late W a l t e r B u c h e r , o n e - t i m e profes-
sor in the D e p a r t m e n t of G e o l o g y at the University of C i n c i n n a t i : " M i l l e r
often c a d g e d a quarter from an a d v o c a t e across the hall to b u y a shot of
b o u r b o n " (Caster 1982, 25). C a s t e r u s e d to give an e x p a n d e d version of t h e
story: w h e n thirsty for a d r i n k . M i l l e r used to go to t h e lawyer's office to bor-
row a quarter. The lawyer w o u l d take a t y p e - s p e c i m e n as collateral. T h e
lawyer w a s s u p p l i e d with m o n e y from t h e W a l k e r M u s e u m i n C h i c a g o .
Miller lived l o n g e n o u g h that every s p e c i m e n m a r k e d " t y p e " w e n t t o C h i -
c a g o . M i c h a e l S . C h a p p a r s , c u r a t o r a t the U n i v e r s i t y o f C i n c i n n a t i G e o l o g y
M u s e u m in 1936, discovered that M i l l e r had not m a r k e d all his types as s u c h .
H e n c e , t h e University o f C i n c i n n a t i got a b o u t h a l f o f Miller's t y p e s b y a c c i -
dent (Caster 1982, pers. c o m m . ) . On the o t h e r h a n d , a 1912 letter from Ray S.
Bassler to C. D. W a l c o t t said that M i l l e r "sold w h e n e v e r i m p e c u n i o u s to
G u r l e y " ( S h e r b o r n 1940, 97); a n d S h e r b o r n i n d i c a t e d that m o s t of M i l l e r s
types were at the University of C h i c a g o in t h e c o l l e c t i o n of W. K E. G u r l e y
(with w h o m M i l l e r had p u b l i s h e d a n u m b e r of scientific papers).
S . A . M i l l e r w i l l e d b o t h his c o l l e c t i o n a n d his library t o t h e U n i v e r s i t y
of C i n c i n n a t i . T h e library is intact, b u t is n o t e a s y to use b e c a u s e ". . .
M i l l e r had b o u n d i t into v o l u m e s , the o n l y c r i t e r i o n o f o r g a n i z a t i o n b e i n g
e n o u g h papers o f the s a m e p a g e - d i m e n s i o n s t o m a k e a c o n v e n i e n t l y s i z e d
v o l u m e . His interests r a n g e d w i d e l y , a n d m o s t v o l u m e s are h i g h l y e c l e c t i c "
(Caster 1982, 26).
A c c o r d i n g t o r e c o r d 61331 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 ,
w h e r e he is b u r i e d , S. A. M i l l e r d i e d on D e c e m b e r 18, 1897, of " c a n c e r of
liver and u r a e m i c p o i s o n i n g . " ( B r a d s h a w , pers. c o m m . ; B r a n d t a n d D a v i s
2007; Caster 1951,1981,1982, pers. c o m m . ; C r o n e i s 1963; C u f f e y , Davis, and
Utgaard 2002; M e r r i l l 1924; Sherborn 1940; Anon. 1875,1878.)

Science in the Hinterland 25

 By now it has b e c o m e o b v i o u s that a n u m b e r of t h r e a d s of o u r story
arc i n t r i c a t e l y i n t e r t w i n e d . V a r i o u s m e m b e r s o f t h e C i n c i n n a t i S c h o o l
h a v e tie-ins w i t h t h e C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y , with the U n i -
versity of C i n c i n n a t i , or w i t h b o t h . A n o t h e r t h r e a d in the skein is W o o d -
w a r d H i g h S c h o o l , a s w e shall see. B u t let u s follow the U n i v e r s i t y o f C i n -
c i n n a t i t h r e a d for a bit.

A. G. W e t h e r b y Albert G a l l a t i n W e t h e r b y ( f i g u r e 2.3A) w a s b o r n in Pittsburgh, Pennsylva-

n i a , in 1833, b u t his family later m o v e d to the C l e v e l a n d , O h i o , area. Alter
g r a d u a t i n g from c o l l e g e , he spent several years t e a c h i n g in a c o u n t r y s c h o o l ,
w i t h s u m m e r s spent f a r m i n g . In 1861 he m o v e d to C i n c i n n a t i and was ap-
p o i n t e d p r i n c i p a l o f W o o d b u r n S c h o o l , o n e o f the p u b l i c s c h o o l s i n the city,
a n d spent s o m e n i n e years there. In a e u l o g y written by G e o r g e W. Harper,
a n o t h e r m e m b e r of the C i n c i n n a t i S c h o o l , it is reported that Wetherby was
a p p o i n t e d professor of n a t u r a l history at the t h e n n e w University of C i n c i n -
nati in 1870 a n d stayed there six years. However, a c c o r d i n g to the University
of C i n c i n n a t i R e c o r d of M i n u t e s N o . 2, a v o l u m e in the archives of the Uni-
versity of C i n c i n n a t i , W e t h e r b y ' s t i m e at the university b e g a n in the a u t u m n
of 1877, a n d he is listed as "Ass't. P r o f A. C. W e t h e r b y of N a t u r a l History." In
January o f 1878 h e was a p p o i n t e d " C u r a t o r o f t h e M u s e u m i n the U n i v e r s i t y , "
a n d in M a r c h of that year, his title w a s c h a n g e d to professor of natural history
(Board o f D i r e c t o r s , University o f C i n c i n n a t i , 46, 65, 77, 8 0 , 1 0 0 , 1 1 7 ) . W e t h -
erby listed h i m s e l f a s " A . M . , Professor o f G e o l o g y and Z o o l o g y , U n i v e r s i t y o f
C i n c i n n a t i " ( W e t h e r b y 1880, 1881). His last entry in the c a t a l o g u e s of the
University of C i n c i n n a t i is for 1884-1885: "Albert G a l l a t i n W e t h e r b y , A . M . ,
Professor of N a t u r a l History."
W e t h e r b y left the U n i v e r s i t y of C i n c i n n a t i to p u r s u e a c a r e e r in b u s i -
ness, f i r s t w i t h the A m e r i c a n a n d E u r o p e a n I n v e s t m e n t C o m p a n y a n d ,
later, a s a m a n a g e r o f s o m e t i m b e r a n d m i n i n g lands o f the Roan M l . Steel
a n d Iron C o . in N o r t h C a r o l i n a . He d i e d on F e b r u a r y 15, 1902, in Magnetic-
City, North Carolina.
Wetherby's interests w e r e m a n y a n d v a r i e d . In a d d i t i o n to b e i n g a
s t u d e n t o f fossils, h e w a s , a t v a r i o u s t i m e s , c u r a t o r o f e n t o m o l o g y a n d c u r a -
for of c o n c h o l o g y at the C i n c i n n a t i Society of Natural History. He co-au-
t h o r e d w i t h a n o t h e r m e m b e r o f the C i n c i n n a t i S c h o o l , John M i c k l e b o r -
o u g h , a list o f t y p e - C i n c i n n a t i a n fossils ( 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). He a u t h o r e d a n u m b e r of o t h e r p a p e r s on fossils, e s p e c i a l l y , but
n o t e x c l u s i v e l y , e c h i n o d e r m s ( W e t h e r b y 1879a, 1879b, 1880, 1881). In the
first-cited of t h o s e , he n a m e d the g e n u s Enoploura, and interpreted the
a n i m a l s o f that g e n u s t o b e c r u s t a c e a n s . W e t h e r b y ' s failure t o r e c o g n i z e
that h e w a s d e a l i n g not w i t h c r u s t a c e a n s , but w i t h e c h i n o d e r m s , b r o u g h t
d o w n o n h i m t h e w r a t h o f H e n r y W o o d w a r d o f the British M u s e u m o f
Natural History. His contributions as a m e m b e r of the C i n c i n n a t i School
o f P a l e o n t o l o g y n o t w i t h s t a n d i n g , m o s t o f W e t h e r b y ' s p u b l i c a t i o n s are not
a b o u t p a l e o n t o l o g y , b u t rather a b o u t p r e s e n t - d a y m o l l u s c s .
W e t h e r b y i s o n e o f m a n y e x a m p l e s o f the f a c t that the lives a n d careers
of the m e m b e r s of the C i n c i n n a t i School were intertwined, f o r example,

26 A Sea without Fish

John M . N i c k l e s s t u d i e d u n d e r W e t h e r b y a t t h e U n i v e r s i t y o f C i n c i n n a t i ,
and G e o r g e W . H a r p e r w r o t e a e u l o g y a b o u t W e t h e r b y . W e t h e r b y w a s o n e
o f a c o m m i t t e e o f ten w h o w r o t e a r e p o r t o n t h e g e o l o g i c a l n o m e n c l a t u r e
o f the t y p e - C i n c i n n a t i a n (S. A . M i l l e r et al. 1879); six o f t h e i n d i v i d u a l s o n
that c o m m i t t e e are g e n e r a l l y r e c o g n i z e d a s m e m b e r s o f t h e C i n c i n n a t i
S c h o o l , a n d all o f t h e m h a v e b e e n listed i n o n e p l a c e o r a n o t h e r a s c o l l e c -
tors o f l o c a l fossils. ( B r a n d 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 . M i l l e r 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; Nickles 1956; W e t h e r b y 1879a, 1879b, 1880, 1881; A n o n . 1876, 1878,
1879.)

John M i c k l e b o r o u g h , P h . D . , w a s t h e p r i n c i p a l o f t h e C i n c i n n a t i N o r m a l John
S c h o o l from 1878 until 1885. T h i s s c h o o l w a s a part of t h e C i n c i n n a t i p u b - Mickleborough
lie school system that w a s d e d i c a t e d t o t r a i n i n g t e a c h e r s . T h e C i n c i n n a t i
Board of Education suspended the operation of the N o r m a l S c h o o l in
1900, but that w a s a d e c a d e a n d a h a l f after D r . M i c k l e b o r o u g h h a d d e -
parted C i n c i n n a t i for N e w York, w h e r e h e b e c a m e t h e p r i n c i p a l o f t h e
Boys High School in Brooklyn.
M i c k l e b o r o u g h h a d b e e n n o m i n a t e d for m e m b e r s h i p i n t h e C i n c i n -
nati S o c i e t y of N a t u r a l History in July of 1876 ( C i n c i n n a t i S o c i e t y of N a t u -
ral History, 90), and he b e c a m e an a c t i v e m e m b e r of t h e s o c i e t y ; for ex-
a m p l e , he served on t h e P u b l i c a t i o n s C o m m i t t e e a n d as a m e m b e r of a
c o m m i t t e e 011 the n o m e n c l a t u r e of t h e r o c k s of t h e t y p e - C i n c i n n a t i a n that
i n c l u d e d five o t h e r i n d i v i d u a l s g e n e r a l l y r e c o g n i z e d a s m e m b e r s o f t h e
C i n c i n n a t i S c h o o l (S. A . M i l l e r et a l . 1879). A n d as n o t e d a b o v e , in t h e
section a b o u t A . G . W e t h e r b y , M i c k l e b o r o u g h a n d W e t h e r b y c o - a u t h o r e d
a n important list o f t y p e - C i n c i n n a t i a n fossils ( M i c k l e b o r o u g h a n d W e t h -
erbv 1878a, b). B u t his m o s t s i g n i f i c a n t p u b l i c a t i o n is h i s 1883 paper on
trilobites, w h i c h i n c l u d e s a d e s c r i p t i o n of a s p e c i m e n of Isotelus f r o m the
t y p e - C i n c i n n a t i a n w i t h p r e s e r v e d a p p e n d a g e s . M i c k l e b o r o u g h w a s rather
a h e a d of his t i m e s in his r e a l i z a t i o n that t h e a p p e n d a g e s of trilobites are
s i m i l a r to those of present-day c h e l i c e r a t e a r t h r o p o d s .
In addition to a u t h o r i n g p u b l i c a t i o n s on fossils, D r . M i c k l e b o r o u g h , as
a professional educator, also wrote in the field of e d u c a t i o n , for e x a m p l e , on
a m e t h o d of t e a c h i n g addition and subtraction in the p r i m a r y grades that was
p r o m o t e d by John B. Peaslee, s u p e r i n t e n d e n t of s c h o o l s in C i n c i n n a t i , a n d
c a l l e d b y Peaslee " T h e l e n s M e t h o d " and b y M i c k l e b o r o u g h " T h e Peaslee
M e t h o d . " (Bassler 1947; Brandt a n d D a v i s 2007; C a s t e r 1982; Lathrop 1900,
1902; M i c k l e b o r o u g h 1883; N i c k l e s 1936; S h o t w e l l 1902; V e n a b l e 1894; A n o n .
1886.)

A l t h o u g h the c a p t i o n o f his p h o t o g r a p h i n the D e p a r t m e n t o f G e o l o g y a t Charles Faber

the University o f C i n c i n n a t i i n d i c a t e s that C h a r l e s F a b e r ( F i g u r e 2.1C) w a s
a realtor, K e n n e t h C a s t e r (1982) c l a i m e d that he w a s a m a n u f a c t u r e r o f
l e a t h e r b e l t i n g . But b o t h s o u r c e s r e c o g n i z e d h i m as a fossil c o l l e c t o r . In
the 1880s a n d 1890s his n a m e a p p e a r e d as a n a u t h o r o f r e c o r d , b o t h a l o n e

Science in the Hinterland 27

 and as a co-author with S. A. Miller. As s u c h , he was involved in the n a m -
i n g o f a n u m b e r o f taxa o f fossils f r o m t h e t y p e - C i n c i n n a t i a n .
A c c o r d i n g t o t h e s a m e p h o t o g r a p h c a p t i o n , F a b e r lived until 1930, late
e n o u g h that Shideler w e n t c o l l e c t i n g with h i m . A c c o r d i n g to Shideler
([1952] 2002, 3), F a b e r w a s , ". . . like t h e t y p i c a l old t i m e r he was[,] very se-
c r e t i v e a n d s u s p i c i o u s . H e w a s n ' t t e l l i n g a n y b o d y a n y t h i n g . I t took m e t w o
years t o g e t h i m s o f t e n e d u p a n d e d u c a t e d s o that h e w a s w i l l i n g t o c o m e
o u t w i t h his i n f o r m a t i o n . S o w e started g o i n g a r o u n d t o a n u m b e r o f t h e
old s e c r e t l o c a l i t i e s w h e r e S . A . M i l l e r got his t y p e s . "
Like most of the other m e m b e r s of the C i n c i n n a t i S c h o o l , Faber was
a s s o c i a t e d w i t h t h e C i n c i n n a t i Society o f N a t u r a l H i s t o r y . I n fact, h e was
p r o p o s e d for m e m b e r s h i p in the society- in 1885, at the s a m e t i m e as C h a r l e s
S c h u c h e r t and Ernst Vaupel, and he was duly elected.
F a b e r sold his o r i g i n a l c o l l e c t i o n t o t h e U n i v e r s i t y o f C h i c a g o for
$5000, a c c o r d i n g to S h i d e l e r , a n d it i n c l u d e d s p e c i m e n s d e s c r i b e d by S. A.
M i l l e r . B e i n g a n inveterate f o s s i l c o l l e c t o r , h o w e v e r , h e p r o c e e d e d t o a m a s s
a second collection. This o n e w a s b e q u e a t h e d t o t h e U n i v e r s i t y o f C i n c i n -
nati. T h e c o l l e c t i o n c a m e w i t h s o m e m o n e y to p r o v i d e for a curatorial posi-
t i o n a n d for p a l e o n t o l o g i c a l p u b l i c a t i o n s . The first holder of the curatorial
p o s i t i o n w a s C a r r o l l L a n e F e n t o n , w h o w e n t o n t o write, a l o n g w i t h his
w i f e , w h a t is arguably t h e b e s t b o o k of its t i m e for a m a t e u r fossil c o l l e c t o r s
( F e n t o n a n d F e n t o n 1958). T h e late K e n n e t h E. C a s t e r w a s also a w e l l -
k n o w n F a b e r c u r a t o r . (Bassler 1947; B e c k e r 1938; C a s t e r 1982; Faber 1886,
1929; S. A. M i l l e r a n d F a b e r 1892a, b, 1894a, b; S h i d e l e r [1952I 2002; A n o n .
1885a, b.)

D. T. D. Dyche D r . D y c h e , o f L e b a n o n , O h i o , i s o n e o f t h e less w e l l - k n o w n m e m b e r s o f
t h e C i n c i n n a t i S c h o o l . H e a u t h o r e d several p a p e r s o n fossil c r i n o i d s o f the
t y p e - C i n c i n n a t i a n . In n a m i n g a s p e c i e s of c o n o d o n t , Prioniodus dychei, U.
P . J a m e s h o n o r e d D r . D y c h e a s o n e " w h o has d o n e s o m u c h i n c o l l e c t i n g
a n d d e v e l o p i n g s o many o f t h e finest C r i n o i d s , etc., f o u n d i n the C i n c i n -
nati G r o u p . . . " (1884c, 1 4 7 - 1 4 8 ) . T h e r e is, i n t h e W a r r e n C o u n t y Historical
M u s e u m , in L e b a n o n , O h i o , what is labeled as the dental cabinet of David
T u l l i s D u r b i n D y c h e . W e h a v e n o t b e e n a b l e t o verify w h e t h e r D . T . D .
D y c h e , the m e m b e r of the C i n c i n n a t i S c h o o l , is the same person as David
Tullis D u r b i n D y c h e , t h e dentist. ( B e c k e r 1938; D y c h e 1892a, b, c; U. P.
J a m e s 1884c.)

E. O. Ulrich E d w a r d O . U l r i c h ( F i g u r e s 2 . 3 C , 2.4B, 2.5D) w a s b o r n F e b r u a r y 1,1857, i n

C i n c i n n a t i , b u t shortly t h e r e a f t e r t h e family m o v e d t o C o v i n g t o n , K e n -
t u c k y , just across t h e O h i o River. T h e "O" stands for " O s c a r , but it was not
a n a m e g i v e n t o h i m b y his p a r e n t s ; E d w a r d U l r i c h g a v e h i m s e l f that n a m e
after a h e r o in o n e of t h e stories he read as a boy. He s e e m s to h a v e b e e n a
sickly- c h i l d , a n d w a s f r e q u e n t l y absent from s c h o o l . He w a s i n t r o d u c e d to
fossils b y h i s m i n i s t e r , t h e R e v e r e n d H e n r y H e r z e r , w h e n h e w a s s e v e n
years old.

28 A Sea without Fish

 After q u i t t i n g s c h o o l , he w a s a s u r v e y o r for a c o u p l e of y e a r s a n d
worked o n t h e E d e n Park R e s e r v o i r , w h i c h , t o this d a y , s u p p l i e s d r i n k i n g
water t o d o w n t o w n C i n c i n n a t i . H e w a s a s t u d e n t a t B a l d w i n - W a l l a c e C o l -
lege for t w o years, hut he did not finish c o l l e g e . D u r i n g t h e 18761877
school year, he was a student in the M e d i c a l C o l l e g e of O h i o in C i n c i n -
nati, an i n d e p e n d e n t institution at that t i m e , b u t a b s o r b e d into t h e U n i v e r -
sity o f C i n c i n n a t i i n 1915 ( B r o a d d u s , pers. c o m m . ) . A g a i n , h e did n o t f i n i s h
work f o r a d e g r e e . F o r m a l e d u c a t i o n a n d h e did n o t g e t o n t o o w e l l , b e -
c a u s e " h e insisted h e was t a u g h t too m u c h h e d i d n ' t w a n t a n d too little that
he d i d " (Bassler 1945, 333).
In 1876, U l r i c h w a s e l e c t e d to m e m b e r s h i p in t h e C i n c i n n a t i S o c i e t y
o f N a t u r a l H i s t o r y . The f o l l o w i n g y e a r h e w a s e l e c t e d c u r a t o r o f p a l e o n t o l -
ogy, a n u n p a i d position. A b o u t that t i m e t h e s o c i e t y a c q u i r e d its o w n b u i l d -
i n g , a n d , in the m i n u t e s of t h e society- for t h e first m e e t i n g h e l d in t h e new
b u i l d 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 N a t u r a l H i s t o r y ) , it
is r e c o r d e d :
" T h e matter of a p p o i n t i n g a janitor for the B u i l d i n g c o m i n g u p , proposi-
tions w e r e r e c e i v e d from Messrs. F. O. Ulrich, Talbot, a n d J. C. S h o r t e n .
"Professor Wetherby m o v e d that the Society p r o c e e d to b a l l o t for a jani-
for, the person e l e c t e d to be subject to s u c h rules as the S o c i e t y m a y a d o p t ,
a g r e e d to. The ballot resulted as follows: Mr. U l r i c h r e c e i v e d 28 votes. M r .
S h o r t e n r e c e i v e d 7 votes. M r . Talbot r e c e i v e d 3 votes a n d t h e r e u p o n Mr.
E. O. U l r i c h was d e c l a r e d e l e c t e d . M r . U l r i c h ' s proposition is as follows,

Cincinnati Nov. 6th 1877

To the Cincinnati Society of Natural History

T h e undersigned is an applicant for the position of Janitor or Custodian of

the Society's building; am willing to devote my entire time to the interests of
the Society, for the consideration of $300.00 per a n n u m , and the Society to
allow me one room for a sleeping apartment.

Respectfully yours,

E. O. Ulrich

Ulrich's association w i t h the C i n c i n n a t i Society o f N a t u r a l History-

brought h i m into c o n t a c t with U.P. James, Joseph F. James, C h a r l e s B. Dyer,
S . A . Miller, and o t h e r m e m b e r s o f t h e C i n c i n n a t i S c h o o l . For e x a m p l e ,
U l r i c h w a s o n e o f six m e m b e r s o f the C i n c i n n a t i S c h o o l w h o served o n a
c o m m i t t e e o n the n o m e n c l a t u r e o f the rocks o f t h e t y p e - C i n c i n n a t i a n (S. A .
Miller e t a l . 1 8 7 9 ) . N o r did his c o n t a c t s c o m e o n l y w i t h C i n c i n n a t i folk. For
e x a m p l e , as early as 1886, Ulrich w e n t c o l l e c t i n g with A u g u s t F Foerste, w h o
went on to b e c o m e o n e of the foremost workers on fossil c e p h a l o p o d s in t h e
United States. A l t h o u g h the position at the s o c i e t y w a s c a l l e d " c u s t o d i a n , "
U l r i c h apparently was in c h a r g e of day-to-day o p e r a t i o n s at their facility.
In addition to the labors a s s o c i a t e d w i t h that j o b , a n d later o n , U l r i c h
w o r k e d at various t i m e s as a c a r p e n t e r , for v a r i o u s state g e o l o g i c a l s u r v e y s ,
a n d p a r t - t i m e for the U n i t e d States G e o l o g i c a l S u r v e y i n T e n n e s s e e . O n e
o f his m a i n s o u r c e s o f i n c o m e , h o w e v e r , was the p r o d u c t i o n o f t h i n - s e e -

Science in the Hinterland 29

 t i o n s of b r y o z o a n s , w h i c h he sold to b u y e r s b o t h in the U n i t e d States and
E u r o p e . I n o r d e r t o c o l l e c t sufficient s p e c i m e n s and m a k e t h i n - s e c t i o n s
f r o m t h e m , U l r i c h e m p l o y e d o t h e r l o c a l a f i c i o n a d o s o f fossils, i n c l u d i n g
Bassler, N i c k l e s , a n d S e h u c h e r t . ( K e n n e t h C a s t e r [1982] has c r e d i t e d U l -
rich w i t h the trait o f e n l i s t i n g the a s s i s t a n c e o f l o c a l y o u t h s , t h e r e i n c h a n g -
i n g their lives. This c a l l s u p t h e i m a g e o f t h e k i n d l y old m a n h e l p i n g t h e
l o c a l kids; i t h a p p e n s , t h o u g h , that t w o o f U l r i c h ' s t h r e e b e s t - k n o w n prote-
g e s , S e h u c h e r t a n d N i c k l e s , w e r e o n l y o n e a n d t w o years y o u n g e r t h a n
U l r i c h , respectively.)
In 1897, U l r i c h w a s h i r e d p e r m a n e n t l y by the United States G e o l o g i c a l
S u r v e y a n d stayed t h e r e for t h e rest o f his c a r e e r , e v e n t u a l l y b e c o m i n g the
h e a d of t h e s t r a t i g r a p h i c s e c t i o n , a n d in effect the arbiter of stratigraphic
decisions in the country. A l t h o u g h s p e c i m e n s and thin-sections provided
by Ulrich are p r e s e n t in many i n s t i t u t i o n s all a c r o s s t h e l a n d , his p e r s o n a l
c o l l e c t i o n w e n t m a i n l y t o the U n i t e d States G e o l o g i c a l S u r v e y a n d t h e n c e
the U n i t e d States N a t i o n a l M u s e u m . U l r i c h officially retired in 1932, but
c o n t i n u e d s c h o l a r l y work as an honorary a s s o c i a t e in p a l e o n t o l o g y at the
S m i t h s o n i a n Institution.
U l r i c h a u t h o r e d or c o - a u t h o r e d many taxa of a n i m a l s of all kinds. Onc-
o l t h e s e w a s a s p e c i e s o f o s t r a c o d c r u s t a c e a n n a m e d alter the m a n o f the
c l o t h w h o h a d i n t r o d u c e d h i m to fossils. U l r i c h w r o t e : "I n a m e it after Rev.
H . H e r z e r , n o w o f B e r e a , O . , w h o w a s the f i r s t t o a w a k e n i n m e t h e latent
love for n a t u r e that has s i n c e g r o w n a l m o s t to a p a s s i o n , a n d b e c o m e an
i n e x h a u s t i b l e s o u r c e o f k e e n e s t e n j o y m e n t " ( U l r i c h 1891, 209).
Ulrich received main honors. Baldwin-Wallace College, which he
h a d a t t e n d e d for a t i m e , a w a r d e d h i m b o t h a n h o n o r a r y master's d e g r e e
a n d an honorary d o c t o r a l d e g r e e in 1886 a n d 1892, respectively. He w a s a
m e m b e r o f t h e N a t i o n a l A c a d e m y o f S c i e n c e s a n d was a w a r d e d their Mary-
C l a r k T h o m p s o n M e d a l . He was elected president of the Paleontological
S o c i e t y for t h e y e a r 1915; this society w a s t h e n a n d is n o w the p r e m i e r p r o -
fessional p a l e o n t o l o g i c a l o r g a n i z a t i o n in t h e U n i t e d States. In 1932, U l r i c h
w a s a w a r d e d t h e P e n r o s e M e d a l o f t h e G e o l o g i c a l Society o f A m e r i c a , the
h i g h e s t h o n o r to w h i c h a g e o l o g i s t may aspire.
O n t h e o t h e r h a n d , U l r i c h s e e m s t o h a v e b e e n i n v o l v e d i n s o m e less-
i h a n - h o n o r a b l e activities. T h e m o s t s u r p r i s i n g i s that h e b a c k e d a n attempt
t o p r e v e n t t h e e l e c t i o n o f C h a r l e s S e h u c h e r t t o t h e presidency o f the G e o -
l o g i c a l S o c i e t y o f A m e r i c a , d e s p i t e t h e fact that S e h u c h e r t h a d b e e n a
p r o t e g e a n d c o l l e a g u e o f U l r i c h f r o m the old C i n c i n n a t i days. T h e proxi-
m a t e c a u s e f o r this w a s that U l r i c h h a d p r o p o s e d r e c o g n i t i o n o f t w o major
c h u n k s of rock b e t w e e n the C a m b r i a n and the O r d o v i c i a n Systems, the
Ozarkian and C a n a d i a n . H o w e v e r , S e h u c h e r t did n o t e n t h u s i a s t i c a l l y
a d o p t the O z a r k i a n a n d the C a n a d i a n S y s t e m s i n t h e p r e s t i g i o u s t e x t b o o k
o n h i s t o r i c a l g e o l o g y o f w h i c h h e w a s c o - a u t h o r . T h e r e also w a s a dispute-
as to w h e t h e r U l r i c h or S e h u c h e r t had " i n v e n t e d " the p a l e o g e o g r a p h i c
map (neither had, but it certainly was Sehuchert who made them
famous).
T h e u l t i m a t e c a u s e , h o w e v e r , w a s t h e fact that U l r i c h w a s extraordi-
narily k n o w l e d g e a b l e o n all t h i n g s s t r a t i g r a p h i c a n d p a l e o n t o l o g i c . F r o m

30 A Sea without Fish

c h i l d h o o d , h e had had a n u n c a n n y m e m o r y . K e n n e t h C a s t e r (pers. c o m m . )
c l a i m e d that, d e c a d e s after v i s i t i n g a l o c a l i t y , U l r i c h c o u l d r e c a l l t h e strati-
g r a p h i c s e c t i o n t h e r e i n c h b y i n c h , w i t h great a c c u r a c y a n d p r e c i s i o n ,
a l o n g with its fossil c o n t e n t s . Thus, Ulrich was supremely self-confident.
I n d i s c u s s i o n s , h e " t o p u t i t m i l d l y , m a d e his p o s i t i o n c l e a r . " H e " w a s i n
c o n g e n i t a l d i s a g r e e m e n t u n t i l t h e day o f his d e a t h o n W a s h i n g t o n ' s B i r t h -
day, 1944" ( C r o n e i s 1963,85). H o w e v e r , o n e s h o u l d n o t c o n j u r e u p a p i c t u r e
o f a g r u m p y old m a n . U l r i c h w a s k n o w n for his g e n i a l d i s p o s i t i o n , a n d ,
i n d e e d , w a s k n o w n a s " U n c l e H a p p y . " (As i t t u r n e d o u t , U l r i c h ' s " n e w "
s y s t e m s n e v e r did g a i n w i d e s p r e a d a c c e p t a n c e , b u t i n t h e m e a n t i m e ,
S c h u c h e r t did get elected.)
U l r i c h was a n early worker o n P a l e o z o i c o s t r a c o d e s a n d c o n o d o n t s . H e
i s e s p e c i a l l y well k n o w n for his w o r k w i t h b r y o z o a n s , h a v i n g b e e n o n e o f
t h e pioneers i n the use o f t h i n - s e c t i o n s t o u n d e r s t a n d t h e a n i m a l s . H o w -
ever, his e x t e n s i v e record o f p u b l i c a t i o n s i n c l u d e s s c i e n t i f i c p a p e r s o n r e p -
resentatives o f a l m o s t e v e r y m a j o r g r o u p o f i n v e r t e b r a t e s ; n o t o n l y t h e o n e s
m e n t i o n e d a b o v e , but also, a n n e l i d w o r m s , b r a c h i o p o d s , m o l l u s c s , s p o n g e s ,
a n d trilobites. (Bassler 1945; B e c k e r 1938; B r a d s h a w 1989; B r a n d t a n d D a v i s
2007; Byers 2001; C a s t e r 1951,1981, 1982; C r o n e i s 1963; C u f f e y , D a v i s , a n d
U t g a a r d 2002; M e r r i l l 1924; S h e r b o r n 1940; S h i d e l e r [1952] 2002; U l r i c h
1891. U l r i c h ' s p e r s o n a l b i b l i o g r a p h y is i m m e n s e ; a g o o d p l a c e to start a
s e a r c h for his works is Bassler 1945.)

C h a r l e s S c h u c h e r t ( F i g u r e 2.5A) w a s b o r n in 1858, t h e s o n o f a c a b i n e t Charles Schuchert

Charles
m a k e r ; the f a m i l y was p o o r , a n d Karl (as he w a s c h r i s t e n e d ) s p e n t his life
well into his t w e n t i e s t r y i n g t o k e e p b o d y a n d s o u l t o g e t h e r . H e a t t e n d e d
s c h o o l t h r o u g h t h e sixth g r a d e , t h e n , a t a g e t w e l v e , w e n t t o a m e r c a n t i l e
s c h o o l to learn b o o k k e e p i n g , at w h i c h p o i n t he b e g a n to w o r k at his father's
f u r n i t u r e factory. H o w e v e r , t h e factory b u r n e d d o w n i n 1877; C h a r l e s re-
v i v e d the enterprise, b u t i t b u r n e d a g a i n i n 1884. M e a n w h i l e , S c h u c h e r t
did take s o m e d r a w i n g c o u r s e s a t t h e O h i o M e c h a n i c s Institute i n C i n c i n -
nati, and he mastered lithography.
S c h u c h e r t ' s i n t r o d u c t i o n to fossils c a m e in 1866, w h e n a l a b o r e r w o r k -
i n g n e a r t h e S c h u c h e r t h o m e tossed t h e e i g h t - y e a r - o l d lad a fossil that h a d
c o m e o u t o f t h e e x c a v a t i o n . S o m e t i m e t h e r e a f t e r , S c h u c h e r t ' s father took
h i m t o see t h e r o o m f u l o f fossils o w n e d b y o n e W i l l i a m F o s t e r " w h i c h
o p e n e d t o m e a n u n k n o w n w o r l d " ( B e c k e r 1938, 193). T h e b o y w a s c o m -
pletely h o o k e d . T h e n , at a b o u t a g e s e v e n t e e n , S c h u c h e r t saw t h e fossils in
the w i n d o w s of t h e e s t a b l i s h m e n t of U. P. James. They m e t , a n d t h e e l d e r
James used s o m e of the y o u n g man's fossils in his p u b l i s h e d work. S c h u c h e r t
also heard a b o u t C B. D y e r ' s c o l l e c t i o n a n d s o u g h t o u t his a c q u a i n t a n c e .
In 1877, he m e t U l r i c h , " w h o w a s to t u r n me f r o m an a m a t e u r into a profes-
sional p a l e o n t o l o g i s t " ( B e c k e r 1938,193).
As a serious a m a t e u r fossil c o l l e c t o r , a n d as o n e t r a i n e d in l i t h o g r a p h y ,
S c h u c h e r t w a s h i r e d b y U l r i c h a s t h e ideal assistant for p r e p a r i n g l i t h o -
g r a p h s for t h e Illinois a n d M i n n e s o t a g e o l o g i c a l s u r v e y s . T h i s w a s 1885
t h r o u g h 1888.

Science in the Hinterland 31

 I n that last year, J a m e s H a l l , t h e clean o f A m e r i c a n p a l e o n t o l o g i s t s ,
visited C i n c i n n a t i a n d w a s s o i m p r e s s e d b y S e h u c h e r t ' s c o l l e c t i o n that h e
h i r e d h i m t o b e c o m e a n assistant for t h e N e w York G e o l o g i c a l S u r v e y (and,
i n c i d e n t a l l y , o b t a i n e d his c o l l e c t i o n o f fossils). I n 1893, S e h u c h e r t j o i n e d
t h e U n i t e d S t a t e s G e o l o g i c a l S u r v e y , a n d , a y e a r later, h e w e n t t o the
U n i t e d States N a t i o n a l M u s e u m , also i n W a s h i n g t o n , D . C . E v e n t u a l l y , h e
c a m e t o o c c u p y t h e m o s t p r e s t i g i o u s g e o l o g i c a l professorship i n N o r t h
A m e r i c a , that a t Y a l e U n i v e r s i t y .
S e h u c h e r t b e g a n a t t e n d i n g m e e t i n g s of the C i n c i n n a t i Society of
N a t u r a l History- in 1878. H o w e v e r , it w a s not u n t i l 1885, after he had left the
f u r n i t u r e b u s i n e s s for g o o d , that he f o r m a l l y w a s p r o p o s e d as a m e m b e r .
A s i t h a p p e n s , C h a r l e s F a b e r , E r n s t V a u p e l , a n d S e h u c h e r t all w e r e n o m i -
nated at the same time.
S e h u c h e r t w a s b o r n in C i n c i n n a t i , but in s o m e respects it is not fair to
c a l l h i m a m e m b e r o f t h e C i n c i n n a t i S c h o o l , b e c a u s e h e did not really p u b -
lish a n y t h i n g locally. H e h a d 234 scientific p u b l i c a t i o n s , b u t n o n e a p p e a r e d
in the local journals. A l t h o u g h he did not invent the p a l e o g c o g r a p h i c m a p ,
he b r o u g h t it to its m a t u r e state. A n d , up until the t i m e of his d e a t h , in 1942,
h e w a s also the f o r e m o s t authority o n fossil b r a c h i o p o d s o f N o r t h A m e r i c a .
L i k e s o m e o t h e r m e m b e r s o f the C i n c i n n a t i S c h o o l , S e h u c h e r t w a s
h o n o r e d d u r i n g his l i f e t i m e . H e w a s a w a r d e d a n h o n o r a r y master's d e g r e e
by Y a l e U n i v e r s i t y in 1904, a n d honorary- d o c t o r a t e s w e r e a w a r d e d h i m by-
N e w York, Harvard, and Yale Universities. He was a m e m b e r of the Na-
tional A c a d e m y of S c i e n c e s , and was elected president both of the Paleon-
tological Society and of the G e o l o g i c a l Society of A m e r i c a , l a k e Ulrich
b e f o r e h i m , S e h u c h e r t w a s a w a r d e d t h e p r e s t i g i o u s P e n r o s e M e d a l o f the
G e o l o g i c a l S o c i e t y o f A m e r i c a i n 1934. Moreover, one of the medals
a w a r d e d b y t h e P a l e o n t o l o g i c a l S o c i e t y b e a r s h i s n a m e . N o t bad for a kid
w h o n e v e r m a d e i t t o h i g h s c h o o l ! (Bassler 1945; B e c k e r 1938; B r a n d t a n d
D a v i s 2007; Byers 2001; C a s t e r 1951, 1981, 1982; C l a r k 1943; C r o n e i s 1963;
C u f f e y , D a v i s , a n d U t g a a r d 2002; D u n b a r 1943; K a e s l e r 1987; K n o p f 1952;
S h i d e l e r [1952] 2002; T w e n h o f e l 1942; Y o c h e l s o n 1973, 1975; A n o n . 1885a.)

John M. Nickles John M i l t o n N i c k l e s ( F i g u r e s 2.4A, 2.5B), on the o t h e r h a n d , clearly was a

m e m b e r o f t h e C i n c i n n a t i S c h o o l . I n d e e d , h e w a s t h e first o f the C i n c i n -
nati S c h o o l actually a s s o c i a t e d w i t h t h e University o f C i n c i n n a t i ; h e re-
c e i v e d a b a c h e l o r ' s d e g r e e in 1882 a n d a master's d e g r e e in 1891. W h i l e at
the university, he studied g e o l o g y u n d e r A. G. Wetherby.
He had attended W o o d w a r d High S c h o o l , in C i n c i n n a t i , and had
b e e n e n c o u r a g e d i n his g e o l o g i c a l interests b y G e o r g e W . H a r p e r , t h e
principal, and by a fellow student. " M y b o y h o o d c o m p a n i o n , Ernst H.
V a u p e l , i n d u c t e d m e into c o l l e c t i n g fossils d u r i n g m y s e c o n d y e a r a t W o o d -
w a r d H i g h S c h o o l . P r e v i o u s l y w e h a d t o g e t h e r c o l l e c t e d snail shells a n d
fresh w a t e r m u s s e l s f r o m t h e O h i o R i v e r a t low water a n d t h e n ' d e e r h o r n s '
(worn c y a t h o p h y l l o i d corals) f r o m t h e drift m a t e r i a l f i l l o f t h e M a r i e t t a a n d
C i n c i n n a t i Railroad (now B & O) m a d e t h r o u g h M i l l C r e e k valley" (Nick-
les 1936).

32 A Sea without Fish

 By t h e t i m e he g r a d u a t e d f r o m h i g h s c h o o l in 1878, he h a d started a
b i b l i o g r a p h i c work o n t h e l o c a l b r y o z o a n s , a n d h e a l r e a d y w a s a c q u a i n t e d
p e r s o n a l l y w i t h U l r i c h a n d S c h u c h e r t . A f t e r stints o f t e a c h i n g i n A r k a n s a s
and then Illinois, w h e r e h e w a s a h i g h s c h o o l p r i n c i p a l , h e r e t u r n e d t o
C i n c i n n a t i , a l t h o u g h for a n u m b e r o f s u m m e r s h e h a d s p e n t v a c a t i o n s w i t h
U l r i c h c o l l e c t i n g b r y o z o a n s all o v e r c e n t r a l a n d e a s t e r n N o r t h A m e r i c a .
I n 1899 N i c k l e s m e t R a y S . Bassler a t t h e r e s i d e n c e o f E . O . U l r i c h i n
N e w p o r t , K e n t u c k y . N i c k l e s a n d Bassler c o l l a b o r a t e d t o p r o d u c e U n i t e d
States G e o l o g i c a l S u r v e y B u l l e t i n 1 7 3 S y n o p s i s of American Fossil Bryo-
zoa ( N i c k l e s a n d Bassler 1900). A b o u t t h e s a m e t i m e , Josua L i n d a h l , t h e
d i r e c t o r o f the C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y a n d f o r m e r state g e -
ologist o f Illinois, asked N i c k l e s t o p r e p a r e a p a p e r o n t h e g e o l o g y o f C i n -
c i n n a t i ; this was p u b l i s h e d in t h e s o c i e t y ' s j o u r n a l in 1902 a n d is u s e d to
this day. In the s u m m e r of 1909, N i c k l e s p r e p a r e d a m a n u s c r i p t g e o l o g i c
m a p o f the W e s t C i n c i n n a t i Q u a d r a n g l e for t h e p r o p o s e d C i n c i n n a t i F o l i o
o f the U n i t e d States G e o l o g i c a l S u r v e y ; this h a s y e t t o b e p u b l i s h e d .
I n 1903 N i c k l e s w a s a p p o i n t e d t o t h e U n i t e d States G e o l o g i c a l S u r v e y
in W a s h i n g t o n , D.C., apparently on the strength of the bryozoan bibliog-
raphy that h a d a p p e a r e d i n 1900; o f c o u r s e , h i s f r i e n d s h i p w i t h U l r i c h did
not hurt. Until his d e a t h in 1945, he d e v o t e d h i m s e l f to c o m p i l i n g b i b l i o g -
raphies, including the Bibliography of North American Geology, t h e An-
notated Bibliography of Economic Geology, and the Bibliography and Index
of Geology Exclusive of North America, w h i c h w e r e p u b l i s h e d o v e r a num-
ber of years. In all, his c o n t r i b u t i o n to t h e s e a m o u n t e d to t h i r t y - e i g h t v o l -
u m e s , c o m p r i s i n g a total o f 14,361 p a g e s a f a n t a s t i c a c c o m p l i s h m e n t !
(And i t all w a s d o n e " t h e o l d - f a s h i o n e d w a y " t h e r e w e r e n o c o m p u t e r s i n
that far-distant day a n d age!)
Nickles authored a respectable pile of publications on b r y o z o a n s , but
h e deliberately s a c r i f i c e d t h e p a l e o n t o l o g i c a l r e p u t a t i o n that w o u l d h a v e
b e e n his in order to serve t h e s c i e n c e of g e o l o g y in t h e t h a n k l e s s task of
c o m p i l i n g the b i b l i o g r a p h i e s , a n d it is for his b i b l i o g r a p h i e s that t h e g e o -
l o g i c a l c o m m u n i t y forever w i l l b e i n d e b t e d t o h i m . (Bassler 1947; B r a n d t
a n d D a v i s 2007; C a s t e r 1951, 1981, 1982; C r o n e i s 1963; C u f f e y , D a v i s , a n d
U t g a a r d 2002; N i c k l e s 1902,1936.)

G e o r g e W . H a r p e r ( F i g u r e 2.3B) w a s a b r i l l i a n t p e d a g o g u e a n d a m a t e u r G e o r g e W. Harper
g e o l o g i s t . A c t u a l l y , c a l l i n g h i m a g e o l o g i s t is t o o n a r r o w an a s s e s s m e n t , for
he was, at one time, the curator of e n t o m o l o g y at the C i n c i n n a t i Society
o f N a t u r a l History a n d , a l a n o t h e r , c u r a t o r o f m e t e o r o l o g y , a n d his n a m e
i s associated w i t h t h e s t u d y o f f r e s h w a t e r m u s s e l s i n t h e C i n c i n n a t i a r e a .
H a r p e r w a s b o r n in F r a n k l i n , O h i o , in 1832, b u t s p e n t t h e vast m a j o r i t y
o f his life i n C i n c i n n a t i . H e g r a d u a t e d f r o m W o o d w a r d C o l l e g e , i n C i n c i n -
nati, i n 1853, v a l e d i c t o r i a n o f his c l a s s , a n d b e g a n t e a c h i n g a t W o o d w a r d .
H e w a s p r i n c i p a l o f W o o d w a r d H i g h S c h o o l f r o m 1865 t h r o u g h 1900.
S o m e w h e r e a l o n g the line he w a s a w a r d e d a master's d e g r e e f r o m D e n i s o n
C o l l e g e (now, D e n i s o n U n i v e r s i t y ) a n d a d o c t o r a t e f r o m P r i n c e t o n U n i v e r -
sity. In 1873, w h e n t h e U n i v e r s i t y of C i n c i n n a t i w a s still n e w , he assisted in

Science in the Hinterland 33

 o r g a n i z i n g classes a n d in p u t t i n g t h e i n s t i t u t i o n in order; in fact, he is
c o u n t e d a s a n i n t e r i m p r e s i d e n t o f the u n i v e r s i t y ( G r a c e a n d H a n d 1995,
139). M o r e o v e r , h e s e r v e d o n t h e b o a r d a n d a s p r e s i d e n t o f the C o l l e g e o f
M e d i c i n e a n d S u r g e r y o f t h e u n i v e r s i t y for a n u m b e r o f years.
H a r p e r w a s e l e c t e d t o m e m b e r s h i p o f the C i n c i n n a t i S o c i e t y o f Natural
History in 1871 ( C i n c i n n a t i S o c i e t y of N a t u r a l History, 20). His association
w i t h the s o c i e t y w a s l o n g a n d extensive. At various t i m e s he served as curator,
librarian, m e m b e r o f the p u b l i s h i n g c o m m i t t e e , v i c e president, a n d presi-
d e n t . M o r e o v e r , h e served a s a m e m b e r o f the c o m m i t t e e o n the n o m e n c l a -
ture o f t h e rocks o f t h e t y p e - C i n c i n n a t i a n c h a i r e d b y a n o t h e r m e m b e r o f the
C i n c i n n a t i S c h o o l a n d i n c l u d i n g four o t h e r s (S. A . M i l l e r e t al. 1879).
W o o d w a r d High S c h o o l , in C i n c i n n a t i , was h e a d e d by " k i n d l y principal
G e o r g e W. H a r p e r , a g e o l o g i s t in his o w n right, w h o s e p a r t i c u l a r desire in
life w a s to train s t u d e n t s of geology" (Bassler 1947, iv). For e x a m p l e , he facili-
tated the progress of John M. N i c k l e s a n d Ray S. Bassler by a l l o w i n g t h e m
to re-arrange their s c h e d u l e s at t h e s c h o o l so as to be able to work with U l r i c h
in p a l e o n t o l o g i c e n d e a v o r s . M o r e o v e r , in 1896, he c o - a u t h o r e d a p a l e o n t o -
l o g i c a l p a p e r w i t h Bassler, t h e n a h i g h s c h o o l senior ( H a r p e r and Bassler
1896). G e o r g e W. H a r p e r died in 1918. (Bassler 1947; C a s t e r 1965,1982; C u f f e y ,
D a v i s , a n d U t g a a r d 2002; D u r y 1910; H a r p e r 1886, 1902; J o h n s o n 2002;
K r a m e r 1918; M a r t i n 1900; A n o n . 1 8 7 6 , 1 8 7 8 , 1 8 8 5 b , 1886a, b.)

Ray S. Bassler R a y m o n d Bassler ( F i g u r e s 2.4A, 2 . 5 C ) , a s h e w a s c h r i s t e n e d , w a s b o r n i n

P h i l a d e l p h i a i n 1878. A t t h e a g e o f t w o h e m o v e d t o C i n c i n n a t i w i t h his
family. H i s father, S i m o n Stein Bassler (that is, S g t . S. S. Bassler, of t h e U . S .
A r m y S i g n a l C o r p s ) , w a s o n e o f t h e f o u n d e r s o f t h e U n i t e d States W e a t h e r
Bureau.
A l t h o u g h a handwritten card in the archives of the University of C i n -
c i n n a t i g i v e s his full n a m e a s " R a y m o n d S m i t h Bassler," h e a s s u m e d the
p r o f e s s i o n a l f o r m o f his n a m e after h a v i n g m a d e a c q u a i n t a n c e w i t h the
scientific works of John Ray and F. R a y Lankester, a n d , thereafter, he
s i g n e d h i m s e l f a s " R a y S . B a s s l e r " ( C a s t e r 1965, P167).
Bassler a t t e n d e d W o o d w a r d H i g h S c h o o l , w h e r e G e o r g e W . H a r p e r
w a s p r i n c i p a l . D u r i n g his f r e s h m a n year, h e m e t U l r i c h a n d b e c a m e his
t e c h n i c a l assistant. Bassler w a s free to w o r k for U l r i c h in t h e a f t e r n o o n s ,
b e c a u s e H a r p e r a l l o w e d h i m t o c o m p r e s s his classes into t h e m o r n i n g s . A s
p r e v i o u s l y n o t e d , w h i l e o n l y a h i g h s c h o o l s e n i o r , Bassler c o - a u t h o r e d a
p a l e o n t o l o g i c a l p a p e r w i t h H a r p e r i n 1896.
I n 1896 Bassler e n t e r e d t h e U n i v e r s i t y o f C i n c i n n a t i , w h i c h h a d n o
g e o l o g y d e p a r t m e n t a t t h e t i m e . Bassler c o n t i n u e d w o r k i n g w i t h U l r i c h ,
h o w e v e r . T h i s w o r k w a s i n v a l u a b l e e x p e r i e n c e ; a s Bassler said, " T h e t h i n -
s e c t i o n s o f P a l e o z o i c B r y o z o a p r e p a r e d t h e h a r d w a y d u r i n g o u r e i g h t years
a s s o c i a t i o n w e r e e q u i v a l e n t t o several c o l l e g e c o u r s e s a t least, a n d t h e t i m e
w a s n o t o t h e r w i s e lost, for o v e r a t h o u s a n d slides w e r e lett for f u t u r e p u b -
l i c a t i o n s " ( C a s t e r 1965, P168).
Bassler s p e n t c o n s i d e r a b l e t i m e a t t h e facilities o f the C i n c i n n a t i S o -
c i e t y of N a t u r a l History. T h e late E l l i s Y o c h e l s o n related a story told to h i m

34 A Sea without Fish

b y Bassler that t h e y o u n g m a n w a s a l o n e i n t h e b u i l d i n g o n e d a y w h e n a
g e n t l e m a n with a brilliant w h i t e b e a r d s h o w e d u p a n d asked t o b e s h o w n
a r o u n d the p l a c e . Bassler d i d so, a n d , a f t e r w a r d s , t h e b e a r d e d visitor d e -
parted on his w a y b a c k to A l b a n y , N e w York. T h u s , Bassler m e t J a m e s Hall
(1811-1898), p e r h a p s t h e f o r e m o s t p a l e o n t o l o g i s t i n t h e c o u n t r y ( Y o c h e l s o n ,
pers. c o m m . ) .
U l r i c h left t h e C i n c i n n a t i area for W a s h i n g t o n , D . C , i n 1900, a n d
Bassler f o l l o w e d in M a r c h of 1901, w i t h d r a w i n g f r o m t h e U n i v e r s i t y of
C i n c i n n a t i , b e f o r e c o m p l e t i n g his s e n i o r year.
Bassler w o r k e d privately for U l r i c h a n d w e n t t o s c h o o l p a r t - t i m e a t
C o l u m b i a n University (now G e o r g e W a s h i n g t o n U n i v e r s i t y ) ; h e w a s a b l e
to transfer credits back to t h e U n i v e r s i t y of C i n c i n n a t i a n d w a s a w a r d e d a
bachelor's d e g r e e i n June o f 1902. A b o u t that s a m e t i m e , Bassler b e g a n
w o r k i n g for the U n i t e d States N a t i o n a l M u s e u m ( w h e r e C h a r l e s S e h u c h e r t
was his i m m e d i a t e s u p e r v i s o r ) . T h i s a s s o c i a t i o n w i t h t h e N a t i o n a l M u -
s e u m lasted for nearly six d e c a d e s , as Bassler rose t h r o u g h the r a n k s to
b e c o m e h e a d c u r a t o r of g e o l o g y in 1929. A f t e r his r e t i r e m e n t , in 1948, he
c o n t i n u e d as an honorary r e s e a r c h a s s o c i a t e u n t i l his d e a t h in 1961. M e a n -
w h i l e , h e e a r n e d his master's a n d d o c t o r a l d e g r e e s a t G e o r g e W a s h i n g t o n
University, in 1903 and 1905, r e s p e c t i v e l y ; t h e r e a f t e r he was a s s o c i a t e d w i t h
t h e university for the next thirty-eight y e a r s , i n c l u d i n g scry ice as professor
and h e a d o f the G e o l o g y D e p a r t m e n t .
Bassler m u s t h a v e had a s e n s e of h u m o r . K e n n e t h C a s t e r p a s s e d on a
story h e h a d h e a r d f r o m Bassler: A s a professor i n W a s h i n g t o n , D . C ,
Bassler u s e d t o take classes t o t h e z o o . O n e d a y h e w a s l e a d i n g s u c h a
g r o u p , and h e n o t i c e d that a n elderly lady, w h o s e e m e d slightly f a m i l i a r ,
was trailing a l o n g alter the g r o u p . In any e a s e , he s t o p p e d by a b o u l d e r of
P r e - C a m b r i a n rock o n the z o o g r o u n d s a n d i n f o r m e d his s t u d e n t s that t h e
rock was o n e billion years old. At this p o i n t , the elderly lady i n t e r r u p t e d :
" B u t , Professor Bassler, I b e l i e v e that y o u h a v e m a d e a m i s t a k e . It h a p p e n s
that I was here last y e a r w h e n y o u b r o u g h t y o u r s t u d e n t s . A t t h a t t i m e y o u
said that the rock w a s a b i l l i o n years old. S o , this year, it m u s t be a b i l l i o n
a n d o n e " (Caster, pers. c o m m . ) .
D u r i n g the s u m m e r o f 1909 Bassler d i d a m a n u s c r i p t g e o l o g i c a l m a p
o f t h e East C i n c i n n a t i Q u a d r a n g l e for t h e p r o p o s e d C i n c i n n a t i F o l i o o f
the U n i t e d States G e o l o g i c a l Survey. This m a p w a s n e v e r p u b l i s h e d .
D u r i n g his life, Bassler was an a u t h o r of o v e r 200 p a p e r s , many of
w h i c h w e r e lengthy- w o r k s . H e w a s t h e f o r e m o s t e x p e r t o n P a l e o z o i c b r y o -
z o a n s and w a s o n e o f the p i o n e e r s i n P a l e o z o i c o s t r a c o d e s a n d i n c o n o -
clonts. thereby e n c o u r a g i n g the d e v e l o p m e n t of m i c r o p a l e o n t o l o g v . It was
Bassler's timely c o m p l e t i o n of t h e b r y o z o a n v o l u m e of t h e Treatise on In-
vertebrate Paleontology that a l l o w e d that p r o j e c t to g e t off t h e g r o u n d ,
w h i c h m i g h t not h a v e h a p p e n e d w i t h o u t t h e a p p e a r a n c e o f Bassler's v o l -
u m e (Bassler 1953). A m o n g his most v a l u a b l e w o r k s is a b i b l i o g r a p h y a n d
i n d e x of A m e r i c a n fossils f r o m t h e O r d o v i c i a n a n d S i l u r i a n (Bassler 1915).
K e n n e t h C a s t e r used to call a 10X h a n d - l e n s a " B a s s l e r o s c o p e " ( C a s t e r ,
pers. c o m m . ) . A c c o r d i n g t o C a s t e r , Bassler r e f u s e d t o u s e a c o m p o u n d
m i c r o s c o p e ; h e n c e , his u n d e r s t a n d i n g o f f o s s i l s w a s a r r i v e d a t w i t h o u t t h e

Science in the Hinterland 35

 b e n e f i t o f h i g h e r m a g n i f i c a t i o n ( C a s t e r 1965,1981). I f that w e r e true a t o n e
t i m e , Bassler m u s t h a v e s e e n t h e l i g h t , at least w i t h respect to b r y o z o a n s :
". . . we c a n not be sure of t h e p o s i t i o n of any form in the s c h e m e of clas-
sification u n t i l w e h a v e l e a r n e d its i n t e r n a l s t r u c t u r e b y m e a n s o f t h i n
s e c t i o n s e x a m i n e d m i c r o s c o p i c a l l y " ( N i c k l e s a n d Bassler 1900, 9). M o r e -
over, a c c o r d i n g to Ellis Y o c h e l s o n , Bassler had a c o m p o u n d m i c r o s c o p e on
his d e s k , a n d it a p p e a r s in p h o t o g r a p h s Y o c h e l s o n h a d seen ( Y o c h e l s o n ,
pers. c o m m . ) .
B a s s l e r w a s r e c o g n i z e d for his g r e a t a c c o m p l i s h m e n t s d u r i n g his life-
t i m e . H e w a s e l e c t e d s e c r e t a r y o f t h e P a l e o n t o l o g i c a l S o c i e t y a n d served i n
that p o s i t i o n f r o m 1910 to 1931, a n d t h e n he b e c a m e president of t h e society.
I n 1933, h e w a s p r e s i d e n t o f t h e G e o l o g i c a l S o c i e t y o f A m e r i c a .
W h e n R a y S . Bassler d i e d i n 1961, t h e C i n c i n n a t i S c h o o l o f P a l e o n t o l -
o g y w a s n o m o r e e x c e p t i n t h e i r vast n u m b e r s o f fossils i n m u s e u m s
a r o u n d t h e w o r l d a n d in their p u b l i c a t i o n s in libraries. He w a s t h e last
survivor. (Bassler 1933; B e c k e r 1938; Brandt and D a v i s 2007; C a s t e r 1965,
1981; C r o n e i s 1963; H a r p e r a n d Bassler 1896; N i c k l e s 1936; N i c k l e s and
Bassler 1900; S h i d e l e r [1952] 2002.)

The Cincinnati A l t h o u g h a n u m b e r o f t h e m e m b e r s o f the C i n c i n n a t i S c h o o l o f P a l e o n t o l -

School in o g y did serve stints as s c h o o l p r i n c i p a l s in real life, t h e C i n c i n n a t i S c h o o l

h a d no c l a s s r o o m s , n o r did it offer c o u r s e s . It did not e v e n h a v e a football
Retrospect
t e a m ! N o n e t h e l e s s , its m e m b e r s d e f i n i t e l y m a d e t h e g r a d e . T h i s i s s o d e -
spite the fact t h a t , t o t h e m a j o r i t y o f t h e C i n c i n n a t i S c h o o l , p a l e o n t o l o g y
w a s n o t a p r o f e s s i o n , b u t r a t h e r a n a v o c a t i o n . T o call t h e s e i n d i v i d u a l s
" a m a t e u r s " i s a t o n c e true a n d u n f a i r . A l t h o u g h t h e y did not m a k e their
l i v i n g s a s p a l e o n t o l o g i s t s , their p u b l i s h e d w o r k s h a v e held u p a s w e l l a s
m u c h o f w h a t was a u t h o r e d b y t h e a c t u a l " p r o f e s s i o n a l s " o f the day. M o r e -
o v e r , s o m e o f t h e m e m b e r s o f t h e C i n c i n n a t i S c h o o l d i d , i n fact, g o o n t o
b e c o m e a m o n g s t t h e l e a d i n g " p r o f e s s i o n a l s " o f their t i m e .
It is primarily t h r o u g h t h e efforts of t h e C i n c i n n a t i S c h o o l of P a l e o n -
tology that the C i n c i n n a t i area is truly w o r l d f a m o u s for its fossils. It was
d u e t o their work that t h e C i n c i n n a t i r e g i o n i s t h e N o r t h A m e r i c a n stan-
dard for t h e s p a n o f g e o l o g i c t i m e d u r i n g w h i c h its rocks w e r e d e p o s i t e d
a n d t h e o r g a n i s m s that w e r e t o b e c o m e its fossils lived.
However, it was not only the m e m b e r s of the C i n c i n n a t i School w h o
h a v e w o r k e d o n t h e r o c k s a n d fossils o f t h e t y p e - C i n c i n n a t i a n . There w e r e ,
a n d c o n t i n u e to b e , a g r e a t m a n y o t h e r s i n v o l v e d . In a p p e n d i x 2 of this
b o o k , we have c o m p i l e d brief biographies of s o m e of these other individu-
als a n d o f i n s t i t u t i o n s a s s o c i a t e d w i t h the study o f t h e g e o l o g y a n d p a l e o n -
tology of the C i n c i n n a t i region.

36 A Sea without Fish

 3
NAMING AND CLASSIFYING
ORGANISMS

W h e n p e o p l e from a n u m b e r o f d i f f e r e n t 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 w i t h o n e a n o t h e r , e v e n t u a l l y t h e r e i s a p r o b l e m , n a m e l y , l a n g u a g e .
D i f f e r e n t p e o p l e s have different n a m e s for t h e s a m e a n i m a l ; for e x a m p l e ,
" c h a t , " "felix," "gato," "gatto," a n d " K a t z e " all refer t o t h e a n i m a l w e c a l l
"cat." M o r e o v e r , the s a m e w o r d m a y b e u s e d t o d e s i g n a t e m o r e t h a n o n e
kind o f a n i m a l ; lor i n s t a n c e , w e use the w o r d " c a t " w h e n t a l k i n g a b o u t a
h o u s e eat, or a l i o n , or a tiger, or a b o b c a t , or a m o u n t a i n l i o n , or. . . .
B e g i n n i n g well over two centuries ago, it gradually was r e c o g n i z e d
that, i f scientists a r o u n d t h e w o r l d w e r e t o c o m m u n i c a t e w i t h o n e a n o t h e r
s u c c e s s f u l l y , e a c h kind o f p l a n t a n d a n i m a l m u s t h a v e its o w n u n i q u e
n a m e , and that e a c h n a m e m u s t refer t o o n e , a n d o n l y o n e , k i n d o f p l a n t
o r a n i m a l . A t that t i m e , all e d u c a t e d E u r o p e a n s k n e w G r e e k a n d , e s p e -
cially, L a t i n , so it was s u g g e s t e d that t h e s e plant n a m e s a n d a n i m a l n a m e s
b e i n o n e o f t h e s e classical l a n g u a g e s ; t h a t w a y , n o o n e m o d e r n l a n g u a g e
w o u l d b e favored. Lor s i m p l i c i t y , h o w e v e r , i t w a s d e c i d e d that G r e e k letters
w o u l d not b e u s e d ; h e n c e , o n l y R o m a n letters w e r e e m p l o y e d i n t h e s e
scientific n a m e s .
T h e n a m e for e a c h basic k i n d o f p l a n t o r a n i m a l c o n s i s t s o f t w o w o r d s .
T o illustrate this s c i e n t i f i c n a m i n g , c o n s i d e r t h e c o m m o n h o u s e c a t , Felis
catus. Felis catus is c a l l e d a s p e c i e s n a m e b e c a u s e e a c h basic k i n d of or-
g a n i s m is c a l l e d a s p e c i e s . The last w o r d , in this e a s e Felis, is t h e g e n e r i c
n a m e ; the s e c o n d w o r d i s c a l l e d t h e s p e c i f i c n a m e o r t r i v i a l n a m e . The
s p e c i e s n a m e , c o n s i s t i n g o f b o t h the g e n e r i c a n d t h e s p e c i f i c (or trivial)
n a m e s , i s c a l l e d a b i n o m e n , literally " t w o n a m e s . " T h e s p e c i e s n a m e o f a n
o r g a n i s m c o m m o n l y i s c a l l e d its s c i e n t i f i c n a m e .
N o t e that t h e s p e c i e s n a m e is in italics. T h i s w a s a g r e e d to by s c i e n -
tists: e a c h b i n o m e n is to be put in a form that s t a n d s o u t f r o m the w r i t i n g
a r o u n d it. G e n e r a l l y this is d o n e w i t h italics, a l t h o u g h s o m e t i m e s w i t h
u n d e r l i n i n g . R e m e m b e r , t h e s p e c i e s n a m e m u s t b e i n R o m a n letters, s o
that e v e n in a Russian or C h i n e s e scientific w o r k , Felis catus w i l l l e a p o u t
of the p a g e at y o u .
T h e initial letter of the g e n e r i c n a m e always is c a p i t a l i z e d . If the s p e c i e s
n a m e of an a n i m a l is printed u s i n g b o t h upper-case and lower-case letters,
then the trivial n a m e always is in lower-case t h r o u g h o u t , e v e n the initial letter.
( T h i s latter c o n v e n t i o n was not universally followed in former t i m e s , espe-
cially if a species was n a m e d after s o m e o n e . For e x a m p l e , in 1872, F. B. M e e k
n a m e d the species Glyptocrinus Dyeri after C. B. D y e r , a w e l l - k n o w n fossil
collector in C i n c i n n a t i about w h o m we wrote in the previous chapter.)

37
 In s o m e scientific w o r k s , y o u m a y see a scientific n a m e followed by a
person's n a m e , a c o m m a , a n d a date, for e x a m p l e , Felis catus L i n n a e u s , 1758.
This m e a n s that it w a s C a r o l u s (or C a r l ) L i n n a e u s w h o n a m e d the species in
t h e tenth e d i t i o n of his b o o k Systema Naturae, p u b l i s h e d in 1758. L i n n a e u s
i n v e n t e d t h e b i n o m i a l system of n a m i n g o r g a n i s m s , and he did so in that
b o o k . B e c a u s e of the great s c o p e a n d i m p o r t a n c e of that work, it is c o m m o n
to abbreviate " L i n n a e u s , 1758" to " L . " H e n c e , y o u m i g h t see Felis catus L.
( T h e r e has b e e n s o m e c o n f u s i o n a s t o L i n n a e u s ' s real n a m e ; see D a v i s
[1992]. As w a s the c u s t o m in S w e d e n at the t i m e , C a r l Linnaeus's father, Nils,
o r i g i n a l l y was c a l l e d N i l s I n g e r m a r s s o n , after his father, Ingermar. As a y o u n g
m a n , N i l s i n t e n d e d to b e c o m e a pastor, a n d , w h e n he registered as a student,
he w a s r e q u i r e d to give a f a m i l y n a m e , rather than just the p a t r o n y m i c . He
c h o s e " L i n n a e u s , " a L a t i n word referring to a l i m e t r e e t h e r e was o n e grow-
i n g i n the f a m i l y g a r d e n . After C a r l w a s f a m o u s a n d e n n o b l e d b y the k i n g ,
he a d o p t e d the honorific form "von L i n n e . " It is for this reason that the n a m e
" L i n n a e u s " s o m e t i m e s is written " L i n n e " | M o o r e , pers. comm.].)
A c t u a l l y , t h e r e g u l a r i z a t i o n o f b i o l o g i c a l n o m e n c l a t u r e (the s c i e n c e
o f n a m i n g t h e g r o u p s into w h i c h o r g a n i s m s arc classified) w a s o n l y o n e o f
the contributions of L i n n a e u s . He also was the inventor of the system we
u s e for that c l a s s i f i c a t i o n . It w o r k s like this: e a c h basic k i n d of o r g a n i s m is
c a l l e d a s p e c i e s . R e l a t e d s p e c i e s are j o i n e d t o g e t h e r in a larger u n i t , a g e n u s
(the plural is " g e n e r a " ) . R e l a t e d g e n e r a are g r o u p e d into an e v e n larger u n i t ,
a f a m i l y . A n d so o n . In s h o r t , L i n n a e u s i n v e n t e d w h a t now is c a l l e d the
L i n n a e a n h i e r a r c h y , a s y s t e m o f c a t e g o r i e s i n w h i c h s m a l l e r g r o u p s o f re-
lated o r g a n i s m s a r e j o i n e d t o f o r m larger, m o r e i n c l u s i v e g r o u p s . Take t h e
h o u s e eat, for e x a m p l e . Felis catus i s t h e n a m e o f t h e s p e c i e s . T h e s p e c i e s
F. catus a n d o t h e r , related cats b e l o n g in t h e g e n u s Felis. T h e g e n u s Felis
a n d o t h e r g e n e r a o f cats a r c a s s i g n e d t o t h e F e l i d a e , t h e cat family. C a t s ,
d o g s , b e a r s , s k u n k s , a n d s o o n , m a k e u p the order C a r n i v o r a . T h e orders
C a r n i v o r a , I n s e c t i v o r a ( s h r e w s , m o l e s , h e d g e h o g s , a n d their kin), P r i m a t e s
( m o n k e y s , a p e s , h u m a n s , a n d their r e l a t i v e s ! , a n d all the other orders o f
h a i r y c r e a t u r e s c o m p r i s e t h e class M a m m a l i a . M a m m a l s , birds, reptiles,
a m p h i b i a n s , f i s h e s , a n d s o o n , c o m p r i s e t h e p h y l u m C h o r d a t a . The chor-
d a t e s , c n i d a r i a n s (corals, a n d s o o n ) , s p o n g e s , a n d m o r e t h a n t w e n t y o t h e r
p h y l a p l u r a l of " p h y l u m " c o m p r i s e the k i n g d o m A n i m a l i a .
L e t u s s u m m a r i z e this i n t a b u l a r f o r m , u s i n g t h e c o m m o n h o u s e c a t
as an e x a m p l e :

Kingdom Animalia
Phylum Chordata
Class Mammalia
Order Carnivora
Family Felidae
Genus Felis
Species Felis catus

The entities listed in the left c o l u m n are levels in the L i n n a e a n hierar-

chy. Each word in t h e right c o l u m n is t h e n a m e of o n e g r o u p at that level in
the hierarchy. Each of t h e s e g r o u p s is a t a x o n (the plural is taxa). S o , for

38 A Sea without Fish

e x a m p l e , " f a m i l y " is the level in t h e L i n n a e a n h i e r a r c h y b e t w e e n " o r d e r "
and "genus," and the " F e l i d a e " is t h e taxon at t h e family-level to w h i c h Felis
catus b e l o n g s .
All o f this i s w i t h i n t h e r e a l m o f t a x o n o m y t h e s c i e n c e o f a s s i g n i n g
organisms t o their p r o p e r b i o l o g i c a l g r o u p s . The w o r d " t a x o n o m y " c o m e s
from t w o G r e e k w o r d s , taxis, m e a n i n g " a r r a n g e m e n t , " a n d norms, meaning
" l a w " o r " s c i e n c e of" ( B r o w n 1956); thus " t a x o n o m y " literally m e a n s the " l a w
o f a r r a n g e m e n t " o r t h e " s c i e n c e o f a r r a n g e m e n t " ; alternative n a m e s are c l a s -
s i f i c a t i o n a n d s y s t e m a t i c s . The real c h a l l e n g e o f t a x o n o m y is, o f c o u r s e ,
f i g u r i n g out the biological relationships o f the o r g a n i s m s b e i n g s t u d i e d . A f t e r
that has b e e n d o n e , and o n l y after that has b e e n d o n e , c a n t h e g r o u p s b e
n a m e d in a truly m e a n i n g f u l fashion, at least, f r o m a b i o l o g i c a l p o i n t of view.
A l t h o u g h s o m e scientists m i g h t separate " n o m e n c l a t u r e , " t h e n a m i n g o f the
groups, from f i g u r i n g o u t the b i o l o g i c a l relationships, t h e t w o activities are
irrevocably i n t e r t w i n e d .
O n e purpose for the L i n n a e a n hierarchy is to simplify the d e s c r i b i n g of
kinds of organisms. I m a g i n e if y o u had to describe a h o u s e cat COMPLETELY; it
w o u l d take reams of paper and m a n y m o n t h s of time. B e c a u s e of the Linnaean
hierarchy, by saying "Felis catus," you c o n v e y to your listener all the character-
istics of the species, g e n u s , family, and so o n , w i t h o u t h a v i n g to use up p a p e r
and time in vast quantities. (Note that the n a m e of the species is Felis catus,
not just catus, w h i c h is only the specific n a m e part of the species name.)
However, the L i n n a e a n hierarchy is not m e r e l y a c o n v e n i e n t w a y to save
words and hours. T h e real s i g n i f i c a n c e is that it g r o u p s together o r g a n i s m s
that are evolutionarily related to o n e another. T h u s , t h e o r g a n i s m s joined to-
gether in a given taxon are m o r e closely related to o n e a n o t h e r than they are
to organisms of any other taxon at the s a m e level in the hierarchy. Put a n o t h e r
way, all m e m b e r s of a given taxon evolved from a c o m m o n ancestor.
T h e r e are s e v e n basic levels i n the L i n n a e a n h i e r a r c h y . W h e n d o i n g
d e t a i l e d work w i t h a g r o u p of o r g a n i s m s , o n e c o m m o n l y finds a n e e d for
m o r e levels. A s a result, extra levels h a v e b e e n i n v e n t e d . F o r e x a m p l e ,
several related f a m i l i e s m a y b e g r o u p e d into a s u p e r f a m i l y . O r , g o i n g i n
t h e o t h e r d i r e c t i o n , a f a m i l y m a y c o n s i s t o f several related s u b f a m i l i e s , a n d
a s u b f a m i l y m a y be c o m p o s e d of several i n f r a f a m i l i e s . A n a l o g o u s l y , a b o v e
t h e class-level, there c a n b e s u p e r c l a s s e s , a n d b e l o w i t c a n b e s u b c l a s s e s
a n d infraclasses. A n d s o o n for t h e o t h e r l e v e l s o f t h e h i e r a r c h y . O n e d o e s
not h a v e t o u s e all t h e levels o f t h e e x p a n d e d h i e r a r c h y , h o w e v e r , just t h e
o n e s n e c e s s a r y best t o classify t h e c r e a t u r e s b e i n g s t u d i e d .
O n e o f the m o r e c o m m o n l y u s e d levels o f the e x p a n d e d L i n n a e a n h i -
erarchy is t h e s u b s p e c i e s . Platystrophia ponderosa auburnensis is a s u b s p e -
cies of P. ponderosa, a l o n g with P. ponderosa ponderosa. N o t e that the s u b -
specific n a m e a p p e a r s in print in exactly the s a m e style as d o c s the specific
name. The word " v a r i e t y " s o m e t i m e s is u s e d as an alternative for " s u b s p e -
cies." ( N o t e that " s u b s p e c i e s " is s i m p l y a level in t h e L i n n a e a n h i e r a r c h y ;
there is no c o n n o t a t i o n that a " s u b s p e c i e s " is inferior in " q u a l i t y " to a " s p e -
cies." Thus, if an o r g a n i s m is assigned to a p a r t i c u l a r s u b s p e c i e s , this m e a n s
that the o r g a n i s m m e r e l y has b e e n a s s i g n e d to a p a r t i c u l a r s u b d i v i s i o n of the
s p e c i e s , not that the o r g a n i s m s o m e h o w is qualitatively inferior.)

Naming and Classifying Organisms 39

W h a t Funny Names! I f y o u h a v e tried t o w r a p y o u r t o n g u e a r o u n d the scientific n a m e s o f fossils,
y o u c a n identify w i t h Jules V e r n e ' s o n e - l i n e r a b o u t scientific t e r m s (al-
t h o u g h h e w a s r e f e r r i n g s p e c i f i c a l l y t o m i n e r a l o g i c a l ones). Part o f t h e
p r o b l e m i s that t h e n a m e s o f fossils d o n o t s e e m t o m a k e s e n s e t h e y a p -

. . . many learned words, p e a r t o b e r a n d o m c o m b i n a t i o n s o f letters. Y e s , t h e y h a v e a u t i l i t a r i a n

half-Greek, half-Latin, significance in d e n o t i n g taxa, but the n a m e s generally have "real" m e a n -
and always difficult to i n g s , t o o . H o w e v e r , t h o s e m e a n i n g s g e n e r a l l y h a v e their roots i n a n c i e n t
pronounce, many unpol- L a t i n or G r e e k or b o t h , w h i c h is u n f o r t u n a t e for the vast majority of us,
ished terms that would w h o are n o t s c h o o l e d i n t h e s e c l a s s i c a l l a n g u a g e s .
scorch a poet's lips.
S o m e n a m e s are e p o n y m o u s , that is, t h e y arc d e r i v e d from t h e n a m e s
Jules V e r n e , [1864] o f p e o p l e . T h e r e i s a g e n u s o f c o m m o n a r t i c u l a t e - b r a c h i o p o d s f o u n d i n the
1992, Journey to the type-Cincinnatian called Rafinesquina after C. S. R a f i n e s q u e (1783-1840),
Centre of the Earth, 4 a naturalist w h o taught at Transylvania C o l l e g e in L e x i n g t o n , Kentucky.
T h e c r i n o i d Pycnocrinus dyeri w a s n a m e d in h o n o r of l o c a l fossil c o l l e c t o r
C . B . D y e r (1806-1883). A n d s o o n .
O t h e r n a m e s d e r i v e f r o m p l a c e s . F o r e x a m p l e , the e d r i o a s t e r o i d s Cin-
cinnatidiscus a n d Isorophus cincinnatiensis were named after a city that,
o n c e u p o n a t i m e , w a s t h e c a p i t a l o f t h e old N o r t h w e s t Territory. ( T h e
L a t i n suffix " - e n s i s " o r " - e n s e " d e n o t e s p l a c e o r locality.)
E a c h g e n u s has w h a t i s k n o w n a s a t y p e - s p e c i e s ; this w a s d e s i g n a t e d
to r e p r e s e n t t h e g e n u s . In s o m e c a s e s , t h e trivial n a m e indicates this special
status, for e x a m p l e , t h e p e l e c y p o d Cymatonota typicalis. (Note, however,
that n o t all g e n e r a h a v e t y p e - s p e c i e s that are n a m e d in s u c h a w a y as to
s i g n a l that status.)
In s o m e instances, a n a m e m a y derive from the rock-unit in w h i c h a
t a x o n o c c u r s . S o , for e x a m p l e , t h e a r t i c u l a t e b r a c h i o p o d Leptaena rich-
mondensis o c c u r s i n r o c k s o f t h e R i c h m o n d i a n S t a g e .
O c c a s i o n a l l y , a n a m e is t a k e n d i r e c t l y f r o m t h e L a t i n or G r e e k . Rana
is a c o m m o n g e n u s of f r o g s , a n d " r a n a " is L a t i n for " f r o g . " The h e a d of a
s p e c i m e n of t h e trilobite Phacops rana, " p e e p i n g " out of a rock, is strongly
r e m i n i s c e n t of t h a t of a frog.
Q u i t e c o m m o n l y a g e n e r i c o r trivial n a m e i s d e s c r i p t i v e . T h e b r y o -
zoan Constellaria b e a r s b u m p s that are d i s p o s e d in star-shaped patterns
("stella," L a t i n for "star"). T h e shell of t h e i n a r t i c u l a t e b r a c h i o p o d Trematis
millepunctata b e a r s m a n y h o l e s ( " t r e m a " = " h o l e " [ G r e e k ] ; " m i l l e " = 1000
[ G r e e k ] ; " p u n c t u m " = " s m a l l h o l e " [Latin]), T h a e r o d o n t a rugosa has w r i n -
k l e s ( " r u g a " = " w r i n k l e " [Latin]). A n d s o o n .
O r t h e n a m e m a y reflect t h e h a b i t s o r h a b i t a t o f a n a n i m a l . T h e trilo-
bite Flexicalymene c o m m o n l y is f o u n d flexed into a b a l l . The i n a r t i c u l a t e
b r a c h i o p o d Petrocrania scabiosa lived a t t a c h e d to o t h e r shells, rather like
t h e s c a b a t t a c h e d t o t h e shin y o u b a r k e d last w e e k .
The m e s s a g e h e r e is that g e n e r i c a n d trivial n a m e s are n o t just c a b a -
listic c o m b i n a t i o n s o f letters u s e d t o refer t o taxa. T h e y a l m o s t a l w a y s h a v e
m e a n i n g s b e y o n d t h a t m e a n i n g s that m a k e sense! (For m o r e i n f o r m a t i o n
o n t h e m e a n i n g s o f s c i e n t i f i c n a m e s , s e e B r o w n [1956].)

40 A Sea without Fish

If y o u hear s o m e o n e t r y i n g to talk a b o u t a s u b j e c t , b u t that p e r s o n r o u t i n e l y Pronouncing
s t u m b l e s over t e c h n i c a l t e r m s o r m i s p r o n o u n c e s t h e m , y o u are b o u n d t o Those Lip-
suspect that h e o r she d o e s n o t k n o w t h e s u b j e c t very w e l l . S o how s h o u l d
Blistering Names
one pronounce the n a m e s of t a x a s o as not oneself to be labeled an
ignoramus?
Proper p r o n u n c i a t i o n of w o r d s that are L a t i n or G r e e k d e p e n d s on a
k n o w l e d g e o f those classical l a n g u a g e s . A l a s ! V e r y few p e o p l e t o d a y h a v e the
requisite k n o w l e d g e . T o m a k e matters w o r s e , e v e n a m o n g those w h o m i g h t
lay a c l a i m to b e i n g well-versed in L a t i n or G r e e k , not e v e r y o n e a g r e e s as to
w h a t constitutes correct p r o n u n c i a t i o n . For e x a m p l e , d e v o t e e s o f " C h u r c h
L a t i n " and those o f " C l a s s i c a l L a t i n " d o n o t sing the s a m e C h r i s t m a s carols
the s a m e w a y . There arc, h o w e v e r , s o m e rules o f t h u m b :

1. Unless you k n o w otherwise, put the emphasis on the antepenulti-

m a t e syllable (that is, t h e s y l l a b l e b e f o r e t h e s y l l a b l e b e f o r e t h e last).
T h u s , Brachiopoda is B r a c h i O p o d a .
2. C ' s a n d G ' s are o r d i n a r i l y hard (as in " c a t " a n d " g u n , " r e s p e c t i v e l y ) .
3. V ' s are p r o n o u n c e d as W ' s .
4. J's s h o u l d s o u n d like the Y in "your."
5. A " l o n g i" in Latin is p r o n o u n c e d like a " l o n g e" in E n g l i s h .
6. The d i p h t h o n g " a e " in L a t i n is p r o n o u n c e d l i k e a " l o n g i" in E n g -
lish, viz., " e y e . "
7. Y is p r o n o u n c e d rather like t h e " o o " in t h e w o r d " l o o k . "

B u t t h e a b o v e " r u l e s " c a n result in s o m e s t r a n g e - s o u n d i n g n a m e s , for

e x a m p l e , the n a m e o f t h e e d r i o a s t e r o i d Cincinnatidiscus w o u l d s o u n d like
" k i n k i n n a t i d i s k u s , " a n d t h e s n a k e Virginia w o u l d b e g i n w i t h t h e syllable
"we're," f o l l o w e d by a hard G. (As an a s i d e , a b u n c h of m a l e g r a d u a t e s of a
c o l l e g e o r university are c a l l e d " a l u m n i , " w i t h the last s y l l a b i c b e i n g p r o -
n o u n c e d " n e e , " w h e r e a s their f e m a l e c l a s s m a t e s are " a l u m n a e , " w i t h t h e
last syllable r h y m i n g w i t h the E n g l i s h w o r d " n i g h . " )
N a m e s derived from the n a m e s of p e o p l e c a n be a real c o m p l i c a t i o n .
The w e l l - k n o w n C i n c i n n a t i " h o r n - c o r a l " Grewingkia w a s n a m e d for t h e
Russian paleontologist C o n s t a n t i n C a s p a r A n d r e a s G r e w i n g k (1819-1887),
w h o p r o n o u n c e d his n a m e "gray-vink." P r e s u m a b l y , t h e n , t h e g e n e r i c n a m e
should be said " G r a y - v i n k - e e - a h . " H o w e v e r , most A m e r i c a n s w o u l d call it
"Grew-wink'-ee-ah."
This raises the most-important c o n c e p t in this s e c t i o n : t h e o v e r a r c h i n g
goal of p r o n u n c i a t i o n is c o m m u n i c a t i o n . It is i m p o r t a n t to p r o n o u n c e t h e
n a m e s s o that those with w h o m y o u w i s h t o c o m m u n i c a t e will u n d e r s t a n d
what y o u m e a n . L i s t e n i n g to the e x p e r t s in the field is g e n e r a l l y a g o o d way
t o learn h o w t o p r o n o u n c e the n a m e s , b u t the g o a l i s c o m m u n i c a t i o n . S o ,
for e x a m p l e , i f those with w h o m y o u are s p e a k i n g refer t o the c o m m o n
C i n c i n n a t i a n b r a c h i o p o d Zygospira w i t h t h e " y " a n d t h e " i " e a c h s o u n d i n g
like "eve," it is okay to do the s a m e (even it your h i g h s c h o o l L a t i n t e a c h e r
taught you that the "y" s h o u l d s o u n d like the "00" in " l o o k , " a n d that t h e " i "
should sound like " e e " ) . W h e n i n C i n c i n n a t i , d o like the C i n c i n n a t i a n s !
Speaking of c o m m u n i c a t i o n , sometimes one encounters words or
phrases that look a bit like s p e c i e s n a m e s , b u t are n o t . F o r e x a m p l e , t h e

Naming and Classifying Organisms 41

 term nomen dubium refers to t h e n a m e of a t a x o n that w a s so p o o r l y d e -
s c r i b e d a n d o t h e r w i s e d o c u m e n t e d that it is n o t c e r t a i n just w h a t c o n s t i -
tutes t h e t a x o n a n d h o w t o r e c o g n i z e it. A d u b i o u s n a m e , i n d e e d .
Another such phrase is nomen nudum. With a term that literally
m e a n s " n a k e d n a m e , " a bit of b a c k g r o u n d is n e c e s s a r y . W h e n a s p e c i e s is
n a m e d , t h e a u t h o r i s e x p e c t e d t o follow c e r t a i n c o n v e n t i o n s that h a v e b e e n
a g r e e d t o b y t h e c o m m u n i t y o f t h e world's biologists. T h e a u t h o r m u s t
i n d i c a t e t h a t he or she is n a m i n g t h e s p e c i e s for t h e first t i m e ( G e n e s i s
2:19-20, n o t w i t h s t a n d i n g ) . T h e s p e c i e s m u s t h a v e a d i a g n o s i s ; this is a
s p e c i a l k i n d o f d e s c r i p t i o n that tells h o w i n d i v i d u a l s o f t h e " n e w " s p e c i e s
differ f r o m all o t h e r m e m b e r s o f t h e g e n u s . A n d o n e o r m o r e t y p e - s p e c i -
mens must be indicated.
T y p e - s p e c i m e n s serve as the material on w h i c h the species is based.
T h e y s h o u l d be d e p o s i t e d in a bona fide m u s e u m so that scientists of future
generations c a n study the exact s p e c i m e n s on w h i c h a given species is
b a s e d . It u s e d to be c o m m o n to b a s e a " n e w " s p e c i e s on a s i n g l e s p e c i m e n .
N o w , g i v e n t h e i n t r a s p e c i f i c v a r i a t i o n that has b e e n f o u n d t o exist w i t h i n
all s p e c i e s , it is m o r e c o m m o n to d e s i g n a t e a suite of t y p e - s p e c i m e n s w h e n
a s p e c i e s is n a m e d . If all t h e t y p e - s p e c i m e n s arc c o n s i d e r e d to be of e q u a l
v a l u e a s r e p r e s e n t i n g t h e s p e c i e s , t h e y arc said t o b e c o t y p e s . O n the o t h e r
h a n d , if t h e r e is s i n g l e t y p e - s p e c i m e n , or o n l y o n e of a suite is c o n s i d e r e d
t o b e t h e " n a m e b e a r e r " for t h e s p e c i e s , i t i s d e s i g n a t e d the h o l o t y p e , a n d
the o t h e r s o f t h e suite are p a r a t y p e s .
B a c k to nomen nudum: It is a n a m e of a taxon that d o e s n o t have the
associated v e r b i a g e , illustrations, d e s i g n a t e d s p e c i m e n s , and s o o n . W i t h o u t
s u c h m a t e r i a l , there is no w a y to k n o w w h a t constitutes the taxon, either
c o n c e p t u a l l y or s p e c i m e n - w i s e . P r o b a b l y the m o s t c o m m o n w a y that a no-
men nudum c o m e s into e x i s t e n c e is t h r o u g h the vagaries of the p u b l i c a t i o n
p r o c e s s . S u p p o s e that a p a l e o n t o l o g i s t is p r e p a r i n g t w o scientific papers for
p u b l i c a t i o n ; o n e is to c o n t a i n c o m p r e h e n s i v e d e s c r i p t i o n s and illustrations
of " n e w " taxa. T h e o t h e r is just a t a b u l a t i o n of the fossils in a g i v e n rock-unit
in a given area; it consists of just t h e n a m e s of t h e taxa formally d e s c r i b e d in
t h e former. A l a s ! By a quirk of fate (or b a d p l a n n i n g ? ) , the list is p u b l i s h e d
q u i c k l y , b u t the p a p e r with the f o r m a l d e s c r i p t i o n s a p p e a r s later, or, even
w o r s e , not at all. A n d a spate of nomina nuda are b o r n .

The G a m e of A m a t e u r fossil c o l l e c t o r s c o m m o n l y g e t m o r e t h a n a little a g g r a v a t e d a t

"Musical N a m e s " p a l e o n t o l o g i s t s for c h a n g i n g t h e n a m e s o f fossils. I n d e e d , o n e o f t h e b u g a -

b o o s o f all folks w h o s t u d y fossils i s that t h e n a m e s s o m e t i m e s c h a n g e w h e n
y o u least e x p e c t it.
Take, for e x a m p l e , the c a s e of a w e l l - k n o w n l o c a l c r i n o i d . In 1872, F.
B . M e e k r e c o g n i z e d a s p e c i e s h e n a m e d Glyptocrinus dyeri. T h e genus
Glyptocrinus h a d b e e n n a m e d by J a m e s H a l l in 1847, a n d t h e t y p e - s p e c i e s ,
t h e s p e c i e s that e x e m p l i f i e s t h e g e n u s , is Glyptocrinus decadactylus H a l l ,
1847. M e e k t h o u g h t that Glyptocrinus dyeri b e l o n g s in the s a m e g e n u s as
G . decadactylus, a n d so, q u i t e properly, h e u s e d t h e s a m e g e n e r i c n a m e .
A b o u t t h e s a m e t i m e (1883, a c t u a l l y ) , S . A . M i l l e r r e c o g n i z e d w h a t h e

42 A Sea without Fish

c o n s i d e r e d to be a separate g e n u s , a n d be c a l l e d it Pycnocrinus. In s u b s e -
q u e n t years, e x p e r t s on fossil c r i n o i d s d e c i d e d that G. dyeri a c t u a l l y is m o r e
closely related to the t y p e - s p e c i e s of Pycnocrinus t h a n to that of Glyptocri-
nus, so t h e s p e c i e s n a m e d by M e e k w a s r e - a s s i g n e d f r o m t h e latter to t h e
f o r m e r g e n u s . H e n c e , its n a m e w a s c h a n g e d to Pycnocrinus dyeri ( M e e k ,
1872)the p a r e n t h e s e s are s h o r t h a n d to tell us that t h e s p e c i e s n a m e d by
M e e k in 1872, w i t h the s p e c i f i c n a m e " d y e r i , " w a s later t r a n s f e r r e d to t h e
g e n u s Pycnocrinus. ( A l t h o u g h p a r e n t h e s e s u s e d i n that w a y m a y not a p p e a r
i n s o m e g u i d e b o o k s for a m a t e u r fossil c o l l e c t o r s , t h e y c a n p r o v i d e a v a l u -
able h i n t t o t h e p a l e o n t o l o g i s t t r y i n g t o track d o w n t h e n o m e n c l a t o r i a l
history of a p a r t i c u l a r species.)
O n the o t h e r h a n d , s o m e t i m e s the s i t u a t i o n i s t h e o t h e r w a y r o u n d . I n
1935, S a b u r o S h i m i z u a n d T a d a h i r o O b a t a r e c o g n i z e d a " n e w " g e n u s o f
fossil cephalopods and named it Orthonybyoceras. In 1942, Rousseau
F l o w e r , a n e m i n e n t e x p e r t o n fossil c e p h a l o p o d s , r e c o g n i z e d a " n e w " g e -
n u s and c a l l e d it Treptoceras. L a t e r w o r k e r s , for e x a m p l e , C u r t T e i c h e r t ,
o n e o f the m o s t f a m o u s p a l e o n t o l o g i s t s o f his clay a n d , i t h a p p e n s , a n e x p e r t
o n fossil c e p h a l o p o d s , c o n c l u d e d that a n i m a l s f o r m e r l y r e c o g n i z e d a s b e -
l o n g i n g in the t w o separate g e n e r a c o m p r i s e d a s i n g l e t a x o n . In s u c h c a s e s ,
biologists apply a c o n v e n t i o n c a l l e d p r i o r i t y , viz., w h e n t h e r e is an o l d e r
n a m e a n d a y o u n g e r n a m e that b o t h h a v e b e e n u s e d t o d e s i g n a t e t h e s a m e
taxon. the o l d e r n a m e b e c o m e s the official n a m e o f the t a x o n . S o , b e c a u s e
it is older, t h e n a m e Orthonybyoceras S h i m i z u a n d O b a t a , 1935 w a s a p p l i e d
to the g e n u s ( T e i c h e r t 1964, K214).
N o t e that t h e g u i d i n g p r i n c i p l e i n b o t h o f t h e s e e x a m p l e s a n d i n all
similar c a s e s i s for a g i v e n t a x o n to h a v e a s i n g l e , u n i q u e n a m e a n d that
a g i v e n n a m e s h o u l d refer to a s i n g l e , u n i q u e t a x o n . If t w o g r o u p s of o r g a n -
isms b e l o n g i n separate taxa, t h e n t h o s e taxa n e e d t o h a v e s e p a r a t e n a m e s
in the a b o v e e x a m p l e , Glyptocrinus a n d Pycnocrinus. If t w o p u t a t i v e l y s e p a -
rate g r o u p s o f a n i m a l s a c t u a l l y c o m p r i s e a s i n g l e t a x o n , t h e n all s h o u l d
parade under a single namein the a b o v e e x a m p l e , Orthonybyoceras.
N a m e s are not c h a n g e d for frivolous reasons. All creatures with the s a m e
species n a m e b e l o n g in the s a m e s p e c i e s . All c r e a t u r e s w i t h the s a m e generic-
n a m e b e l o n g in the s a m e g e n u s . That way, w h e n a particular g e n u s or s p e c i e s
is m e n t i o n e d by n a m e , e v e r y o n e e v e r y w h e r e k n o w s e x a c t l y w h a t is b e i n g
discussed. The goal of z o o l o g i c a l n o m e n c l a t u r e is c o m m u n i c a t i o n !

Naming and Classifying Organisms 43

Figure 4 . 1 . Cincinnatian stratigraphic nomenclature from 1955 through 1986. From Davis and Cuffey (1998).
From Schumacher (1984, figure 2), courtesy of the Ohio Department of Natural Resources Division of Geo-
logical Survey. This chart shows stratigraphic subdivisions of the Cincinnatian Series proposed by different
researchers for different parts of the Cincinnati Arch region. Subdivisions in the Caster et al. (1955) column
were based largely on differences in fossil content. Broader subdivisions such as those of the Brown and
Lineback (1966), Hatfield (1968), Gray (1972), Peck (1966), and Lee (1974) columns were based on general
characteristics of the rocks (lithology) and bedding. Hay (1981) and Tobin (1986) used both lithologic as well
as paleontologic aspects. In the Hatfield column, the vertical lines indicate parts of the section excluded
from his study. Units separated by jagged lines indicate lateral changes in rock characteristics (faces)

44 A Sea without Fish

ROCKS, FOSSILS, AND TIME

Fossils i n m a n y c o l l e c t i o n s a n d m u s e u m e x h i b i t s are o f t e n i m p r e s s i v e for

f i n e l y p r e s e r v e d detail a n d e v e n b e a u t y , b e c a u s e t h e y h a v e u n d e r g o n e
p a i n s t a k i n g p r e p a r a t i o n b y w h i c h e v e r y t r a c e o f t h e stony m a t r i x h a s b e e n
r e m o v e d . H o w e v e r , a fossil so isolated f r o m its e m b e d d i n g m a t r i x a l s o loses
m u c h of its s i g n i f i c a n c e as a m e a n s by w h i c h to u n d e r s t a n d w h e n a n d how-
it lived. Only t h r o u g h i n v e s t i g a t i o n of the fossil in the rock c a n we a t t a i n a
c l e a r u n d e r s t a n d i n g o f t h e s i g n i f i c a n c e o f t h e a b u n d a n t O r d o v i c i a n fossils
of the C i n c i n n a t i A r c h r e g i o n , or a n y fossils for that m a t t e r . In this c h a p t e r
w e will e x p l o r e the n a t u r e o f t h e r o c k s i n w h i c h C i n c i n n a t i a n fossils are
f o u n d , t h e m e a n s b y w h i c h t h e y are s u b d i v i d e d , a n d t h e a p p l i c a t i o n s o f
this study t o u n d e r s t a n d i n g t h e e n v i r o n m e n t s i n w h i c h t h e y w e r e f o r m e d
and t o d e t e r m i n i n g their g e o l o g i c a g e .
T w o words effectively d e s c r i b e t h e b e d r o c k o f the C i n c i n n a t i A r c h re-
g i o n for e v e n first-time observer: monotonous a n d layered. C i n c i n n a t i rocks
are i n d e e d m o n o t o n o u s as their entire t h i c k n e s s of o v e r 250 m (820 feet)
consists o f a p p a r e n t l y s i m i l a r b l u e - g r e y l i m e s t o n e s a n d shales. These two
c o m m o n s e d i m e n t a r y rocks form t h i n , a l t e r n a t i n g layers ( b e d s or strata) that
appear to be h o r i z o n t a l and c o n t i n u o u s across an e x p o s u r e s u c h as a roadcut.
The overall impression is that of a layer c a k e , a n d in fact C i n c i n n a t i a n b e d -
rock has l o n g b e e n k n o w n as a classic of " l a y e r c a k e g e o l o g y . " C l o s e r e x a m i -
nation by scores of geologists over m o r e t h a n 150 years r e v e a l e d that like m o s t
first impressions, the a c t u a l c h a r a c t e r of C i n c i n n a t i a n strata is q u i t e differ-
ent. G e o l o g i s t s vigorously p u r s u e d the d e t a i l e d d e s c r i p t i o n , s u b d i v i s i o n , a n d
classification of these s e e m i n g l y u n i f o r m strata for a variety of p u r p o s e s , re-
sulting in a l o n g and c o m p l e x history of study. G e o l o g i s t s e m p l o y different
m e t h o d s to d e s c r i b e a n d s u b d i v i d e rocks. For layered s e d i m e n t a r y rocks, t h e
c h i e f m e t h o d s arc relative age, rock type, a n d fossils.

A l t h o u g h the history o f t h e Earth is part o f a c o n t i n u o u s flow o f t i m e , t h e Geeoollooggiicc Time Units

G
e v i d e n c e we h a v e for that past history, the rocks a n d fossils, r e p r e s e n t s o n l y
f r a g m e n t s of that history. For this r e a s o n , g e o l o g i s t s u s e a d u a l set of t e r m s
for divisions of c o n t i n u o u s t i m e a n d t h e r o c k s that r e p r e s e n t p r e s e r v e d in-
tervals of t i m e (see F i g u r e 1.1). T h e c o n t i n u o u s flow of g e o l o g i c t i m e is
d i v i d e d into m a j o r d i v i s i o n s c a l l e d e o n s , w h i c h are i n t u r n d i v i d e d into
eras and p e r i o d s , the f u n d a m e n t a l u n i t s o f E a r t h history. P e r i o d s are s u b -
divided into e p o c h s (Early, M i d d l e , Late), a n d a g e s . I n g e o l o g i c t i m e t e r m s ,
the C i n c i n n a t i a n i s part o f t h e L a t e O r d o v i c i a n E p o c h o f t h e O r d o v i c i a n
Period. T h e O r d o v i c i a n Period i s t h e n e x t - t o - o l d e s t p e r i o d o f t h e P a l e o z o i c

45
 E r a , a n d the P a l e o z o i c Era i s the earliest era o f t h e P h a n e r o z o i c E o n ( m e a n -
i n g " t i m e o f r e v e a l e d life," for t h e a b u n d a n c e o f fossils i n those strata). T h e
o t h e r set of t e r m s , time-stratigraphic units, applies to the a c t u a l rocks that
g e o l o g i s t s assign to p a r t i c u l a r intervals of t h e g e o l o g i c t i m e s c a l e , and are
r o u g h l y e q u i v a l e n t t o t h e divisions o f c o n t i n u o u s t i m e , e x c e p t that there are
m a n y g a p s i n t h e record o f t i m e that result from i n c o m p l e t e preservation.

Relative A g e A c c o r d i n g t o a f u n d a m e n t a l p r i n c i p l e o f g e o l o g y , s u p e r p o s i t i o n , the l o w e s t
r o c k layers i n a n u n d i s t u r b e d v e r t i c a l s e q u e n c e w e r e d e p o s i t e d b e f o r e lay-
ers l y i n g a b o v e t h e m . T h u s , for t h e C i n c i n n a t i " l a y e r c a k e , " w e k n o w that
layers e x p o s e d i n t h e b e d o f t h e O h i o R i v e r f o r m e d b e f o r e layers e x p o s e d
h i g h e r a l o n g t h e h i l l s i d e s , b u t w e d o n o t k n o w e x a c t l y how old t h e layers
a r e , or how much older are l o w e r layers t h a n h i g h e r layers. We k n o w o n l y
that l o w e r layers are relatively older t h a n h i g h e r layers. G e o l o g i s t s e s t a b -
l i s h e d t h e r e l a t i v e a g e s e q u e n c e o f t h e m a j o r s e d i m e n t a r y layers o f the
E a r t h ' s c r u s t l o n g b e f o r e a n a l y t i c a l m e t h o d s (chiefly r a d i o m e t r i c d a t i n g )
were 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 of rocks.

Time-Stratigraphic R o c k s that f o r m e d d u r i n g a p a r t i c u l a r interval o f g e o l o g i c t i m e are c a l l e d

Units t i m e - s t r a t i g r a p h i c u n i t s . T h u s , rocks d e p o s i t e d d u r i n g t h e O r d o v i c i a n Pe-

riod c o n s t i t u t e the O r d o v i c i a n S y s t e m , w h i c h is s u b d i v i d e d into S e r i e s (usu-
ally L o w e r , M i d d l e , and U p p e r , but s o m e t i m e s n a m e d , as in the C i n c i n n a t i a n
Series). S t a g e s are t h e f u n d a m e n t a l subdivisions of series u s e d for correlation
on a c o n t i n e n t a l a n d i n t e r c o n t i n e n t a l scale. C i n c i n n a t i a n stages are the E d e -
n i a n , M a y s v i l l i a n , and R i c h m o n d i a n , i n order from oldest t o y o u n g e s t .

Rock- R o c k s c a n also b e s u b d i v i d e d o r classified b y r o c k t y p e o r l i t h o l o g y . L i t h o l -

stratigraphic or o g y u s u a l l y i n c l u d e s t h e c o m p o s i t i o n o f t h e r o c k , its c o n s t i t u e n t p a r t i c l e s ,
their size, shape, and a r r a n g e m e n t . The fundamental lithostratigraphic
Lithostratigraphic
u n i t is t h e f o r m a t i o n , d e f i n e d by a d i s t i n c t i v e l i t h o l o g i c c h a r a c t e r as w e l l
Units
as a t h i c k n e s s s u f f i c i e n t to be s h o w n at a c o n v e n i e n t s c a l e of g e o l o g i c m a p -
p i n g . F o r m a t i o n s c a n b e c o m b i n e d a s g r o u p s , o r s u b d i v i d e d into m e m -
b e r s . F o r m a t i o n s d o n o t n e c e s s a r i l y c o i n c i d e w i t h a s p e c i f i c interval o f
t i m e , a n d i n d e e d are o f t e n t i m e - t r a n s g r e s s i v e , that is, their l o w e r o r u p p e r
b o u n d a r i e s are n o t t i m e - e q u i v a l e n t o r i s o c h r o n o u s s u r f a c e s .

Biostratigraphic R o c k s c a n b e s u b d i v i d e d o n t h e basis o f fossil c o n t e n t . T h e t h i c k n e s s o f

Units strata in w h i c h a p a r t i c u l a r k i n d of fossil o c c u r s d e f i n e s the fossil z o n e (or

b i o z o n e ) , w h i c h is the f u n d a m e n t a l biostratigraphic unit. T h e lower and
u p p e r l i m i t s o f a p a r t i c u l a r fossil d o n o t n e c e s s a r i l y represent t h e s a m e t i m e
horizons in different stratigraphic sections. However, t h r o u g h the use of
t h e o v e r l a p p i n g r a n g e z o n e s o f m a n y s p e c i e s , b i o s t r a t i g r a p h e r s establish
t h e b o u n d a r i e s o f t i m e - s t r a t i g r a p h i c u n i t s s u c h a s stages.

46 A Sea without Fish

T h e earliest s t u d i e s o f the r o c k s a n d fossils o f t h e C i n c i n n a t i r e g i o n d a t e Early Studies
to the early 1800s ( D r a k e 1825; R i d d e l l 1837; L o c k e 1838; L y e l l 1845) a n d
c o i n c i d e w i t h t h e very b e g i n n i n g s o f g e o l o g y a s a s c i e n c e . T h e s e early
workers m a d e n o a t t e m p t t o s u b d i v i d e t h e strata a r o u n d C i n c i n n a t i , a n d
referred t o t h e m s i m p l y a s t h e " B l u e M i a m i L i m e s t o n e " ( R i d d e l l 1837),
" B l u e L i m e s t o n e " ( L o c k e 1838), o r " G r e a t L i m e s t o n e D e p o s i t e " ( B r i g g s
1838). F . B . M e e k a n d A . H . W o r t h e n (1865), b o t h p i o n e e r s o f A m e r i c a n
p a l e o n t o l o g y , first u s e d t h e t e r m C i n c i n n a t i G r o u p for t h e O h i o strata.
E d w a r d O r t o n ( O r t o n 1873), O h i o ' s third state g e o l o g i s t , p r o p o s e d a four-
fold s u b d i v i s i o n o f w h a t h e t e r m e d t h e C i n c i n n a t i b e d s p r o p e r : R i v e r
Q u a r r y B e d s ( l o w e r m o s t ) , M i d d l e ( E d e n ) S h a l e s , Hill Q u a r r y B e d s , a n d
L e b a n o n Beds ( u p p e r m o s t ) .

T h e a b u n d a n t and w e l l - p r e s e r v e d fossils o f t h e C i n c i n n a t i r e g i o n a t t r a c t e d Subdivision Based

m o r e and m o r e a t t e n t i o n d u r i n g t h e latter p a r t o f t h e n i n e t e e n t h c e n t u r y , on Fossils
and as a result, d e t a i l e d k n o w l e d g e of t h e d i s t r i b u t i o n of p a r t i c u l a r fossil
s p e c i e s i n t h e strata w a s a c c u m u l a t e d t h r o u g h t h e efforts o f t h e C i n c i n n a t i
s c h o o l of early p a l e o n t o l o g i s t s (see c h a p t e r 2). In 1902 John M. N i c k l e s
published a comprehensive report on the g e o l o g y of C i n c i n n a t i , an excel-
lent r e v i e w a n d c o m p i l a t i o n o f e a r l i e r s t u d i e s ( N i c k l e s 1902).
B y the t i m e o f Niekles's report, the C i n c i n n a t i G r o u p h a d c o m e t o b e
r e c o g n i z e d as " o n e of the major divisions of the O r d o v i c i a n or L o w e r Silurian
Era, with the title of C i n c i n n a t i P e r i o d " ( N i c k l e s 1902, 64). N i c k l e s s u b d i -
vided the Cincinnati Period into three major div I S I O N S . e a c h t e r m e d a g r o u p :
Utica g r o u p ( l o w e r m o s t , 260 feet thick), L o r r a i n e g r o u p (310 feet thick), and
R i c h m o n d g r o u p ( u p p e r m o s t , 2 0 0 - 3 0 0 feet thick). The t e r m s U t i c a a n d Lor-
raine were both derived from c o m p a r i s o n s w i t h the O r d o v i c i a n of New York.
W i n c h e l l and U l r i c h (1897) h a d p r o p o s e d the t e r m R i c h m o n d (from e x c e l -
lent exposures at R i c h m o n d , Indiana) for the u p p e r m o s t C i n c i n n a t i strata
b e c a u s e L e b a n o n had previously b e e n used for older O r d o v i c i a n strata in
Tennessee. The f o l l o w i n g s e c t i o n from Niekles's report c o n t a i n s f u n d a m e n -
tal statements that w e r e the basis for subdivision of the s e c t i o n :

There is considerable variation in the different groups in the proportions of

limestone and shale. Shale greatly predominates in the Utica, but from the
lower beds of the Lorraine on, the proportion of limestone gradually increases.
This shows that there was a gradual change from more or less turbulent condi-
tions prevailing at the close of the Trenton to the time of the Lower Rich-
mond, when quiet seas permitted the a c c u m u l a t i o n of the materials for
closely succeeding beds of limestone. As the period c a m e to a close, there
came anew turbulent conditions. The fauna of the different groups indicates
the same succession of changes. (Nickles 1902, 65)

B o t h the c h a r a c t e r o f t h e rock ( l i t h o l o g y ) a n d t h e fossil c o n t e n t differenti-

ate subdivisions. N i c k l e s listed s p e c i e s that c o u l d b e f o u n d t h r o u g h o u t t h e
entire s e c t i o n , but t h e s e w e r e o n l y 4 p e r c e n t o f all f o r m s k n o w n f r o m t h e
C i n c i n n a t i p e r i o d , a n d " t h e great b u l k o f forms u s u a l l y h a v e a l i m i t e d verti-
c a l r a n g e " ( N i c k l e s 1902, 65). His f u r t h e r s u b d i v i s i o n o f the g r o u p s w a s
b a s e d o n a c o m b i n a t i o n o f l i t h o l o g i c a n d p a l e o n t o l o g i c criteria. T h e U t i c a

Rocks, Fossils, and Time 47

 g r o u p ". . . m a y be d i v i d e d into t h r e e s u b d i v i s i o n s , m o r e easily r e c o g n i z e d
faunally t h a n l i t h o l o g i c a l l y , t h o u g h c l o s e s t u d y s h o w s l i t h o l o g i c a l differ-
e n c e s , w h i c h s o o n c o m e t o b e felt, b u t are n o t easily d e s c r i b e d " ( N i c k l e s
1902, 69) T h e s e w e r e n a m e d the L o w e r Utica or Aspidopora newberryi
B e d s , t h e M i d d l e U t i c a or Batostoma jamesi B e d s , a n d the U p p e r U t i c a or
Dekayella ulrichi B e d s . T h e L o r r a i n e w a s l i k e w i s e s u b d i v i d e d into six
" b e d s , " e a c h g i v e n a l o c a l g e o g r a p h i c n a m e as w e l l as a c h a r a c t e r i s t i c fossil
s p e c i e s , a n d he i n d i c a t e d t h a t t h e g r o u p ". . . is easily s e p a r a b l e on f a u n a l
g r o u n d s , w i t h c o r r e s p o n d i n g m o r e o r less w e l l - m a r k e d l i t h o l o g i c a l c h a r a c -
ters" ( N i c k l e s 1902, 75) H e d i v i d e d t h e R i c h m o n d into l o w e r , m i d d l e , a n d
u p p e r d i v i s i o n s b a s e d o n t h e i r f a u n a s , b u t i n d i c a t e d that study has b e e n
i n s u f f i c i e n t t o establish their b o u n d a r i e s o r l i t h o l o g i c a l c h a r a c t e r s .
In 1903 A u g u s t F. Foerste i n t r o d u c e d for t h e first t i m e t h e t e r m C i n c i n -
n a t i a n S e r i e s for t h e e n t i r e s e c t i o n , and referred t o t h e U t i c a , L o r r a i n e , a n d
R i c h m o n d a s stages ( F o e r s t e 1903). F o e r s t e (1906) t e r m e d t h e s a m e t h r e e
g r o u p s f o r m a t i o n s , w i t h their s u b d i v i s i o n s a s m e m b e r s . H e also d i s c a r d e d
t h e New York t e r m L o r r a i n e a n d r e p l a c e d it w i t h M a y s v i l l e . In t h e s a m e
y e a r R a y S. Bassler (1906) e l e v a t e d t h e n a r r o w e r s u b d i v i s i o n s to f o r m a t i o n s
a n d d e f i n e d t h e C o v i n g t o n C r o u p t o i n c l u d e t h e U t i c a , E d e n , Fairview,
and M c M i l l a n Formations, overlain by the R i c h m o n d C r o u p , including
the A r n h e i m , Waynesville, Liberty, Whitewater, and Saluda Formations.
In time, further subdivisions of these formations were designated as m e m -
b e r s , a l t h o u g h d i f f e r e n t w o r k e r s c o n t i n u e d t o u t i l i z e different stratigraphic
c l a s s i f i c a t i o n s ( W e i s s a n d N o r m a n 1960b).
A g u i d e b o o k to the fossils a n d strata of the C i n c i n n a t i area p r e p a r e d
i n 1939 b y Prof. W a l t e r H . B u c h e r , w i t h illustrations b y K e n n e t h F . C a s t e r ,
assisted b y S t e w a r t Jones, w a s f i r s t p r i n t e d i n f o r m a l l y b y t h e D e p a r t m e n t
o f G e o l o g y o f t h e University o f C i n c i n n a t i ( B u c h e r e t al. 1939). The b o o k l e t
w a s p u b l i s h e d i n 1945 b y t h e C i n c i n n a t i M u s e u m o f N a t u r a l H i s t o r y u n d e r
the title Elementary Guide to the Fossils and Strata in the Vicinity of Cin-
cinnati, Ohio ( B u c h e r et al. 1945); later r e v i s e d by C a s t e r , D a l v e , a n d P o p e
(1955, 1961), it b e c a m e t h e s t a n d a r d for w h a t has b e c o m e k n o w n as t h e
" t r a d i t i o n a l " o r " b i o s t r a t i g r a p h i c " c l a s s i f i c a t i o n o f t h e C i n c i n n a t i a n (Fig-
ure 4.1). T h e s e e d i t i o n s of t h e Elementary Guide i n c l u d e the s t a t e m e n t that
" T h e l i t h o l o g y , i.e., r o c k c h a r a c t e r i s t i c s , o f the b e d s a s i n d i c a t e d o n t h e
charts varies considerably w h e n traced away from the Tri-Statc Area.
T h e r e f o r e , l i t h o l o g i c i d e n t i f i c a t i o n o f t h e C i n c i n n a t i a n f o r m a t i o n s and
b e d s i s u n r e l i a b l e . H o w e v e r , t h e s e q u e n c e o f fossils persists t h r o u g h o u t the
a r e a a n d f u r n i s h e s a r e l i a b l e basis for i d e n t i f y i n g e q u i v a l e n t b e d s , a n d thus
t h e p o s i t i o n i n t h e s t r a t i g r a p h i c s e q u e n c e " ( C a s t e r , D a l v e , a n d P o p e 1955,
15; C a s t e r , D a l v e , a n d P o p e 1961, 15). S u b s e q u e n t revisions of this b o o k by
R. A. D a v i s (1985, 1992), u n d e r t h e title Cincinnati Fossils r e t a i n e d t h e
t i m e - h o n o r e d C i n c i n n a t i a n stratigraphic chart of the older work, but
p o i n t e d o u t h o w i t differs f r o m m o d e r n s t r a t i g r a p h i c u s a g e .
M o d e r n p r a c t i c e s i n t h e f i e l d o f stratigraphy e m e r g e d d u r i n g t h e post-
W o r l d W a r I I era a s efforts w e r e m a d e t o e l i m i n a t e c o n f u s i o n r e s u l t i n g
f r o m n o n - u n i f o r m u s a g e a n d t o s t a n d a r d i z e s t r a t i g r a p h i c classification a n d
t e r m i n o l o g y . T h e history o f C i n c i n n a t i a n stratigraphy ( W e i s s a n d N o r m a n

48 A Sea without Fish

 Figure 4 . 2 . Average per-
cent composition of the
allochem fraction of fau-
nal groups and algae in
the upper (Saluda and
Whitewater 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.

1960b) p r o v i d e s a s t r o n g c a s e d e m o n s t r a t i n g this n e e d ! G u t s t a d t (1958) a n d

W e i s s (1961) p o i n t e d o u t that t r a d i t i o n a l C i n c i n n a t i a n s t r a t i g r a p h i c classi-
f i c a t i o n w a s e s s e n t i a l l y a b i o s t r a t i g r a p h i c z o n a t i o n , sorely i n n e e d o f revi-
sion i n k e e p i n g w i t h n e w c o n c e p t s a n d p r a c t i c e s . P u b l i c a t i o n o f t h e C o d e
of Stratigraphic N o m e n c l a t u r e ( A m e r i c a n C o m m i s s i o n on Stratigraphic
N o m e n c l a t u r e 1961) a n d the I n t e r n a t i o n a l S t r a t i g r a p h i c G u i d e ( S a l v a d o r
1994) c u l m i n a t e d many years of c o l l a b o r a t i v e work b e t w e e n e a r t h scientists
in the United States a n d w o r l d w i d e .
C i n c i n n a t i a n l i t h o s t r a t i g r a p h i c u n i t s are d e f i n e d b y data o n t h e fol-
lowing characteristics obtained from measured sections: limestone types
p r e s e n t , clastic ratio, a n d b e d d i n g f e a t u r e s . L i m e s t o n e s i n t h e C i n c i n n a -
tian display w i d e v a r i a t i o n , r a n g i n g f r o m f i n e - g r a i n e d t o c o a r s e - g r a i n e d
and c o m p o s e d o f v a r y i n g m i x e s o f fossils ( F i g u r e s 4.2, 4.3). A c c e l e r a t e d
research i n l i m e s t o n e ( c a r b o n a t e ) p e t r o g r a p h y e n a b l e d g r e a t l y r e f i n e d d e -
scription o f C i n c i n n a t i a n l i m e s t o n e s ( W e i s s a n d N o r m a n 1960a; M a r t i n
1975; T o b i n 1982). The elastic ratio is t h e p r o p o r t i o n of s h a l e t h i c k n e s s in
relation to l i m e s t o n e t h i c k n e s s in a s e c t i o n . B e d d i n g f e a t u r e s i n c l u d e the
b e d d i n g i n d e x , c a l c u l a t e d as the n u m b e r of b e d s (x 100) in a g i v e n interval
divided b y the t h i c k n e s s o f the i n t e r v a l , a n d b e d f o r m , s u c h a s p l a n a r o r
wavy. T h e s e l i t h o l o g i c c h a r a c t e r i s t i c s c a n b e q u a n t i f i e d a n d p l o t t e d o n
vertical s e c t i o n s o r m a p s .
T h e n e w clarification and codification of stratigraphic usage m e a n t
that C i n c i n n a t i a n f o r m a t i o n s that h a d b e e n d e f i n e d l a r g e l y o n t h e r a n g e s
o f c h a r a c t e r i s t i c fossils w e r e i n v a l i d . The d i s t i n c t i o n w a s m u c h m o r e t h a n
a semantic problem: attempts to r e c o g n i z e the traditional C i n c i n n a t i a n
stratigraphic f o r m a t i o n s b e y o n d t h e i m m e d i a t e vicinity o f C i n c i n n a t i w e r e

Rocks, Fossils, and Time 49

 Figure 4 . 3 . Variation in limestone deposition in relation to benthic fossil as-
semblages. Smaller circles depict the character of different limestone types
as seen in petrographic thin section under the microscope. All limestones are
not the same. Different types of limestone will form depending on the rela-
tive contributions of skeletal grains and other allochems, fine-grained car-
bonate sediment (mud), and calcium carbonate cement (spar). A packstone
has skeletal grains in contact (grain-supported) with mud filling the spaces
between grains. A grainstone is grain-supported with cement filling the in-
tergranular spaces. A wackestone has skeletal grains "floating" in a mud ma-
trix (mud-supported) with more than 10% grains, and a mudstone has less
than 10% grains. In the interior of a benthic assemblage, water movement is
retarded by the dense growth of organisms, causing fine-grained sediment
to be deposited along with skeletal grains, resulting in either packstones or
wackestones. More variable water movement at margins of assemblages will
produce a wider variety of limestones. A grainstone usually reflects the high-
est level of water motion, by which mud is winnowed, leaving only skeletal
grains to be cemented. From Martin and Hauer (2006), courtesy of Wayne
D. Martin.

50 A Sea without Fish

fraught w i t h difficulties. E v e n the i n t e r n a t i o n a l l y r e c o g n i z e d C i n c i n n a t i a n
stages ( E d e n i a n , M a y s v i l l i a n , a n d Richmondian) c o u l d he called into
q u e s t i o n a s valid t i m e - s t r a t i g r a p h i c u n i t s b e c a u s e t h e y w e r e e q u i v a l e n t t o
the E d e n , M a y s v i l l e , a n d R i c h m o n d " C r o u p s " w h i c h c o m p r i s e d " f o r m a -
tions" that l a c k e d t i m e - s t r a t i g r a p h i c s i g n i f i c a n c e . C o n s e q u e n t l y , geologic-
research o n the C i n c i n n a t i a n d u r i n g t h e 1960s resulted in m a j o r revisions
of the stratigraphic classification and n o m e n c l a t u r e .

T h r e e m a j o r r e s e a r c h p r o g r a m s o f t h e 1960s m a r k e d a n e w p h a s e i n t h e Lithostratigraphy
d e v e l o p m e n t a n d u n d e r s t a n d i n g o f C i n c i n n a t i a n s t r a t i g r a p h y ; all w e r e ini- and "Stateline
tiated o u t s i d e o f C i n c i n n a t i . S e v e r a l g e o l o g i s t s a t T h e O h i o S t a t e U n i v e r -
Boundaries"
sity (notably S t i g M . B e r g s t r o m , W a l t e r C . S w e e t , M a l c o l m P . W e i s s , a n d
their students) p u b l i s h e d a series o f p a p e r s a i m e d a t r e v i s i n g t h e A m e r i c a n
Upper Ordovician Standard in both lithostratigraphic and biostratigraphy
terms. I n t h e 1960s t h e K e n t u c k y G e o l o g i c a l S u r v e y a n d t h e U n i t e d States
G e o l o g i c a l S u r v e y initiated a m a j o r p r o j e c t t o p r o v i d e n e w g e o l o g i c m a p s
o f t h e e n t i r e state o f K e n t u c k y a t s c a l e o f t h e 7.5 m i n u t e q u a d r a n g l e
(1:24,000 scale). In order to a c c o m p l i s h this e n o r m o u s task, it w a s n e c e s s a r y
to p r o v i d e a u n i f o r m s t r a t i g r a p h i c c l a s s i f i c a t i o n of m a p p a b l e lithostrati-
g r a p h i c units. I n k e e p i n g w i t h t h e n e w S t r a t i g r a p h i c C o d e , g e o l o g i s t s as-
s o c i a t e d w i t h the m a p p i n g p r o g r a m d e v e l o p e d a n e w s t r a t i g r a p h i c classifi-
cation for the Ordovician rocks of Kentucky based on mappable,
lithologically defined formations. N e w formational n a m e s were proposed,
such as the K o p e Formation, based on type sections in O h i o and Kentucky,
w h i c h r e p l a c e d t h e old Latonia o r E d e n F o r m a t i o n ( W e i s s a n d S w e e t 1964).
S o m e traditional n a m e s , s u c h a s t h e F a i r v i e w (Ford 1967), w e r e r e t a i n e d
a n d given f o r m a l d e f i n i t i o n a s f o r m a t i o n s . I n K e n t u c k y m o s t o f t h e n e w
formations w e r e thick p a c k a g e s o f strata c o m p r i s i n g e q u i v a l e n t s o f older,
t h i n n e r units that either c o u l d n o t b e t r a c e d into K e n t u c k y o r e l s e h a d
b e e n b a s e d o n fossil c o n t e n t ( F i g u r e 4.1). A l s o i n t h e 1960s t h e I n d i a n a
G e o l o g i c a l Survey undertook restudy of the U p p e r O r d o v i c i a n in south-
eastern I n d i a n a . This work p r o d u c e d a revised l i t h o s t r a t i g r a p h i c classifica-
tion in w h i c h a s i n g l e f o r m a t i o n , the D i l l s b o r o , s p a n n e d the M a y s v i l l i a n
a n d R i c h m o n d i a n S t a g e s ( F i g u r e 4.1; B r o w n a n d L i n e b a c k 1966).

A l t h o u g h the n e w research programs yielded stratigraphic units in

k e e p i n g w i t h m o d e r n l i t h o s t r a t i g r a p h i c p r a c t i c e , t h e status o f C i n c i n n a -
tian stratigraphy in t h e late 1960s p r e s e n t e d new c h a l l e n g e s to a n o v e r a l l
u n d e r s t a n d i n g o f C i n c i n n a t i a n g e o l o g i c history. C r e a t i o n o f s u c h t h i c k ,
broadly defined formations tended to obscure m a n y c h a n g e s o c c u r r i n g at
smaller scales of s t r a t i g r a p h i c r e s o l u t i o n , a n d t h e a b u n d a n t fossil c o n t e n t
o f these rocks w a s largely i g n o r e d . A n y a t t e m p t t o m a p t h e C i n c i n n a t i a n
over the entire tri-state r e g i o n r e v e a l e d that f o r m a t i o n s e n d e d a b r u p t l y a t
state lines, r e s u l t i n g in c o n f u s i o n not u n l i k e that of a visitor to t h e city of
C i n c i n n a t i w h o f i n d s that m a n y street n a m e s c h a n g e across m a j o r i n t e r s e c -
tions! O n e m i g h t also b e led t o c o n c l u d e that m a j o r faults f o l l o w e d t h e state
b o u n d a r i e s o r t h e c o u r s e o f t h e O h i o River! S t r a t i g r a p h i c c o l u m n s e s t a b -

Rocks, Fossils, and Time 51

 l i s h e d for t h e C i n c i n n a t i a n s e c t i o n i n e a c h state o f t h e tri-state r e g i o n h a d
to be correlated.

N e w A d v a n c e s in A l t h o u g h t h e early z o n a t i o n o f t h e C i n c i n n a t i a n b a s e d o n s u c h g r o u p s a s

Biostratigraphy b r y o z o a n s a n d b r a c h i o p o d s h a d p r o v e n i n a d e q u a t e for c o r r e l a t i o n very far

o u t s i d e t h e l o c a l C i n c i n n a t i a r e a , b e g i n n i n g in t h e late 1950s, n e w research
into t h e b i o s t r a t i g r a p h y o f t h e C i n c i n n a t i a n e n a b l e d a s i n g l e time-strati-
g r a p h i c f r a m e w o r k t o b e a p p l i e d across t h e e n t i r e r e g i o n . I n order t o u n -
d e r s t a n d h o w this n e w a d v a n c e c a m e a b o u t , w e m u s t c o n s i d e r w h a t char-
a c t e r i s t i c s are r e q u i r e d for a fossil to be a reliable tool for l o n g d i s t a n c e
c o r r e l a t i o n o f strata.
Fossils that are m o s t u s e f u l for c o r r e l a t i o n of strata over great d i s t a n c e s
i d e a l l y m u s t h a v e t w o c h a r a c t e r i s t i c s : a short v e r t i c a l (time) r a n g e and a
w i d e lateral d i s t r i b u t i o n . Fossils h a v i n g a short vertical r a n g e w i l l be g r o u p s
w h o s e p r e s e r v a b l e m o r p h o l o g y e v o l v e s relatively rapidly o v e r t i m e . Fossils
w i t h a w i d e lateral d i s t r i b u t i o n w i l l be g r o u p s that either tolerate a broad
r a n g e o f e n v i r o n m e n t s o r else are c a p a b l e o f w i d e dispersal t h r o u g h a free-
s w i m m i n g larval stage o r a d u l t m o d e o f life. T h e short v e r t i c a l r a n g e o f
m a n y C i n c i n n a t i a n fossils p r o d u c e d a d e t a i l e d b i o s t r a t i g r a p h i c s u b d i v i s i o n
o f t h e strata. H o w e v e r , a s a l r e a d y m e n t i o n e d , m a n y o f t h e fossil z o n e s es-
t a b l i s h e d i n t h e i m m e d i a t e C i n c i n n a t i a r e a c o u l d n o t b e t r a c e d very far.
B e c a u s e l i t h o l o g i e s also c h a n g e d laterally, it is very likely that the fossils
(chiefly b o t t o m - d w e l l i n g b r y o z o a n s a n d b r a c h i o p o d s ) are s t r o n g l y c o n -
trolled by e n v i r o n m e n t a l c h a n g e . M o d e r n t e c h n i q u e s in biostratigraphy
s h o w e d that g r o u p s s u c h a s m i c r o p l a n k t o n i c a l g a e ( c h i t i n o z o a n s a n d acri-
tarchs), c o n o d o n t s ( S w e e t a n d B e r g s t r o m 1971), a n d graptolites ( M i t c h e l l
a n d B e r g s t r o m 1977; G o l d m a n a n d B e r g s t r o m 1997) are better suited for
l o n g d i s t a n c e c o r r e l a t i o n (even i n t e r c o n t i n e n t a l ) o f O r d o v i c i a n strata b e -
c a u s e t h e y h a v e v e r y w i d e d i s p e r s a l . ( M o s t o f t h e s e g r o u p s lived i n the
w a t e r c o l u m n a b o v e t h e sea floor, e i t h e r d r i f t i n g a s p l a n k t o n o r actively-
s w i m m i n g . ) T h e vertical ranges of these groups through the O r d o v i c i a n
t e n d t o b e rather l o n g , s o t h a t t h e y c a n n o t b e u s e d t o s u b d i v i d e t h e C i n c i n -
n a t i a n v e r y e f f e c t i v e l y b e l o w t h e stage l e v e l , b u t t h e y d o form t h e basis o f
m o d e r n t i m e - s t r a t i g r a p h i c c o r r e l a t i o n o f t h e C i n c i n n a t i a n i n relation t o
o t h e r r e g i o n s o f N o r t h A m e r i c a a n d w o r l d w i d e ( W e b b y et al. 2004).

Cycles and D e s p i t e t h e a p p a r e n t m o n o t o n y o f C i n c i n n a t i a n l i m e s t o n e s and shales, m a -

Depositional jor r e p e a t e d patterns in the l i t h o l o g i e s had b e e n r e c o g n i z e d s i n c e the earliest

Sequences w o r k of O r t o n ( O r t o n 1873). Nickles (1902) related these l i t h o l o g i c patterns to

c h a n g e s in t u r b u l e n c e . In 1969 A n s t e y a n d F o w l e r r e c o g n i z e d that a transi-
tion f r o m t h e l i m e s t o n e - r i c h L e x i n g t o n L i m e s t o n e u n d e r l y i n g the C i n c i n -
n a t i a n t o t h e s h a l e - d o m i n a t e d E d e n S h a l e (now the K o p e F o r m a t i o n ) trace-
able throughout the tri-state area of O h i o , Indiana, and Kentucky,
c o r r e s p o n d e d to a transition from s h a l l o w water, m o r e t u r b u l e n t c o n d i t i o n s
to d e e p e r water, m o r e q u i e s c e n t c o n d i t i o n s . In s h a l l o w water, greater w a v e -
g e n e r a t e d t u r b u l e n c e was e x p e c t e d to p r e v e n t s e t t l e m e n t of fine-grained sedi-

52 A Sea without Fish

 Figure 4.4. Cincinnatian
shoaling-upward cycles.
From Tobin (1982, figure
74). The base of the Cin-
cinnatian is at the base of
the 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
are: brick pattern = lime-
stone, dashed pattern =
shale, wavy pattern =
wavy-bedded limestone,
rhombic pattern = dolo-
mite, small circles = nod-
ular limestone. More re-
cent work by Holland has
recognized six major
shoaling-upward cycles
or sequences over the
same stratigraphic inter-
merits, l e a v i n g coarser, shelly s e d i m e n t , w h i l e in d e e p e r water, fine-grained val in which Tobin delin-
s e d i m e n t s , chiefly clay particles, are m o r e likely to settle to t h e b o t t o m . A eated three cycles (see
return to s h a l l o w e r water w a s m a r k e d by i n c r e a s i n g l i m e s t o n e in t h e overly- Figure 15.1).

i n g F a i r v i e w to B e l l e v u e , f o l l o w e d by a d e e p e n i n g in t h e s h a l e - r i c h C o r -
ryville a n d a s h a l l o w i n g i n the l i m e s t o n e o f t h e M t . A u b u r n . D e e p e n i n g
again o c c u r r e d in the shales A r n h e i m to Waynesville, followed by a shallow-
i n g with i n c r e a s i n g l i m e s t o n e in the Liberty to S a l u d a , and a slight d e e p e n -
ing in t h e u p p e r m o s t W h i t e w a t e r interval. Hay (1981) a n d T o b i n (1982) r e c o g -
n i z e d b a s i c a l l y t h e s a m e c y c l i c p a t t e r n s ( F i g u r e s 4 . 4 , 4.5). Thus, major
c h a n g e s in sea level c o u l d a c c o u n t for t h e l a r g e - s c a l e p a t t e r n s of r e p e a t e d
lithologies o f the C i n c i n n a t i a n S e r i e s . T h e r e are several possible c a u s e s for
these major c h a n g e s in sea level, i n c l u d i n g t e c t o n i c e v e n t s in t h e A p p a l a -
c h i a n o r o g e n i c belt to t h e cast, fluctuations in s e d i m e n t supply, a n d g r o w t h
o f ice sheets i n t h e s o u t h e r n p o l a r r e g i o n s d u r i n g the L a t e O r d o v i c i a n .
The m o d e r n c o n c e p t s o f s e q u e n c e s t r a t i g r a p h y u s e t h e i m p o r t a n c e
o f sea level c h a n g e s t o e x p l a i n t h e p a t t e r n s o f r e p e a t e d d e p o s i t i o n i n t h e

Rocks, Fossils, and Time 53

Figure 4 . 5 . Geological
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.

54 A Sea without Fish

 Figure 4.6. A. Three
scales of Cincinnatian
stratigraphic cyclicity.
From Tobin (1982, figure
30). In storm cycle, basal
bed 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-
ters. Tapering wedges
beside each column indi-
cate direction of shallow-
ing (wider = shallower).
Distal facies contain ma-
terial transported farther
from place of formation
than proximal fades.

C i n c i n n a t i a n , a s w e l l a s t h e m a r k e d lateral c h a n g e s that h a d s o g r e a t l y
frustrated earlier efforts to a c h i e v e a s i n g l e s t r a t i g r a p h i c f r a m e w o r k for t h e
entire o u t c r o p r e g i o n . B e c a u s e t h e O r d o v i c i a n sea floor w a s a g e n t l e r a m p
that g e n e r a l l y b e c a m e d e e p e r from s o u t h t o n o r t h , p r e s u m a b l y e u s t a t i c
(global) c h a n g e s i n sea level b r o u g h t a b o u t d i f f e r e n t d e p o s i t i o n a l c o n d i -
tions over different parts of the r e g i o n at the s a m e t i m e . T h e a b r u p t transi-
tions from shallow t o d e e p e r w a t e r m a r k s e q u e n c e b o u n d a r i e s that c a n b e
used t o c o r r e l a t e s e c t i o n s l o c a t e d o n different r e g i o n s o f t h e p a l e o s l o p e .
H o l l a n d (1993, 1997, 1998) r e c o g n i z e d six m a j o r d e p o s i t i o n a l s e q u e n c e s
w i t h i n the C i n c i n n a t i a n that c u t across l i t h o s t r a t i g r a p h i c c o n t a c t s (see
F i g u r e 15.1). T h e C i n c i n n a t i a n c o m p r i s e s mainly the l o w e r four s e q u e n c e s .
T h e C1 sequence (Edenian) comprises the K o p e Formation in the C i n c i n -
nati area. T h e C2 s e q u e n c e ( M a y s v i l l i a n ) c o m p r i s e s t h e F a i r v i e w - B e l l e v u e
s e q u e n c e , and the C3, C o r r y v i l l e - M t . A u b u r n s e q u e n c e . The R i c h m o n -

Rocks, Fossils, and Time 55

Figure 4.7. A. Storm
deposition and erosion
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 =
rare.

dian consists o f three s e q u e n c e s : C 4 , S u n s e t - O r e g o n i a , C 5 , Waynesville-

Liberty-Whitewater, and C 6 , U p p e r W h i t e w a t e r - E l k h o r n , truncated by the
e r o s i o n a l u n c o n f o r m i t y w i t h the S i l u r i a n .
W i t h i n t h e s e six m a j o r C i n c i n n a t i a n s e q u e n c e s , c y c l i c p a t t e r n s o f
stratification a l s o o c c u r o n s m a l l e r s c a l e s ( f i g u r e 4.6A). C o u p l e t s o f l i m e -
s t o n e a n d s h a l e b e d s on a s c a l e less t h a n o n e m e t e r t h i c k are very c h a r a c -
teristic of m u c h of t h e C i n c i n n a t i a n . 1 lay (1981) a n d T o b i n (1982) first rec-
o g n i z e d that a t least s o m e o f t h e s e c o u p l e t s w e r e t h e p r o d u c t o f storm
p r o c e s s e s ( F i g u r e s 4 . 6 A , 4.7). T h e s h a l l o w , s u b t r o p i c a l C i n c i n n a t i a n sea
floor m u s t h a v e b e e n s w e p t f r e q u e n t l y b y large s t o r m s , e v e n h u r r i c a n e s .
D u r i n g t h e s e s t o r m s , w a v e s a n d s t r o n g b o t t o m c u r r e n t s d i s r u p t e d sea floor
c o m m u n i t i e s o f o r g a n i s m s , s w e e p i n g f i n e - g r a i n e d s e d i m e n t s into s u s p e n -
sion w h i l e l e a v i n g shells a n d o t h e r skeletal d e b r i s a s c o a r s e c a l c a r e o u s l a g
d e p o s i t s . A s the s t o r m s u b s i d e d , f i n e - g r a i n e d s e d i m e n t s settled o n t o t h e sea
floor, s m o t h e r i n g c o m m u n i t i e s a n d f o r m i n g what w o u l d b e c o m e a s h a l e
b e d o v e r l y i n g a fossiliferous l i m e s t o n e (see F i g u r e 15.2). T o b i n (1982) r e c o g -
n i z e d that t h e C i n c i n n a t i a n c o n t a i n s a w i d e variety o f t h e s e s t o r m c y c l e s ,
s o m e c o n s i s t i n g of s i m p l y a s h a l e b e d o v e r h a n g a l i m e s t o n e , in a d d i t i o n to
o t h e r m o r e c o m p l e x p a t t e r n s ( F i g u r e 4.7B). S o m e p a l e o e c o l o g i s t s like Har-
ris and M a r t i n (1979) e n v i s i o n e d an e c o l o g i c a l s u c c e s s i o n f o l l o w i n g such
storm e v e n t s , i n w h i c h s h e l l y o r g a n i s m s r e p o p u l a t e d the m u d d y b o t t o m
and were in turn colonized by encrusting and attached epifauna (Figure
4.8). M a n y c a s e s o f c o m p l e t e l y p r e s e r v e d fossils s u c h a s c r i n o i d s , edrioast-
e r o i d s , a n d trilobites i n t h e C i n c i n n a t i a n are u n d o u b t e d l y the p r o d u c t o f

56 A Sea without Fish

 Figure 4 . 8 . Colonization
of a soft mud substratum
by Dalmanella (low di-
versity) is followed by the
development of a shell
pavement and succession
of erect bryozoans.
Higher diversity is
achieved in the mature
community by crinoids
growing on a firm sub-
stratum. From Harris and
Martin (1979) and re-
printed by permission of
MATURE COMMUNITY
the Society for Sedimen-
tary Geology.

SUCCESSION

COLONIZATION

rapid b u r i a l b y f i n e - g r a i n e d s e d i m e n t s d u r i n g s t o r m e v e n t s ( B r a n d t 1985;
M e y e r 1990; S c h u m a c h e r a n d S h r a k e 1997; H u g h e s a n d C o o p e r 1999).
I n m a n y parts o f t h e C i n c i n n a t i a n s e c t i o n , p a r t i c u l a r l y t h e K o p e For-
m a t i o n , c y c l i c i t y at a s c a l e of a b o u t o n e to t h r e e m e t e r s is v e r y a p p a r e n t as
o n e passes l o n g h i g h w a y r o a d c u t s . D i f f e r e n t w o r k e r s h a v e i n t e r p r e t e d this
cyclicity in d i f f e r e n t w a y s . H a y (1981) r e c o g n i z e d a r e g u l a r s p a c i n g of
c o a r s e - g r a i n e d l i m e s t o n e b u n d l e s ( F i g u r e 4.6B). T o b i n (1982) t e r m e d t h i s

Rocks, Fossils, and Time 57

58 A Sea without Fish
s c a l e of cyclicity t h e m e g a c y c l e , a n d d e s c r i b e d it as a " f i n i n g - u p w a r d " Figure 4.9. Kope cyclicity
p a c k e t h a v i n g a basal c o m p o n e n t o f c o a r s e - g r a i n e d l i m e s t o n e ( g r a i n s t o n e ) , and Kope-Fairview for-

a middle c o m p o n e n t with thin interbedded limestones (packstones) and mational contact trace-
able over broad area of
shales, a n d a n u p p e r s h a l e - r i c h c o m p o n e n t ( F i g u r e s 4 . 6 A , B). J e n n e t t e a n d
Cincinnati Arch.
Pryor (1993) felt that t h e c y c l e w a s " c o a r s e n i n g - u p w a r d , " b e g i n n i n g w i t h
A. Geological Society of
shale, o v e r l a i n b y i n t e r b e d d e d t h i n l i m e s t o n e a n d s h a l e , a n d c a p p e d b y
America Field Trip, 1981.
c o a r s e - g r a i n e d l i m e s t o n e ( F i g u r e 4.6B). I n their i n t e r p r e t a t i o n t h e c y c l e Participants examining
w a s of a regressive n a t u r e , c o r r e s p o n d i n g to s h o r t - t e r m fluctuations of sea meter-scale cyclicity in
level. M o r e recent s t u d i e s b y H o l l a n d a n d o t h e r s (1997) h a v e c o n t i n u e d t o the Kope Formation
r e c o g n i z e storm-related f e a t u r e s w i t h i n t h e s e m e t e r - s c a l e c y c l e s , b u t n o t e along Kentucky Route
445 near the Ohio River,
that n o single pattern o f c o a r s e n i n g o r f i n i n g u p w a r d i s d o m i n a n t . T h e h i g h
Campbell County, Ken-
variability in c y c l e c h a r a c t e r p r o b a b l y reflects a c o m p l e x i n t e r p l a y of sea
tucky. B. Contact of
level c h a n g e and fluctuations in s t o r m intensity a n d f r e q u e n c y . A r e m a r k -
Kope Formation with
able result of all t h e s e s t u d i e s is that d e s p i t e t h e l o n g - h e l d v i e w that i n d i - overlying Fairview Forma-
vidual strata w i t h i n the C i n c i n n a t i a n are n o t laterally persistent, m e t e r - s c a l e tion (arrow) along Rad-
cycles w i t h i n the K o p e F o r m a t i o n c a n i n d e e d b e c o r r e l a t e d o v e r v i r t u a l l y cliff Drive, Cincinnati,
the entire o u t c r o p area for tens of k i l o m e t e r s (Brett a n d A l g e o 2001b; H o l - Ohio. Photo by Paul E.
l a n d , M i l l e r , and M e y e r 2001; W e b b e r 2002). V a r i a t i o n s i n p r o p o r t i o n s o f Potter.

limestone to shale within the K o p e and Fairview Formations define e v e n

larger-scale c y c l e s , on t h e order of 1 0 - 2 0 m t h i c k ( " d e c a m e t e r - s c a l e c y c l e s " )
that e n a b l e lateral c o r r e l a t i o n o v e r l o n g d i s t a n c e s ( F i g u r e 4.9; J e n n e t t e a n d
Pryor 1993; D a t t i l o 1996; Holland et al. 1997; Brett a n d A l g e o 2001b).

T h e m o s t r e c e n t s t u d i e s o f t h e C i n c i n n a t i a n h a v e , i n a s e n s e , c o m e full Tracing Biofacies

circle in r e t u r n i n g to its a b u n d a n t fossil c o n t e n t as a k e y to u n d e r s t a n d i n g and Event Beds
h o w C i n c i n n a t i a n strata w e r e d e p o s i t e d . I n c o n t r a s t t o earlier lithostrati-
g r a p h i c studies i n w h i c h t h i c k u n i t s w e r e d e f i n e d o n t h e basis o f l i m e s t o n e -
to-shale ratios with few internal s u b d i v i s i o n s , t h e latest s t u d i e s arc b a s e d
on h i g h r e s o l u t i o n , b e d - b y - b e d l o g g i n g of s t r a t i g r a p h i c s e c t i o n s in t e r m s of
lithologic as well as paleontologic features. In the K o p e F o r m a t i o n , l o n g
regarded as a s i n g l e , t h i c k a n d m o n o t o n o u s s h a l e - r i c h u n i t , h i g h r e s o l u t i o n
c e n s u s e s o f fossil a s s e m b l a g e s ( b i o f a c i e s ) r e v e a l e d t r e n d s that are n o t i m -
mediately obvious from lithologic features (lithofacies) (Holland, Miller,
M e y e r , and D a t t i l o 2001). T h e s e b i o f a c i e s t r e n d s a p p e a r t o reflect d e p t h -
related f a u n a l p r e f e r e n c e s a n d c a n b e t r a c e d w i t h i n t h e C i n c i n n a t i a r e a
a n d also e a s t w a r d t o M a y s v i l l e , K e n t u c k y ( H o l l a n d e t al. 2001; W e b b e r
2002). W i t h i n a n o t h e r shale-rich u n i t , the M i a m i t o w n S h a l e , D a t t i l o (1996)
also d e m o n s t r a t e d t h e utility o f fossil a s s e m b l a g e s , t o g e t h e r w i t h l i t h o f a c i e s
analysis, in c o r r e l a t i o n of m e t e r - s c a l e c y c l i c i t y w i t h i n a 30 km radius.
Earlier C i n c i n n a t i a n workers r e c o g n i z e d m a n y t h i n b e d s c h a r a c t e r i z e d
by an a b u n d a n c e of a p a r t i c u l a r fossil s p e c i e s a n d restricted in stratigraphic
o c c u r r e n c e . T h e s e are m a r k e d i n the traditional b i o s t r a t i g r a p h y section ( D a -
vis 1992) and are well k n o w n a m o n g local c o l l e c t o r s , yet f o u n d little use dur-
ing the phase of C i n c i n n a t i a n Lithostratigraphy. Exceptions were the "shin-
gled Rafinesquina" beds in the M i a m i t o w n S h a l e a n d a thin z o n e of a b u n d a n t
Platystrophia ponderosa o c c u r r i n g at the base of t h e M t . A u b u r n that w e r e
used by Ford (1967) as m a r k e r b e d s that w e r e useful for correlation.

Rocks, Fossils, and Time 59

 I n r e c e n t years several p a l e o e c o l o g i c a n d t a p h o n o m i c s t u d i e s b r o u g h t
r e n e w e d a t t e n t i o n to t h i n , fossil-rich h o r i z o n s that are t r a c e a b l e over a w i d e
a r e a . Frey (1987b) t r a c e d t h e n a u t i l o i d - r i c h Treptoceras duseri s h a l e w i t h i n
the W a y n e s v i l l e Formation in Warren and C l i n t o n C o u n t i e s , O h i o , and
c o r r e l a t e d it w i t h a lithologically e q u i v a l e n t trilobite-rich s h a l e to the west
in I n d i a n a . A s t r o p h o m e n i d b r a c h i o p o d shell p a v e m e n t e n c r u s t e d by the
s a m e t h r e e e d r i o a s t e r o i d s p e c i e s o c c u r s i n the u p p e r C o r r y v i l l e f o r m a t i o n
at F l o r e n c e , K e n t u c k y , a n d a g a i n 22 km to t h e n o r t h n e a r the I-275 b e l t w a y
n o r t h w e s t o f C i n c i n n a t i ( M e y e r 1990). I n t h e u p p e r K o p e f o r m a t i o n , a
persistent b e d o f c a l c a r e o u s siltstone r e p l e t e w i t h t h e U - s h a p e d trace fossil
Diplocraterion p r o v i d e d a k e y m a r k e r h o r i z o n for c o r r e l a t i o n from n o r t h -
e r n K e n t u c k y t o s o u t h w e s t e r n O h i o ( J e n n e t t e a n d Pryor 1993). S e v e r a l
e x a m p l e s o f p a l e o n t o l o g i c a l e v e n t h o r i z o n s i n t h e C i n c i n n a t i a n are dis-
c u s s e d i n Brett a n d B a i r d (1997). T h e s e i n c l u d e : a t r a c e a b l e storm h o r i z o n
( M i l l e r 1997); a o n e - m e t e r t h i c k interval c h a r a c t e r i z e d by the b r a c h i o p o d
Onniella meeki t r a c e a b l e 135 km f r o m O h i o into I n d i a n a (Frey 1997b); the
R i c h m o n d i a n f a u n a l " i n v a s i o n " ( H o l l a n d 1 9 9 7 ) ; a n d Isotelus-bearing s h a l e s
in the W a y n e s v i l l e f o r m a t i o n that c a n be t r a c e d for at least 40 km across
s o u t h w e s t e r n O h i o ( S c h u m a c h e r a n d S h r a k e 1997). D a t t i l o (1996) u s e d the
restricted o c c u r r e n c e of t h e b r a c h i o p o d Heterorthina fairmountensis as a
d a t u m for c o r r e l a t i o n o f t h e M i a m i t o w n S h a l e f r o m s o u t h w e s t e r n O h i o t o
n o r t h e r n K e n t u c k y . Brett a n d A l g e o (2001b) d i s c u s s e d several o t h e r k e y
f o s s i l b e d s o r e p i b o l e s i n t h e K o p e a n d f a i r v i e w f o r m a t i o n s that facilitate
c o r r e l a t i o n o v e r a w i d e area i n O h i o a n d K e n t u c k y . A l l o f t h e s e m a r k e r
b e d s are e x c e p t i o n a l for their w i d e g e o g r a p h i c d i s t r i b u t i o n , b u t h a v e a va-
riety o f c a u s e s . S o m e r e s u l t e d f r o m c o n d i t i o n s favorable t o a p a r t i c u l a r
s p e c i e s for a short t i m e . O t h e r s reflect a w i d e s p r e a d event s u c h as a storm
that s m o t h e r e d the sea floor o v e r a w i d e area. The w i d e e x t e n t of these b e d s
is also related to t h e v e r y g r a d u a l c h a n g e in d e p t h o v e r t h e e n t i r e r e g i o n ,
c r e a t i n g vast a r e a s o f s i m i l a r d e p t h a n d b o t t o m type.
O t h e r t y p e s of e v e n t h o r i z o n s in addition to z o n e s of restricted fossil
o c c u r r e n c e are also w i d e l y t r a c e a b l e across the C i n c i n n a t i A r c h region. I n
p a r t i c u l a r , Jennette a n d Pryor (1993) u s e d a b e d with w e l l - d e v e l o p e d basal
g u t t e r casts as a d a t u m for c y c l e c o r r e l a t i o n n e a r the top of the K o p e f o r m a -
tion i n t h e v i c i n i t y o f C i n c i n n a t i a n d n o r t h e r n K e n t u c k y . C u t t e r casts are
s e d i m e n t f i l l i n g s o f t r o u g h s e r o d e d into t h e sea floor. M o r e recently Brett
a n d A l g e o (2001b) d e m o n s t r a t e d the utility of several l\ pes of event h o r i z o n s
in the C i n c i n n a t i a n . T e m p e s t i t e s or storm b e d s i n c l u d e a variety of s e d i m e n -
tary features s u c h as s h i n g l e d b r a c h i o p o d b e d s , rippled b e d s , g r a d e d b e d s ,
g u t t e r casts, a n d s m o t h e r e d b o t t o m o r o b r u t i o n d e p o s i t s a l l p r o d u c t s o f
short-term b u t w i d e s p r e a d e p i s o d e s o f intense d i s t u r b a n c e o f the sea f l o o r .
O t h e r e v e n t h o r i z o n s i n c l u d e distinctive trace fossil h o r i z o n s (the a f o r e m e n -
tioned Diplocraterion), b r a c h i o p o d shell p a v e m e n t s , c o n c r e t i o n a r y layers,
h a r d g r o u n d s ( b e d s f o r m e d by early c e m e n t a t i o n on the sea floor), and beds
indicative of s e i s m i c e v e n t s . F e a t u r e s possibly c a u s e d by seismic d e f o r m a t i o n
o f t h e sea floor i n c l u d i n g b e d s w i t h i n t e n s e internal d e f o r m a t i o n and so-
c a l l e d b a l l a n d p i l l o w s t r u c t u r e s o c c u r over a w i d e area i n s o u t h e r n O h i o
and n o r t h e r n K e n t u c k y ( S c h u m a c h e r 1992; P o p e et al. 1997). T h e s e s e i s m i t e s

60 A Sea without Fish

m a y have resulted from e a r t h q u a k e s c a u s e d by t e c t o n i c activity in the rising
A p p a l a c h i a n m o u n t a i n s far to t h e east. U s i n g all of t h e s e e v e n t h o r i z o n s ,
Brett and A l g e o (2001a) w e r e able to correlate d e c a m e t e r a n d m e t e r - s c a l e
cycles w i t h i n the K o p e a n d F a i r v i e w F o r m a t i o n s for d i s t a n c e s up to 80 km
a l o n g the AA Highway ( K e n t u c k y R o u t e 9) in n o r t h e r n K e n t u c k y .
To a certain d e g r e e , the m o s t r e c e n t studies of C i n c i n n a t i a n stratigraphy
c o n f i r m and v i n d i c a t e the heavy e m p h a s i s that was p l a c e d on fossil c o n t e n t
by g e n e r a t i o n s of earlier workers in the C i n c i n n a t i a n . C l e a r l y , no study of
C i n c i n n a t i a n stratigraphy c a n afford to ignore the p l e t h o r a of i n f o r m a t i o n
offered by the a b u n d a n c e and diversity of fossils t h r o u g h o u t t h e r e g i o n . At
the t i m e of this w r i t i n g , efforts to d e v e l o p a h i g h l y d e t a i l e d r e g i o n a l strati-
g r a p h i c framework for the C i n c i n n a t i a n rely on a synthesis of all e v i d e n c e
available, i n c l u d i n g l i t h o l o g y , s e d i m e n t o l o g y , a n d p a l e o n t o l o g y . S t u d i e s in-
c o r p o r a t i n g this "total e v i d e n c e " a p p r o a c h i n c l u d e those o f D a t t i l o (1996),
Holland (1993 Holland (1997) Holland. M i l l e r , a n d M e y e r (2001), Brett a n d
A l g e o (2001b), W e b b e r (2002), a n d M c L a u g h l i n a n d Brett (2007).
T h r o u g h these studies, t h e r e c o g n i t i o n that the C i n c i n n a t i a n i s c o n -
structed of a series of stratal " p a c k a g e s " d e l i m i t e d by relatively short-term sea
level c h a n g e s that m a r k s e q u e n c e b o u n d a r i e s also to s o m e extent revives t h e
old c o n c e p t of the stratigraphic " l a y e r c a k e " that typified m u c h of the origi-
nal work on the C i n c i n n a t i a n ( A l g e o and Brett 2001). C l e a r l y the C i n c i n -
natian is not m e r e l y a j u m b l e d mosaic- of c a r b o n a t e and shale f a c i e s with
poor lateral continuity. A l t h o u g h lateral facies variations do exist, they c a n
b e u n d e r s t o o d i n the c o n t e x t o f r e g i o n a l p a l e o b a t h y m e t r y a n d s e q u e n c e
architecture. S e q u e n c e , c y c l i c , and e v e n t stratigraphy offer great p r o m i s e for
d e v e l o p m e n t of a h i g h resolution t i m e - s t r a t i g r a p h i c f r a m e w o r k for t h e C i n -
c i n n a t i a n in w h i c h many interesting q u e s t i o n s of e v o l u t i o n a r y p a l e o n t o l o g y ,
p a l e o e c o l o g y , and s e d i m e n t o l o g y c a n b e addressed.

How old arc the fossils of the C i n c i n n a t i a n ? H o w l o n g a g o did the C i n c i n - A g e of t h e

natian A g e b e g i n , and w h e n did i t e n d ? H o w m u c h t i m e d o e s a n y part o f t h e Cincinnatian
C i n c i n n a t i a n S e r i e s a f o r m a t i o n , a s e q u e n c e , or a s i n g l e b e d r e p r e s e n t ?
T h e s e are questions of the absolute a g e of C i n c i n n a t i a n fossils a n d strata.
To say that the C i n c i n n a t i a n is L a t e O r d o v i c i a n in a g e is to m a k e a
statement a b o u t its relative a g e . The g e o l o g i c t i m e scale of e o n s , eras, a n d
periods was c o n s t r u c t e d d u r i n g the n i n e t e e n t h c e n t u r y o n t h e basis o f super-
position and fossil s u c c e s s i o n . Initially, p i o n e e r i n g g e o l o g i s t s like the E n g -
l i s h m a n W i l l i a m S m i t h r e c o g n i z e d that fossils c o u l d b e used t o c h a r a c t e r i z e
strata and correlate t h e m from o n e region to a n o t h e r . T h r o u g h the w o r k of
Smith and other pioneers like C u v i e r and B r o g n i a r t in F r a n c e t h e geologic-
succession of strata was established by r e c o g n i z i n g that strata with character-
istic sets of fossils always o c c u r r e d in the s a m e vertical s e q u e n c e f r o m o l d e s t
to y o u n g e s t . T h e p r i n c i p l e of o r g a n i c e v o l u t i o n as later put forth by C h a r l e s
D a r w i n and others was u n k n o w n t o t h e s e f o u n d e r s o f g e o l o g y , b u t t h e y w e r e
able to use the c h a n g i n g m a k e u p of lite on E a r t h as a relative g a u g e of a n -
cient history. O r g a n i c e v o l u t i o n is the u n d e r l y i n g m e c h a n i s m that provides
a directional c o m p o n e n t , a sort of c l o c k , by w h i c h to p l a c e fossils a n d their

Rocks, Fossils, and Time 61

 e n c a s i n g rocks o n the g e o l o g i c t i m e scale. H o w e v e r , the rate o f e v o l u t i o n a r y
c h a n g e varies greatly a m o n g l i v i n g o r g a n i s m s , and s o m e groups show marked
c h a n g e t h r o u g h a vertical s u c c e s s i o n of strata w h i l e others do not; c o n s e -
q u e n t l y , different g r o u p s h a v e different c l o c k s . E v o l u t i o n a r y c h a n g e thus
c a n n o t p r o v i d e a u n i f o r m m e a s u r e of t i m e by w h i c h to e s t i m a t e the absolute
a g e of fossils (despite t h e m o d e r n use of " m o l e c u l a r c l o c k s " to set the a g e for
c o m m o n a n c e s t o r s o f different l i v i n g groups).
A b s o l u t e a g e d a t i n g o f a n c i e n t r o c k s relies o n t h e c o n s t a n t rate o f ra-
dioactive decay of unstable isotopes of certain e l e m e n t s contained in m i n -
erals f o u n d i n v o l c a n i c a n d o t h e r t y p e s o f i g n e o u s rocks. B e c a u s e t h e s e
m i n e r a l s d o n o t u s u a l l y f o r m i n t h e shells a n d s k e l e t o n s o f o r g a n i s m s o r i n
t h e s e d i m e n t s , d i r e c t d a t i n g of fossils a n d most s e d i m e n t a r y rocks is very
difficult. A t b e s t , w e c a n h o p e t o f i n d d a t a b l e layers s u c h a s v o l c a n i c ash
b e d s or lava flows interbedded w i t h fossiliferous strata. It m a y t h e n be pos-
sible to state that a f o s s i l - b e a r i n g layer lies e i t h e r a b o v e or b e l o w a dated
h o r i z o n , m a k i n g t h e fossil e i t h e r relatively y o u n g e r o r older. I n t h e m a r i n e
s e d i m e n t a r y r e c o r d t h e b e s t o p p o r t u n i t i e s for this m e t h o d o f a g e d e t e r m i -
nation c o m e from u n i q u e potassium-rich clay beds k n o w n as K-bentonites,
f o r m e d t h r o u g h c h e m i c a l a l t e r a t i o n o f v o l c a n i c ash falls into t h e sea.
K - b e n t o n i t e s c o n t a i n biotites a n d z i r c o n s that are d a t a b l e u s i n g u r a n i u m -
lead and potassium-argon dating techniques.
I n N o r t h A m e r i c a , ash b e d s p r e s e r v e d a s K - b e n t o n i t e s are f o u n d
t h r o u g h o u t t h e O r d o v i c i a n b u t are p a r t i c u l a r l y well k n o w n from t h e M i d -
d l e O r d o v i c i a n ( H u f f e t al. 1992). The closest dated K-bentonite to the
C i n c i n n a t i a n is a remarkable bed k n o w n as the Millbrig K-bentonite oc-
curring at the base of the M i d d l e Ordovician Lexington Limestone of
K e n t u c k y O r d o v i c i a n ( H u f f e t al. 1992). T h e M i l l b r i g i s f o u n d t h r o u g h o u t
m u c h of the eastern m i d - c o n t i n e n t of North A m e r i c a and is equivalent to
a n ash b e d f o u n d w i d e l y i n w e s t e r n E u r o p e . R a d i o m e t r i c a g e d a t e s ( U - P b
and 4 0 A r - A r m e t h o d s ) o f a b o u t 4 5 4 m i l l i o n y e a r s for this b e d i n N o r t h
3 9

A m e r i c a a n d E u r o p e i n d i c a t e that i t r e p r e s e n t s o n e o f t h e largest ash-pro-

d u c i n g volcanic eruptions k n o w n in the Phanerozoic record. The eruption
took p l a c e d u r i n g t h e T a c o n i a n o r o g e n i c e v e n t a s the a n c i e n t c o n t i n e n t s
Baltica and Laurentia collided, closing the ancient Iapetus Sea. On the
basis o f this d a t e d h o r i z o n , t h e C i n c i n n a t i a n A g e b e g a n s o m e t i m e m o r e
r e c e n t l y t h a n 454 m i l l i o n y e a r s a g o .
U n f o r t u n a t e l y for a c l o s e r d a t i n g of t h e C i n c i n n a t i a n , K-bentonites are
scarce in the U p p e r O r d o v i c i a n in both N o r t h A m e r i c a and E u r o p e (Berg-
s t r o m , pers. c o m m . ) , p r o b a b l y r e f l e c t i n g a lull i n t e c t o n i c activity b e t w e e n
the T a c o n i a n and A c a d i a n (Middle Devonian) orogenic (mountain build-
ing) pulses. H o w e v e r , a z o n e of two or three impure K-bentonites, k n o w n
a s t h e B e a r C r e e k K - b e n t o n i t e z o n e , o c c u r s w i t h i n t h e Point P l e a s a n t For-
m a t i o n f r o m 12.5 m to 13.7 m a b o v e t h e L e x i n g t o n / P o i n t Pleasant c o n t a c t ,
a n d just below the b a s e o f t h e C i n c i n n a t i a n ( S c h u m a c h e r a n d C a r l t o n
1991). This z o n e has b e e n i d e n t i f i e d i n c o r e s i n O h i o and n o r t h e r n K e n -
t u c k y , a n d i n t h e B e a r C r e e k Q u a r r y , C l e r m o n t C o u n t y , O h i o , b u t has n o t
b e e n dated, as it contains no zircons. The only bentonite k n o w n to o c c u r
w i t h i n t h e C i n c i n n a t i a n itself w a s r e c o r d e d i n the l o w e r R i c h m o n d i a n

62 A Sea without Fish

from a d r i l l c o r e i n S e n e c a C o u n t y , O h i o ( B e r g s t r o m a n d M i t c h e l l 1992),
but it, t o o , has not b e e n d a t e d .
B y u s i n g r a d i o m e t r i c dates f r o m o t h e r r e g i o n s t o c a l i b r a t e t h e t i m e -
stratigraphic record b a s e d on r a n g e s of c o n o d o n t s a n d graptolites, it is p o s -
sible to p l a c e further constraints on the C i n c i n n a t i a n t i m e 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 b o u t 488.3 +/-1.7 m i l l i o n years
a g o and its e n d at 443.7 +/-1.5 m i l l i o n years a g o (International C o m m i s s i o n
o n Stratigraphy 2004). The t i m e scale p r o d u c e d b y t h e U N E S C O - s p o n s o r e d
International G e o l o g i c a l C o r r e l a t i o n P r o g r a m m e ( W e b b y , C o o p e r , B e r g -
strom, and Paris 2004) places the b e g i n n i n g of t h e C i n c i n n a t i a n at a b o u t
452.5 m i l l i o n years ago. H o w e v e r , the top of the section in the C i n c i n n a t i
A r c h region d o e s not i n c l u d e the u p p e r m o s t O r d o v i c i a n record b e c a u s e o f
pre-Silurian erosion, and an entire stage, k n o w n as the H i r n a n t i a n , is miss-
ing. A c c o r d i n g t o the t i m e s c a l e p r o v i d e d b y W e b b y , C o o p e r , B e r g s t r o m , a n d
Paris (2004) the base of t h e H i r n a n t i a n is at a b o u t 445 m i l l i o n years a g o .
These c o n s i d e r a t i o n s yield an approximate duration for t h e Edenian,
M a y s v i l l i a n , and R i c h m o n d i a n of a b o u t 7.5 m i l l i o n years (452.5 my445 my).
The fact that dates for n e i t h e r the lower or u p p e r b o u n d a r i e s of the C i n c i n -
natian c o m e from s a m p l e s in the local r e g i o n s h o u l d alert t h e reader for
future refinements! H o w e v e r , t h e dates a n d 7.5 m i l l i o n year e s t i m a t e s e e m
to be the best i n f o r m a t i o n c u r r e n t l y available.
H o l l a n d a n d P a t z k o w s k y (1996) c a l i b r a t e d b i o s t r a t i g r a p h i c z o n a t i o n o f
the C i n c i n n a t i a n b a s e d on c o n o d o n t s ( S w e e t 1984) w i t h a r a d i o m e t r i c a l l y
dated K - b e n t o n i t e t o d e r i v e e s t i m a t e s o f t h e d u r a t i o n o f t h e six C i n c i n n a -
tian stratigraphic s e q u e n c e s . B y their e s t i m a t e s , t h e d u r a t i o n o f t h e C 1 se-
q u e n c e ( E d e n i a n ) w a s a b o u t 2.5 m i l l i o n y e a r s , t h a t o f t h e C 2 s e q u e n c e
( F a i r v i e w - B e l l e v u e ) a b o u t 1.7 m i l l i o n y e a r s , t h a t o f t h e C 3 s e q u e n c e
( C o r r y v i l l e - M t A u b u r n ) a b o u t 0.6 m i l l i o n y e a r s , that o f t h e C 4 ( S u n s e t -
O r e g o n i a ) s e q u e n c e a b o u t 0.8 m i l l i o n y e a r s , t h a t o f t h e C 5 s e q u e n c e
( W a y n e s v i l l e - L i b e r t y - W h i t e w a t e r ) a b o u t 1.5 m i l l i o n y e a r s , a n d that o f t h e
C 6 s e q u e n c e ( U p p e r W h i t e w a t e r - E l k h o r n ) a b o u t 0.7 m i l l i o n y e a r s . T h e i r
total duration for the six s e q u e n c e s is 7.8 m i l l i o n y e a r s v e r y c l o s e to t h e
m o r e r e c e n t e s t i m a t e of 7.5 m i l l i o n y e a r s .
If the C i n c i n n a t i a n c a n be b r a c k e t e d in t i m e b e t w e e n 452 a n d 445 m i l -
lion years a g o , is it possible to assign dates to s u b d i v i s i o n s of t h e C i n c i n n a -
tian, and to d e t e r m i n e the a m o u n t of t i m e represented by i n d i v i d u a l c y c l e s
or beds? Unless any directy datable c o m p o n e n t s , s u c h as ash b e d s , are f o u n d
w i t h i n the C i n c i n n a t i a n , n o absolute a g e s c a n b e d e t e r m i n e d . Instead, w e
arc forced to rely on a p r o c e s s of i n t e r p o l a t i o n , w o r k i n g up f r o m t h e earliest
r a d i o m e t r i c a l l y d e t e r m i n e d c a l i b r a t i o n p o i n t o r d o w n from t h e t e r m i n a l
calibration point. A s s u m i n g that s e d i m e n t a t i o n rates w e r e c o n s t a n t d u r i n g
the C i n c i n n a t i a n , o n e c o u l d assign i n t e r m e d i a t e dates b a s e d o n stratal thick-
ness. However, there is a b u n d a n t e v i d e n c e from studies of m o d e r n s e d i m e n -
tation rates that the a s s u m p t i o n of u n i f o r m s e d i m e n t a t i o n rate c a n n o t be
m a d e . A single b e d of C i n c i n n a t i a n l i m e s t o n e m i g h t well represent the prod-
uct of a single major storm e v e n t , or it c o u l d represent a l o n g - t e r m a c c u m u l a -
tion of shell material. Likewise, a shale bed c o u l d he the p r o d u c t of a single
depositional event, or alternatively represent very slow a c c u m u l a t i o n . E a c h

Rocks, Fossils, and Time 63

 b e d d i n g c o n t a c t itself p o t e n t i a l l y represents a break in s e d i m e n t a t i o n or Hia-
tus o f i n d e t e r m i n a t e d u r a t i o n d a y s , m o n t h s , years, o r m o r e . M a n y C i n c i n -
n a t i a n l i m e s t o n e s h a v e irregular, pitted u p p e r surfaces, s o m e t i m e s b e a r i n g
e n c r u s t i n g o r g a n i s m s , w h i c h strongly suggest formation of a h a r d g r o u n d on
t h e sea floor o v e r a l o n g t i m e interval d u r i n g w h i c h very little s e d i m e n t ac-
c u m u l a t e d . I n g e n e r a l , g i v e n t h e a b u n d a n c e o f e v i d e n c e f o r storm-related
d e p o s i t i o n in the C i n c i n n a t i a n ( T o b i n 1982; Jennette and Pryor 1993). m o s t
l i m e s t o n e s p r o b a b l y represent m o r e t i m e t h a n t h e i n t e r b e d d e d shales, b u t
d e t e r m i n a t i o n of absolute d u r a t i o n s is very u n c e r t a i n .
W h e r e v e r s e d i m e n t a r y strata display a strongly c y c l i c pattern of repeated
sets of b e d s h a v i n g u n i f o r m t h i c k n e s s or variation in lithologies, geologists
h a v e s o u g h t a possible link to c y c l i c or e p i s o d i c c a u s e s reflecting s e a s o n a l ,
a n n u a l , or l o n g e r t i m e s c a l e s of periodicity. M o s t i n t r i g u i n g is the possibility
that variations in the E a r t h ' s orbital p a r a m e t e r s c o u l d exert i n f l u e n c e on
c l i m a t e c h a n g e s that in t u r n c a u s e c y c l i c or p e r i o d i c s e d i m e n t a t i o n pro-
cesses ( G r o t z i n g e r et al. 2007). If s e d i m e n t a t i o n c o u l d be s h o w n to respond
t o this k i n d o f a s t r o n o m i c a l m e t r o n o m e , interpolation b e t w e e n calibration
points o n the t i m e s c a l e c o u l d b e m a d e accurately, and the a m o u n t o f t i m e
represented by p a r t i c u l a r sets of strata c o u l d be d e t e r m i n e d . However, cycles
o f this t y p e , k n o w n a s M i l a n k o v i t c h c y c l e s , h a v e not a s yet b e e n d e m o n -
strated to exist w i t h i n the cyclic C i n c i n n a t i a n strata

64 A Sea without Fish

66 A Sea without Fish
 5
ALGAE: THE BASE OF
THE FOOD CHAIN

A l g a e are u n c o m m o n fossils i n t h e C i n c i n n a t i a n , b u t are p o t e n t i a l l y sig- Figure 5.1. A - C . Cincin-

nificant a s p r i m a r y p r o d u c e r s o f t h e O r d o v i c i a n e c o s y s t e m a n d a s i n d i c a - natian acritarchs, all Ede-

tors o f i m p o r t a n t e n v i r o n m e n t a l c o n d i t i o n s s u c h a s w a t e r d e p t h . A l g a e nian, Kope Formation,

Wayne County, Indi-
i n c l u d e s i n g l e - c e l l e d as well as m u l t i - c e l l e d p l a n t s that are c o n f i n e d to
ana. A. Veryhachium
a q u a t i c or moist habitats b e c a u s e t h e y lack internal c a n a l s for w a t e r storage
edenense Colbath, x
and transport. T h e y arc therefore t e r m e d n o n v a s c u l a r . B e c a u s e t h e y o b t a i n
569 (from Colbath [1979,
their e n e r g y t h r o u g h p h o t o s y n t h e s i s , a l g a e r e q u i r e a d e q u a t e e x p o s u r e t o plate 13, figure 1]).
sunlight. This essential r e q u i r e m e n t g e n e r a l l y restricts t h e m t o very shal- B. Ordovicium gracile
l o w water. A l g a e i n c l u d e t h e b l u e - g r e e n a l g a e (division C y a n o p h y t a ) , g r e e n Colbath, x 507 (from Col-
a l g a e (division C h l o r o p h y t a , s i n g l e - a n d m u l t i - c e l l e d ) , red a l g a e (division bath [1979, plate 8, fig-
R h o d o p h y t a ) , and the b r o w n a l g a e (division P h a e o p h y t a ) . ure 4]). C. Multipli-

Blue-green a l g a e are u b i q u i t o u s in m o d e r n m a r i n e e n v i r o n m e n t s , w h e r e cisphaeridium

micraulaxum Colbath, x
they form a " t u r f " or m a t of very fine filaments less t h a n 1 mm in l e n g t h .
1038 (from Colbath
These mats are an i m p o r t a n t food s o u r c e for m a n y g r a z i n g a n i m a l s s u c h as
[1979, plate 7, figure 10]).
gastropods. A l t h o u g h algal filaments arc not often fossilized, the p r e s e n c e of
A-C reprinted by permis-
algal mats is c o m m o n l y r e c o r d e d in the g e o l o g i c record as finely l a m i n a t e d sion of E. Schweizer-
m o u n d s or sheets c a l l e d s t r o m a t o l i t e s . A l g a l m a t s h a v e sticky, velcro-like bart'sche. Note: in all
surfaces that trap very fine s e d i m e n t . As e a c h thin layer of s e d i m e n t a c c u - figures, x indicates the
m u l a t e s , the algal filaments grow t h r o u g h it to form a new m a t , a n d the magnification factor.
D-F. Cincinnatian chitino-
process c a n b e repeated indefinitely. O v e r t i m e the layers ( l a m i n a e ) c a n b e
zoans, all Maysvillian.
c e m e n t e d , s o m e t i m e s w i t h t h e h e l p of the a l g a e , to form a stromatolite.
Stromatolites are a m o n g the oldest k n o w n e v i d e n c e of life, d a t i n g to o v e r 3.5 D. Cyathochitina sp. cf.
C. campanulaeformis,
billion years old, and arc e s p e c i a l l y c h a r a c t e r i s t i c o f P r e c a m b r i a n s e d i m e n -
OSU 32534, x 747 (from
tary rocks. At the end of the P r e c a m b r i a n , stromatolites d e c l i n e d drastically
Miller [1976, plate 5, fig-
in a b u n d a n c e and variety of g r o w t h forms, possibly as a c o n s e q u e n c e of the
ure 6]). E. Hercochi-
evolution o f large g r a z i n g a n i m a l s . After t h e b e g i n n i n g o f the P a l e o z o i c , tina turnbulli Jenkins,
stromatolites b e c a m e e v e n less c o m m o n a n d w e r e u s u a l l y restricted t o very OSU 32568, x 516 (from
shallow water e n v i r o n m e n t s , such as tidal flats, w h e r e a l g a l m a t s w e r e less Miller [1976, plate 13,
susceptible to g r a z i n g . Interestingly, stromatolites are v i r t u a l l y absent f r o m figure 2a]). F. Conochi-

C i n c i n n a t i a n strata, a l t h o u g h t h e y o c c u r i n t h e tidal f l a t facies o f t h e M i d d l e tina hirsuta Laufeld,

OSU 32542, x 287 (from
O r d o v i c i a n H i g h B r i d g e G r o u p n e a r L e x i n g t o n , K e n t u c k y ( C r e s s i n a n and
Miller [1976, plate 8, fig-
N o g e r 1976). O n e b l u e - g r e e n alga that is preserved as c a l c a r e o u s filaments,
ure 1]). G-l. Cincinna-
Girvanella, o c c u r s in t h e W h i t e w a t e r F o r m a t i o n in t h e u p p e r m o s t C i n c i n -
tian dasycladacean al-
natian in a z o n e c o n t a i n i n g biscuit-like l a m i n a t e d o n c o l i t e s also attributed gae. G. Lepidolites
to a l g a e ( B l a c k w e l l et al. 1984). dickhauti Ulrich, William
A m o n g the m a n y s p e c i e s o f g r e e n a l g a e are s o m e s p e c i e s c a p a b l e o f Heimbrock collection,

precipitating c a l c i u m c a r b o n a t e w i t h i n their p l a n t tissues. S a n d - s i z e d f l a k e s Edenian, Kope Forma-

tion, Kenton County,
and m i n u t e n e e d l e s p r o d u c e d b y t h e s e a l g a e are responsible for h u g e a c c u -
Kentucky, x 3.8. H. Cy-
m u l a t i o n s o f c a l c a r e o u s s e d i m e n t s i n m o d e r n shallow tropical seas. O n e

67
clocrinites darwinii g r o u p , t h e D a s y c l a d a c e a e , is represented in t h e C i n c i n n a t i a n by several spe-
(Miller), Stephen Felton cies (Cross e t a l . 1996). M o d e r n d a s y c l a d s o c c u r n o d e e p e r than a b o u t 3 0 m ,
collection, Maysvillian,
a n d f r e q u e n t l y less t h a n 5 m ( W r a y 1 9 7 7 ) . Cyclocrinites is the largest and
Mt. Auburn Formation,
most c o m m o n dasyclad, found in the Bellevue Limestone and throughout
Butler County, Ohio, x
t h e R i c h m o n d C r o u p ( F i g u r e s 5 . 1 H , I). This alga r e s e m b l e s a g o l f ball in
1.3. I. C. darwinii, sur-
face detail of H, polygo- size, s h a p e , a n d its d i m p l e d s u r f a c e . In w e l l - p r e s e r v e d s p e c i m e n s the d i m -

nal facet diameter ~0.4 pled s u r f a c e reveals a pattern of r h o m b o i d a l plates. T h e s e plates arc a c t u a l l y
mm. e x p a n d e d e n d s of b r a n c h e s that radiate from a central axis. Several species
w e r e o r i g i n a l l y d e s c r i b e d u n d e r the n a m e Pasceolus, but Nitecki (1970) re-
ferred m o s t of t h e s e to a s i n g l e s p e c i e s , Cyclocrinites darwini (Miller).
T h r e e o t h e r g e n e r a o f d a s y c l a d a l g a e o c c u r i n the C i n c i n n a t i a n (Cross
et al. 1996). Lepidolites dickhautii U l r i c h has a s a u s a g e - l i k e s h a p e , a b o u t 2
cm l o n g , b u t is u s u a l l y flattened ( F i g u r e 5 . 1 G ) . Its s u r f a c e has a scaly a p -
p e a r a n c e , a n d it o c c u r s in t h e K o p e F o r m a t i o n . Anomaloides reticulatus
U l r i c h is r e p o r t e d f r o m t h e l o w e r F a i r v i e w F o r m a t i o n a n d has a s i m i l a r
s c a l y s u r f a c e . H o w e v e r , it is c l u b - s h a p e d , a n d c a n r e a c h several c e n t i m e t e r s
in length. Ischadites circularis (Emmons) has b e e n reported from the
F a i r v i e w , C o r r y v i l l e , a n d M t . A u b u r n F o r m a t i o n s ( H a l v e 1948) but its taxo-
n o m i c status h a s n o t b e e n r e c e n t l y r e v i e w e d .
O n e red a l g a , Solenopora richmondensis ( M i l l e r ) , o c c u r s in t h e W h i t e -
w a t e r a n d E l k h o r n F o r m a t i o n s o f t h e u p p e r m o s t R i c h m o n d G r o u p ( B l a c k - w e l l e t al. 1982
cerium richmondense). T h e skeletal microstructure of this a l g a , with parallel
l a m i n a e a n d v e r t i c a l pillars, c o u l d easily b e m i s t a k e n for a s t r o m a t o p o r o i d
(see c h a p t e r 6).

T w o o t h e r a l g a e o f u n c e r t a i n affinities are reported from t h e C i n c i n -

n a t i a n : Cyclindrocoelia covingtonensis Ulrich from the u p p e r F a i r v i e w For-
m a t i o n , a n d Faheria anomala M i l l e r from t h e u p p e r Fairview, B e l l e v u e ,
C o r r y v i l l e , M t . A u b u r n , A r n h e i m , Liberty, W h i t e w a t e r , and Elkhorn F o r m a -
tions ( D a l v e 1948). A l t h o u g h a l g a e are n o t c o m m o n i n the C i n c i n n a t i a n ,
the fact that a l g a e h a v e b e e n reported t h r o u g h o u t the s e c t i o n , from facies
r e g a r d e d as d e e p e r water as well as s h a l l o w e r , suggests that the entire section
represents s h a l l o w d e p t h s w i t h i n t h e p h o t i c z o n e . Notably, a l g a e are m o s t
c o m m o n i n facies o c c u r r i n g a t the top o f the m a j o r s h a l l o w - u p w a r d cycles:
the Bellevue, Mt. A u b u r n , and Whitewater-Elkhorn Formations, where
s h o a l i n g d e p t h s h a v e b e e n p o s t u l a t e d o n t h e basis o f o t h e r s e d i m e n t a r y
evidence.
S i n g l e - c e l l e d a l g a e o c c u r a s m i c r o f o s s i l s i n t h e C i n c i n n a t i a n a n d pro-
v i d e t h e o n l y fossil e v i d e n c e for t h e p h y t o p l a n k t o n that m u s t h a v e b e e n t h e
basis for t h e m a r i n e food c h a i n . T h e s e microfossils fall into t w o g r o u p s : t h e
a c r i t a r c h s a n d t h e c h i t i n o z o a n s . B o t h g r o u p s are r e c o v e r e d from acid-re-
sistant r e s i d u e s o f rock s a m p l e s . B e c a u s e t h e s e p l a n k t o n i c microfossils were
w i d e l y d i s t r i b u t e d b y o c e a n c u r r e n t s , t h e y a r c very u s e f u l for biostrati-
graphic correlation.
A c r i t a r c h s ( m e a n i n g " u n c e r t a i n o r i g i n " ) i n c l u d e a w i d e variety of m i -
c r o s c o p i c fossil cysts w i t h o r g a n i c ( n o n - m i n e r a l i z e d ) walls; t h e y m a y repre-
sent several different g r o u p s of a l g a e ; c o n s e q u e n t l y acritarchs are not a formal
t a x o n o m i c g r o u p ( W i l l i a m s 1978; C r o s s et al. 1996). A c r i t a r c h s are g e n e r a l l y

68 A Sea without Fish

spherical in s h a p e , less than 100 m i c r o n s in d i a m e t e r ( m i c r o n = m i l l i o n t h of
a meter), and h a v e p r o j e c t i n g spines that often b r a n c h ( F i g u r e s 5 . 1 A - C ) .
A l t h o u g h acritarchs are s i m i l a r to the resting stages of t h e d i n o f l a g e l l a t e s
(division Pyrrophyta), o n e o f the major c o m p o n e n t s o f t h e m o d e r n m a r i n e
phytoplankton, they c a n n o t be d i a g n o s t i c a l l y related to any p a r t i c u l a r phy-
toplankton g r o u p . A c r i t a r c h s o c c u r t h r o u g h o u t t h e C i n c i n n a t i a n a n d their
diversity is c o n s i d e r a b l e : fifty-two s p e c i e s w e r e r e c o r d e d by C o l b a t h (1979)
in a c o r e from the K o p e F o r m a t i o n of I n d i a n a ; J a c o b s o n (1978) reported
forty-four s p e c i e s from the C i n c i n n a t i r e g i o n ; a n d L o e b l i c h a n d T a p p a n
(1978) d e s c r i b e d forty n e w s p e c i e s f r o m the C i n c i n n a t i a n o f O h i o , I n d i a n a ,
and K e n t u c k y . Jacobson (1979) f o u n d that fluctuations in a b u n d a n c e of dif-
ferent acritarch species c o r r e s p o n d e d to p a l e o e n v i r o n m e n t a l variations in
relative water d e p t h .
D e s p i t e the n a m e , c h i t i n o z o a n s are n e i t h e r c o m p o s e d o f t h e p r o t e i n
c h i t i n , nor d o they d e f i n i t e l y r e p r e s e n t a n a n i m a l g r o u p . T h e a c t u a l c o m -
position of the m i n u t e ( 5 0 - 2 0 0 0 m i c r o n s ) , b o t t l e - s h a p e d test is s i m i l a r to
c h i t i n but is t e r m e d a p s e u d o c h i t i n . C h i t i n o z o a n s are t h o u g h t m o s t likely
t o b e s o m e t y p e o f p h y t o p l a n k t o n , b u t t h e i r e x a c t b i o l o g i c a l affinities are
u n r e s o l v e d (Jansonius a n d J e n k i n s 1978; C r o s s e t al. 1996). L i k e a c r i t a r c h s ,
c h i t i n o z o a n s are f o u n d t h r o u g h o u t t h e C i n c i n n a t i a n ( M i l l e r 1 9 7 6 ; J a c o b -
son 1979) a n d p r o v i d e u s e f u l i n f o r m a t i o n for w o r l d w i d e b i o s t r a t i g r a p h i c
correlation (Figures 5 . 1 D - F ) .

Algae 69
70 A Sea without Fish
PORIFERANS AND CNIDARIANS:
SPONGES, CORALS, AND JELLYFISH

A l t h o u g h s p o n g e s are r e g a r d e d a s t h e 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 - Sponges

tive o f m u l t i c e l l e d a n i m a l s , t h e y p l a y a n e s s e n t i a l role a s " s a n i t a r y e n g i -
n e e r s " i n a q u a t i c e n v i r o n m e n t s , l i v i n g a s a c t i v e s u s p e n s i o n f e e d e r s o r filter
f e e d e r s (Plate 3A). B y r e m o v i n g m i n u t e o r g a n i c p a r t i c l e s f r o m t h e w a t e r ,
sponges prevent decay products from p o i s o n i n g the e n v i r o n m e n t . This i s
a l o n g - r u n n i n g role, as s p o n g e s first a p p e a r in the fossil record d u r i n g t h e
late P r e c a m b r i a n , o v e r 540 m i l l i o n years a g o .
T h e b o d y o f a s p o n g e lacks d i s t i n c t c e l l layers, b u t i s c o m p o s e d o f dif-
ferent s p e c i a l i z e d t y p e s o f c e l l s that p e r f o r m d i f f e r e n t life f u n c t i o n s . T h e
f u n d a m e n t a l s p o n g e c e l l i s t h e c o l l a r c e l l , e q u i p p e d w i t h a w a v i n g flagel-
l u m that draws w a t e r into a c o n e f o r m e d o f m i c r o v i l l i ( F i g u r e 6.1). T h e
simplest sponge is a hollow tube, o p e n at o n e e n d . C o l l a r cells line the
interior o f the t u b e a n d c r e a t e a f e e d i n g c u r r e n t t h a t passes t h r o u g h t h e
b o d y w a l l via o p e n i n g s c a l l e d ostia a n d t u b u l a r c e l l s c a l l e d p o r o c y t e s . T h e
c o l l a r c e l l s r e m o v e f o o d p a r t i c l e s that are d i g e s t e d b y a m e b o c y t e s . T h e
feeding current carries wastewater, depleted of nutrients, out of the sponge
cavity t h r o u g h o n e o r m o r e c h i m n e y - l i k e o p e n i n g s c a l l e d o s c u l a . B e c a u s e
Figure 6 . 1 . A simple
s p o n g e s are f i x e d t o t h e s u b s t r a t u m a n d d o n o t m o v e a b o u t , t h e y are often
sponge, showing a cross-
r e g a r d e d as inert or n o n l i v i n g . In fact t h e y are a c t i v e l y c i r c u l a t i n g w a t e r section of the body wall.
and p r o c e s s i n g it for n u t r i e n t s (Plate 3A). Inset shows a magnified
T h e body of a s p o n g e is m o s t l y c o m p o s e d of a fibrous p r o t e i n c a l l e d view of incurrent canals
s p o n g i n , w h i c h i s also s e c r e t e d b y s p e c i a l i z e d c e l l s . ( T h i s i s w h a t m a k e s u p (ostia), collar cells and

a natural b a t h s p o n g e . ) A f t e r d e a t h , s p o n g i n r e a d i l y d e c a y s , s o t h a t m a n y collar-cell chambers.

Drawing by John Agnew.
s p o n g e s h a v e little c h a n c e o f b e c o m i n g f o s s i l i z e d . M o s t s p o n g e s a l s o se-
crete m i n u t e , m i n e r a l i z e d s p i c u l e s that are e m b e d d e d w i t h i n t h e s p o n g i n
n e t w o r k . S p i c u l e s c a n be as s i m p l e in form as a s i n g l e n e e d l e , b u t c a n also
be very c o m p l e x , burr-like, a n d e v e n f u s e d to f o r m a basket-like l a t t i c e ;
their c o m p o s i t i o n i s e i t h e r c a l c i u m c a r b o n a t e o r s i l i c o n d i o x i d e . O n c e a
s p o n g e d e c a y s , t h e s p i c u l e s are r e l e a s e d into t h e s e d i m e n t w h e r e t h e y c a n
b e preserved a s m i c r o f o s s i l s . S p o n g e s that h a v e d e n s e o r f u s e d n e t w o r k s o f
spicules are m o r e likely to be p r e s e r v e d i n t a c t , a n d it is t h e s e t y p e s t h a t
m a k e u p m o s t o f the fossil record o f s p o n g e s .
In the late 1960s biologists d i v i n g on t h e coral reefs of J a m a i c a discov-
ered a n e w g r o u p of l i v i n g s p o n g e s that c o m p l e t e l y defied t h e c o n c e p t of a
typical s p o n g e (Plate 3B). M o r e o v e r , t h e s e n e w s p o n g e s p r o v i d e d a n i m p o r -
tant link to s o m e fossils that had l o n g b e e n misclassified. A m o n g t h e s e fossils
are s o m e found i n the C i n c i n n a t i a n . T h e n e w s p o n g e s are c a l l e d sclcro-
sponges or coralline s p o n g e s b e c a u s e they form a massive c a l c a r e o u s skeleton
like that of coral. T h e s p o n g e b o d y is restricted to a t h i n s u r f a c e layer in

71
Figure 6.2. Reconstruc-
tion of a living stromato-
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 lami-
nae. Compare to Figure
6.3D, below. Drawing by
John Agnew.

w h i c h typical s p o n g e cells carry out f i l t e r f e e d i n g ( f i g u r e 6.2). Wastewater

c a n a l s that c o n v e r g e on the o s c u l a leave starburst patterns of g r o o v e s in t h e
c a l c a r e o u s skeleton that m a t c h w i t h structures c a l l e d astrorhizae in t h e fossil
g r o u p k n o w n a s s t r o m a t o p o r o i d s ( f i g u r e s 6.2, 6.3A, D ) . These and other
similarities e n a b l e d the s t r o m a t o p o r o i d s to be r e c o g n i z e d c o r r e c t l y as a new
g r o u p o f s p o n g e s , after t h e y h a d b e e n classified with c n i d a r i a n s , b r y o z o a n s ,
a n d e v e n p r o t o z o a n s . S t r o m a t o p o r o i d s f i r s t a p p e a r i n strata o f O r d o v i c i a n
a g e , a n d w e r e t h o u g h t t o h a v e b e c o m e e x t i n c t i n the C r e t a c e o u s , until the
l i v i n g s c l e r o s p o n g e s w e r e f o u n d . D u r i n g the S i l u r i a n and D e v o n i a n periods,
Figure 6.3. Cincinnatian
s t r o m a t o p o r o i d s w e r e m a j o r reef b u i l d e r s a l o n g w i t h c o r a l s , but the m o d e r n
sponges and stromato-
s c l e r o s p o n g e s ( m o r e distantly related to t h e P a l e o z o i c forms) are restricted to
poroids. A. A cylindri-
cal stromatoporoid, Aul- d e e p reef e n v i r o n m e n t s a n d play a s m a l l e r role in reef b u i l d i n g .
acera undulata S p o n g e s are n o t c o m m o n f o s s i l s i n the C i n c i n n a t i a n , a l t h o u g h t h e y
(Billings), MUGM 29618, often m a y be overlooked b e c a u s e they can resemble bryozoans or corals,
Richmondian, horizon a n d a l s o h a v e a rather n o n d e s c r i p t a p p e a r a n c e ( f i g u r e 6 . 3 C ) . Five g e n e r a
and locality unknown, x o f s p o n g e s a n d t h r e e g e n e r a o f s t r o m a t o p o r o i d s are r e c o g n i z e d i n the C i n -
0.3. B. A large sponge,
c i n n a t i a n ( D a l v e 1 9 4 8 ; R i g b y 1996). Brachiospongia is t h e largest a n d m o s t
Brachiospongia tuber-
s t r i k i n g s p o n g e ( f i g u r e 6.3B). This s p o n g e has a hollow c e n t r a l c a v i t y from
culata James, holotype,
w h i c h radiate 6 - 1 2 straight o r c u r v e d , f i n g e r - l i k e p r o j e c t i o n s ; its d i a m e t e r
CMC IP 209, Richmon-
c a n r e a c h 2 8 c m (11 in). O t h e r C i n c i n n a t i a n s p o n g e s arc d e s c r i b e d i n
dian, Liberty Formation,
Clinton County, Ohio, x d e t a i l a n d illustrated b y R i g b y ( 1 9 9 6 ) .

0.3. C. A sponge, Pat- S t r o m a t o p o r o i d s are n o t c o m m o n i n t h e C i n c i n n a t i a n a n d m a y super-

tersonia tuberosa ficially r e s e m b l e b r y o z o a n s in h a v i n g a m o u n d - l i k e or e n c r u s t i n g form
(Beecher), MUGM, ( F i g u r e 6 . 3 D ) . T h e y differ f r o m b r y o z o a n s i n b e i n g d e n s e l y c o v e r e d w i t h
Maysvillian, Fleming t u b e r c l e s . C r o s s - s e c t i o n s o f C i n c i n n a t i a n s t r o m a t o p o r o i d s are c o m p o s e d
County, Kentucky, x
o f blister-like skeletal d e p o s i t s c a l l e d v e s i c l e s , w h e r e a s b r y o z o a n s h a v e a
0.3. D. Polished cross-
t u b u l a r s t r u c t u r e ( F i g u r e 6.2). Aulacera is a very d i s t i n c t i v e , large C i n c i n -
section of a stromato-
n a t i a n s t r o m a t o p o r o i d w i t h a c y l i n d r i c a l s h a p e ( f i g u r e 6.3A). In life, Aul-
poroid, Labechia huron-
acera grew u p r i g h t on the sea floor like a tree or a s t a l a g m i t e , but they are
ensis (Billings), MUGM
634A2t, Richmondian, p r e s e r v e d l y i n g h o r i z o n t a l l y like fallen logs. Aulacera f o u n d in O r d o v i c i a n

Montgomery County, strata slightly y o u n g e r t h a n t h e y o u n g e s t C i n c i n n a t i a n b e d s o n A n t i c o s t i

Ohio, x 0.3. Island in the Gulf of St. L a w r e n c e r e a c h e d g i g a n t i c s i z e s , up to 28 cm in

72 A Sea without Fish

d i a m e t e r a n d 1 - 2 m i n l e n g t h ( C a m e r o n a n d C o p p e r 1994). I n t h e C i n c i n -
nati region they arc restricted to the R i c h i n o n d i a n f o r m a t i o n s , the Elkhorn
f o r m a t i o n i n p a r t i c u l a r , a n d d i a m e t e r s u p t o 5 - 1 0 c m arc k n o w n .

T h e stony c o r a l s are the best k n o w n m e m b e r s o f the p h y l u m C n i d a r i a , Cnidarians

w h i c h also i n c l u d e s jellyfish, sea a n e m o n e s , a n d m a n y g r o u p s like I n -

Poriferans and Cnidarians 73

 d r o i d s , sea f a n s , sea w h i p s , a n d soft c o r a l s that are often m i s t a k e n l y t h o u g h t
mouth to be s e a w e e d s . D e s p i t e s u c h a b e w i l d e r i n g array of f o r m s , c n i d a r i a n s share
several features that i n d i c a t e their c o m m o n r e l a t i o n s h i p a s m e m b e r s o f o n e
o f t h e s i m p l e s t m u l t i - c e l l e d a n i m a l ( m e t a z o a n ) p h y l a . C n i d a r i a n s stand
a p a r t f r o m o t h e r a n i m a l s i n h a v i n g t w o b o d y f o r m s , the p o l y p a n d the
corallum
medusa. T h e p o l y p , as typified by a sea a n e m o n e or stony c o r a l (Plate 3C;
F i g u r e 6.4), is c y l i n d r i c a l in s h a p e a n d a t t a c h e d at t h e b a s e to a hard s u b -
stratum. T h e body w a l l c o n s i s t s o f o n l y t w o c e l l layers ( u n l i k e the t h r e e
layers f o u n d in all o t h e r m e t a z o a n s ) , s e p a r a t e d by a n o n - c e l l u l a r jelly layer
c a l l e d t h e m e s o g l e a . T h e r e is a s i n g l e o p e n i n g ( m o u t h ) into the b o d y cavity
t h r o u g h w h i c h f o o d is i n g e s t e d a n d w a s t e is e x p e l l e d . A r i n g of t e n t a c l e s
s u r r o u n d i n g t h e m o u t h serves for f o o d c a p t u r e a n d d e f e n s e . T h e m e d u s a
or jellyfish has t h e s a m e s t r u c t u r e as the p o l y p (with t h i c k e r m e s o g l e a ) but
Figure 6.4. Reconstruc- i s f r e e - l i v i n g , s w i m m i n g b y m u s c u l a r p u l s a t i o n s w i t h the m o u t h o r i e n t e d
tion of the Cincinnatian
d o w n w a r d s . B o t h p o l y p a n d m e d u s a f o r m s arc p r e s e n t a t different stages
solitary rugose coral,
d u r i n g t h e life c y c l e o f s o m e c n i d a r i a n s p e c i e s .
Grewingkia canaden-
T h e t h r e e m a j o r c l a s s e s o f c n i d a r i a n s differ i n their expression o f t h e
sis, showing the polyp
p o l y p and m e d u s a stages. H y d r o z o a n s (including hydroids, Portuguese Man-of-War, a
with extended tentacles.
Drawing by Kevina (true jellyfish) restrict t h e p o l y p to a larval s t a g e a n d live m o s t l y as m e d u -
Vulinec. sae. A n t h o z o a n s ( a n e m o n e s , c o r a l s , "soft c o r a l s " ) live e x c l u s i v e l y a s p o l y p s ,
a n d often a s c o l o n i e s o f m u l t i p l e p o l y p s that are g e n e t i c c l o n e s o f a s i n g l e
initial p o l y p . O n e o t h e r f e a t u r e c o m m o n t o all c n i d a r i a n s p r o v i d e s a c l u e
t o t h e i r m o d e o f life. M i c r o s c o p i c s t i n g i n g c e l l s (cnidoblasts) are c o n c e n -
trated in t h e t e n t a c l e s of all c n i d a r i a n s . U p o n c o n t a c t w i t h a foreign o b j e c t
t h e s t i n g i n g c e l l r e l e a s e s a h a r p o o n - l i k e h o l l o w t h r e a d that p i e r c e s soft
tissue a n d injects a t o x i n . S m a l l o r g a n i s m s are s t u n n e d o r k i l l e d b y t h e
s t i n g i n g c e l l s , t h e n g r a s p e d b y t h e t e n t a c l e s a n d stuffed into t h e m o u t h .
T h u s c n i d a r i a n s live a s p r e d a c e o u s c a r n i v o r e s , a l t h o u g h p o l y p s a n d w e a k l y
s w i m m i n g jellyfish m u s t rely o n w a t e r m o t i o n t o s u p p l y their prey. B e c a u s e
t h e prey o f m o s t c n i d a r i a n s i s m a i n l y m i n u t e z o o p l a n k t o n , t h e y are also
c o n s i d e r e d t o b e passive s u s p e n s i o n f e e d e r s .

I n t h e C i n c i n n a t i a n strata, t w o m a j o r g r o u p s o f c n i d a r i a n s arc repre-

s e n t e d : t h e a n t h o z o a n c o r a l s a n d the s c y p h o z o a n c o n u l a r i i d s . The corals
i n c l u d e b o t h solitary a n d c o l o n i a l s p e c i e s .

Solitary Corals C i n c i n n a t i a n solitary corals, c o m m o n l y c a l l e d horn corals o r c u p corals, a n d

o n e c o l o n i a l coral b e l o n g to the order R u g o s a . R u g o s a n s first a p p e a r e d in the
L a t e O r d o v i c i a n a n d b e c a m e extinct b y the e n d o f the P e r m i a n . The c o m m o n
n a m e " h o r n c o r a l " r e f e r s t o the c o n i c a l o r c y l i n d r i c a l s h a p e o f the calcitic
skeleton ( c o r a l l u m ) ; these corals are c o m m o n l y m i s t a k e n for a fossilized cow's
h o r n ( F i g u r e s 6.4, 6.5). T h e p o l y p o c c u p i e d a depression (calice) at the wide-
e n d o f the c o r a l l u m . A s the coral grew, the p o l y p deposited successive l a y e r s
of skeletal material b e n e a t h it, thereby i n c r e a s i n g the l e n g t h of the c o r a l l u m .
O l d e r parts of the c o r a l l u m did not c o n t a i n living tissue and thus were subject
to physical abrasion or e n c r u s t a t i o n a n d b o r i n g by other o r g a n i s m s .

74 A Sea without Fish

Figure 6.5. Cincinnatian rugose corals. A. Streptelasma divaricans (Nicholson), USNM 70211, several cor-
alla attached to bryozoan, Richmondian, Waynesville Formation, Clinton County, Ohio, x 2.0 (from Elias
[1982, plate 3, figure 5]). B. S. divaricans, USNM 135767, three coralla attached to margin of brachiopod
Rafinesquina, Richmondian, Whitewater Formation, Wayne County, Indiana, x 1.3 (from Elias [1982, plate
3, figure 9[). C. S. divaricans, USNM 40086, two coralla attached to brachiopod Lepidocyclus, Rich-
mondian, Whitewater Formation, Butler County, Ohio, x 2.2 (from Elias [1982, plate 3, figure 8[). D.
Grewingkia canadensis (Billings), CMC IP 50667, Weaver Collection, well-preserved corallum, Richmon-
dian, Adams County, Ohio, x 0.6, inset showing epithecal growth-lines, scale in mm. E. G. canadensis,
CMC IP 45413, typical abraded corallum with broken rim, showing circular borings of Trypanites, Rich-
mondian, Whitewater-Elkhorn Formations, Wayne County, Indiana, x 1.0 (from Elias [1982, plate 9, figure
10]). A-C reprinted by permission of the Paleontological Research Institution.

Poriferans and Cnidarians 75

76 A Sea without Fish
 A l t h o u g h the soft p o l y p o f r u g o s a n s i s n e v e r p r e s e r v e d , r u g o s a n s are Figure 6.6. Cincinnatian
k n o w n t o b e c o r a l s b e c a u s e t h e c a l i c e has m u l t i p l e r a d i a t i n g p a r t i t i o n s colonial corals. A-E.
Corallites as seen on ex-
c a l l e d septa that are f o u n d in l i v i n g c o r a l s . S e p t a are s e c r e t e d by soft tissue
ternal surface or in cross
partitions o f t h e internal b o d y c a v i t y c a l l e d m e s e n t e r i e s . I n l i v i n g a n t h o z o -
sections of corallum, at
a n s , m e s e n t e r i e s serve i m p o r t a n t f u n c t i o n s i n d i g e s t i o n a n d r e p r o d u c t i o n .
same scale x 3.7. A - E
The n u m b e r a n d a r r a n g e m e n t o f t h e septa are traits u s e d i n t h e classifica- from Elias (1998). A.
tion of c o r a l s . In r u g o s a n s t h e septa h a v e a r o u g h l y four-fold s y m m e t r y , Cyathophylloides, a co-
c o m p a r e d t o l i v i n g c o r a l s that h a v e six-fold s y m m e t r y . lonial rugosan, Richmon-
T w o s p e c i e s o f solitary r u g o s a n s o c c u r c o m m o n l y i n t h e C i n c i n n a t i a n , dian. B. Foerstephyl-
b o t h in t h e R i c h m o n d i a n strata ( F i g u r e 6.5; E l i a s 1982, 1998). Grewingkia lum, a tabulate,
Richmondian. C. Cala-
canadensis (Billings) i s t h e largest a n d m o s t c o m m o n r u g o s a n ( F i g u r e s
poecia, a tabulate, Rich-
6 . 5 D , E). C o r a l l a reach l e n g t h s o v e r 13 cm (5 in) b u t are g e n e r a l l y in the
mondian. D. Nycto-
r a n g e 1 0 - 6 0 m m ( 0 . 5 - 2 in); t h e d i a m e t e r r a n g e s f r o m 2 2 t o 4 0 m m ( 0 . 9 - 1 , 6
pora, a tabulate,
in). S p e c i m e n s are a l m o s t a l w a y s f o u n d l y i n g o n t h e i r sides a n d a p p e a r Richmondian. E. Tetra-
highly abraded, encrusted, and bored (Figure 6.5E). External c o n c e n t r i c dium, a tabulate, Rich-
g r o w t h lines are rarely p r e s e r v e d ( F i g u r e 6 . 5 D , inset). Like s o m e l i v i n g mondian. F. Coral bed,
solitary c o r a l s , Grewingkia p r o b a b l y lived upright, partly buried in soft Richmondian, Madison

sediment with the polyp exposed. S o m e encrustation and b o r i n g took County, Kentucky, length
of hammer 25 cm (from
p l a c e d u r i n g life but c o n t i n u e d after t h e c o r a l w a s e x h u m e d b y s t o r m activ-
Elias [1998, figure 5]).
ity a n d d e p o s i t e d o n its side. B r y o z o a n s are t h e m o s t c o m m o n e n c r u s t e r s
G. Protaraea richmon-
a n d a w o r m p r o b a b l y f o r m e d t h e b o r i n g s (trace fossil n a m e Trypanites, s e e
densis Foerste, MUGM
Elias 1986). Field studies d e m o n s t r a t e that Grewingkia s p e c i m e n s on s i n g l e 5435, a tabulate encrust-
b e d d i n g s u r f a c e s are o r i e n t e d i n preferred d i r e c t i o n s t h a t p r o b a b l y resulted ing brachiopod shell,
from a l i g n m e n t o f c o r a l s d u r i n g s t o r m s . Richmondian, Liberty For-
Streptelasma divaricans ( N i c h o l s o n ) is t h e o t h e r solitary r u g o s e c o r a l mation, Preble County,
Ohio, x 1.7. H. Octago-
f o u n d i n t h e R i c h m o n d i a n s e c t i o n ( F i g u r e s 6 . 5 A - C ; E l i a s 1982, 1998).
nal tool house built ca.
U n l i k e Grewingkia, Streptelasma is f o u n d in g r o w t h p o s i t i o n , on the u p p e r
1900 in John Paul Park,
s u r f a c e s o f l i m e s t o n e s . T h e c o r a l l a are u s u a l l y 6 - 1 3 m m ( 0 . 2 5 - 0 . 5 in) i n
Madison, Indiana, con-
l e n g t h , rarely e x c e e d i n g 2 5 m m ( 1 in), w i t h a d i a m e t e r o f 1 3 m m (0.5 in). structed entirely of colo-
C o r a l l a o c c u r i n d i v i d u a l l y and i n c l u s t e r s , often a t t a c h e d t o b r a c h i o p o d s , nial corals from the Rich-
b r y o z o a n s , or e v e n c o r a l l a of Grewingkia. In m a n y c a s e s t h e b r a c h i o p o d or mondian coral beds
host w a s l i v i n g a t t h e t i m e t h e c o r a l s a t t a c h e d . T h e o u t e r layer o f t h e c o r a l - exposed in the vicinity.
l u m (epitheca) s h o w s septal g r o o v e s a n d interseptal ridges i n c o n t r a s t t o t h e A - F reprinted by permis-
sion of the Mid-America
smooth, worn e p i t h e c a of Grewingkia.
Paleontology Society.

Most C i n c i n n a t i a n colonial corals b e l o n g to another extinct coral group Colonial Corals

c a l l e d the T a b u l a t a . Tabulates are c o m m o n l y c a l l e d h o n e y - c o m b c o r a l s
b e c a u s e o f their m u l t i p l e , p o l y g o n a l c o r a l l i t e s (skeletal t u b e s s e c r e t e d b y
individual polyps) ( F i g u r e s 6.6, 6 . 7 B ) . Tabulates o r i g i n a t e d i n t h e E a r l y
O r d o v i c i a n a n d b e c a m e e x t i n c t b y t h e e n d o f the P e r m i a n . C o r a l l a o f t a b u -
lates vary in s h a p e f r o m sheet-like to h e m i s p h e r i c a l to s p h e r i c a l , r e a c h i n g
d i a m e t e r s of a b o u t 4 m e t e r s (13 ft). I n d i v i d u a l t a b u l a t e p o l y p s b u i l t tall,
n a r r o w corallites. P o l y p s p e r i o d i c a l l y d e p o s i t e d a t r a n s v e r s e basal plate
(tabula) a s t h e y g r e w ; t h u s , b r o k e n c o r a l l a o r l o n g i t u d i n a l p o l i s h e d s e c t i o n s
h a v e a c h a r a c t e r i s t i c l a d d e r - l i k e a p p e a r a n c e ( F i g u r e 6.7B). In life, t a b u l a t e
p o l y p s w e r e truly c o l o n i a l b e c a u s e c o r a l l i t e w a l l s are s h a r e d a n d h a v e in-
t e r c o n n e c t i n g pores. S e p t a are not well d e v e l o p e d , l e a d i n g s o m e s p e c i a l i s t s

Poriferans and Cnidarians 77

Figure 6.7. Cincinnatian colonial corals. A. Cyathophylloides stellata (Hall), MUGM 5285, a colonial
rugosan, Richmondian, Liberty Formation, Nelson County, Kentucky, x 3. B. Foerstephyllum vacuum
(Foerste), MUGM 5301, a tabulate in vertical section, showing tabulae, Richmondian, Liberty Formation,
Nelson County, Kentucky, x 0.8.

78 A Sea without Fish

t o q u e s t i o n w h e t h e r t a b u l a t e s w e r e i n fact c o r a l s . A n e x t r a o r d i n a r y d i s c o v -
ery of soft tissue p o l y p s p r e s e r v e d in a S i l u r i a n t a b u l a t e ( C o p p e r 1985)
settled the d e b a t e for m o s t t a b u l a t e s , a l t h o u g h s o m e , like t h e C i n c i n n a t i a n
Tetradium, are very s i m i l a r to s o m e l i v i n g s p o n g e s that b u i l d a c a l c a r e o u s
skeleton w i t h a t a b u l a t e s t r u c t u r e .
C o l o n i a l corals o c c u r exclusively i n the R i c h m o n d i a n W a y n e s v i l l e , Lib-
ertv, W h i t e w a t e r , S a l u d a , and Elkhorn F o r m a t i o n s . W i t h i n these f o r m a t i o n s ,
there are as m a n y as tour distinct h o r i z o n s w h e r e c o l o n i a l corals are c o n c e n -
trated into "coral b e d s " up to a b o u t 4 m (12 ft) thick, t r a c e a b l e for great dis-
tances a l o n g the o u t c r o p belt o f the R i c h m o n d i a n a r o u n d the C i n c i n n a t i
A r c h ( F i g u r e 6.6F; B r o w n e 1964, 1965; Hatfield 1968; E l i a s 1998). F o u r
genera of massive c o l o n i a l tabulates (Foerstephyllum, Calapoecia, Nyctopora,
and Tetradium) and o n e c o l o n i a l r u g o s a n (Cyathophylloides) are f o u n d in
these b e d s ( F i g u r e s 6 . 6 A - F ) . A n o t h e r t a b u l a t e , Protaraea, o c c u r s in the
R i c h m o n d i a n but did not form m a s s i v e c o l o n i e s . Instead, Protaraea e x c l u -
sively e n c r u s t s t h e shells o f b r a c h i o p o d s a n d o t h e r objects ( F i g u r e 6 . 6 G ) . I n
John Paul Park, in M a d i s o n , I n d i a n a , there is a u n i q u e , o c t a g o n a l tool h o u s e
built entirely of c o l o n i a l corals g a t h e r e d from the c o r a l b e d e x p o s e d north
of the town ( F i g u r e 6.6H). C o l o n i a l corals are also i n c o r p o r a t e d into stone-
walls beside s o m e o f the e l e g a n t h o u s e s i n M a d i s o n .

A l t h o u g h the coral beds of the R i c h m o n d i a n have s o m e characteristics of A r e Reefs Present in

reefs, t h e y are not c o n s i d e r e d to be true reefs. W h y is t h i s s o , a n d are t h e r e t h e Cincinnatian?
o t h e r reef-like c o n c e n t r a t i o n s o f fossils e l s e w h e r e i n t h e C i n c i n n a t i a n ? T o
a n s w e r t h e s e c o n d q u e s t i o n f i r s t , t h e o n l y o t h e r reef-like s t r u c t u r e s re-
p o r t e d in t h e C i n c i n n a t i a n arc s m a l l m o u n d s , o n l y 0.3 m h i g h by 3 in
across, w h i c h w e r e c o m p o s e d of t r e p o s t o m e b r y o z o a n s (Cuffey 1998).
T h e s e m o u n d s o c c u r in the Maysvillian Grant Lake Limestone near
M a y s v i l l e , K e n t u c k y , b u t u n f o r t u n a t e l y the o u t c r o p h a s b e e n d e s t r o y e d .
T h e r e are t h r e e r e a s o n s w h y t r u e reefs m i g h t b e e x p e c t e d i n t h e C i n -
c i n n a t i a n . First, o r g a n i s m s w i t h r e e f b u i l d i n g p o t e n t i a l c e r t a i n l y e x i s t e d i n
a b u n d a n c e throughout the C i n c i n n a t i a n , i n c l u d i n g corals, sponges, and
b r y o z o a n s . S e c o n d , t h e s e s a m e o r g a n i s m s w e r e c o n s t r u c t i n g t r u e reefs b y
Early O r d o v i c i a n t i m e i n o t h e r r e g i o n s w o r l d w i d e (Plate 2 ; C o p p e r 1 9 9 7 ) .
B y M i d d l e O r d o v i c i a n t i m e , b r y o z o a n s a s s u m e d a m a j o r role i n r e e f b u i l d -
ing in regions as close to the C i n c i n n a t i A r c h as T e n n e s s e e (Alberstadt et
al. 1974). By C i n c i n n a t i a n t i m e , a d i v e r s e east of r e e f - b u i l d i n g o r g a n i s m s
h a d a s s e m b l e d that w o u l d d o m i n a t e r e e f b u i l d i n g w o r l d w i d e d u r i n g t h e
ensuing Silurian and D e v o n i a n ( C o p p e r 1997). T h i r d , the tropical to sub-
tropical p a l e o l a t i t u d e o f the C i n c i n n a t i r e g i o n d u r i n g the L a t e O r d o v i c i a n
was well w i t h i n t h e c l i m a t i c r a n g e w h e r e r e e f b u i l d i n g m i g h t b e e x p e c t e d
and i n d e e d w a s o c c u r r i n g e l s e w h e r e w o r l d w i d e (Plate 2 ; C o p p e r 2 0 0 1 ;
W e b b y 2002).
The coral b e d s o f t h e R i c h m o n d i a n , a s w e l l a s t h e M a y s v i l l i a n b r y o -
z o a n m o u n d s , h a v e not b e e n r e g a r d e d a s t r u e reefs b e c a u s e t h e y did n o t
g r o w into a n i n t e r c o n n e c t e d f r a m e w o r k that d e v e l o p e d s i g n i f i c a n t relief i n
relation to the s u r r o u n d i n g sea floor. R a t h e r , t h e y w e r e r e s t r i c t e d to l o w -

Poriferans and Cnidarians 79

Figure 6.8. Conulariid,
Conularia formosa
Miller and Dyer, Univer-
sity of Cincinnati collec-
tions, Maysvillian, Cor-
ryville Formation, Butler
County, Ohio, x 7.2.

relief c o l o n i e s l i v i n g very c l o s e t o sea level (Hatfield 1968). W i t h i n the c o r a l

b e d s , c o n c e n t r a t i o n s o f c o r a l s are n o m o r e t h a n a few m e t e r s w i d e ( F i g u r e
6.6F). T h e r e is no differentiation of the coral cluster as a reef "core" from
t h e b e d s l y i n g a d j a c e n t to it. A r e e f c o r e u s u a l l y i n d i c a t e s the c o r a l s c o n -
s t r u c t e d a m o u n d h a v i n g relief greater than that of a s i n g l e c o l o n y , less than
o n e m e t e r . C o r a l clusters p r o b a b l y existed as s m a l l p a t c h e s on a level sea
floor, s i m i l a r t o s m a l l l i v i n g p a t c h reefs. T h e s i z e o f s o m e t a b u l a t e c o l o n i c s

80 A Sea without Fish

is c o m p a r a b l e to that of m a n y l i v i n g c o r a l s in p a t c h reefs. H a t f i e l d (1968)
s h o w e d that the c o r a l z o n e w i t h i n t h e S a l u d a F o r m a t i o n a c t e d as a low,
coral barrier s u r r o u n d i n g a c e n t r a l l a g o o n w h e r e f i n e - g r a i n e d c a r b o n a t e
s e d i m e n t s a c c u m u l a t e d . I n this w a y t h e c o r a l z o n e a c t e d a s d o p r e s e n t - d a y
reefs t o i n f l u e n c e w a t e r m o v e m e n t a n d s e d i m e n t d e p o s i t i o n . A p p l i c a t i o n
of the t e r m s p a t c h reefs a n d b i o s t r o m e s to t h e R i c h m o n d i a n c o r a l b e d s is
therefore q u i t e r e a s o n a b l e .
The inability o f C i n c i n n a t i a n c o r a l s a n d o t h e r p o t e n t i a l r e e f b u i l d e r s
to c o n s t r u c t m a j o r reefs has several p o s s i b l e e x p l a n a t i o n s . First, c o r a l s w e r e
present i n t h e region o n l y d u r i n g R i c h m o n d i a n t i m e ( W e b b y 2002). T h e
a b s e n c e o f corals d u r i n g E d e n i a n a n d M a y s v i l l i a n t i m e i s p u z z l i n g , b e -
c a u s e o f t h e similarity o f t h e rest o f t h e f a u n a t h r o u g h o u t t h e C i n c i n n a t i a n .
E n v i r o n m e n t a l c o n d i t i o n s i n the C i n c i n n a t i A r c h r e g i o n m a y h a v e b e e n
u n s u i t a b l e for solitary a n d c o l o n i a l c o r a l s d u r i n g t h e E d e n i a n a n d M a v s v i l -
lian, b u t it is d i f f i c u l t to identify t h e factors r e s p o n s i b l e . A b u n d a n c e of
f i n e - g r a i n e d s e d i m e n t s a n d f r e q u e n t d i s t u r b a n c e o f t h e sea floor b y s t o r m s
are t w o factors that m i g h t h a v e restricted t h e p r e s e n c e o f c o r a l s . H o w e v e r ,
b o t h factors are p e r v a s i v e t h r o u g h o u t t h e C i n c i n n a t i a n , a n d it is n o t cer-
tain that either d e c r e a s e d s i g n i f i c a n t l y d u r i n g t h e R i c h m o n d i a n . A c c o r d -
i n g t o E l i a s (1982) solitary c o r a l s w e r e i n t r o d u c e d d u r i n g a n e a r l y R i c h -
m o n d i a n invasion from s o u r c e s t o t h e west. S o l i t a r y c o r a l s are p r e s e n t i n
the E d e n i a n - M a y s v i l l i a n strata o f the M a q u o k e t a G r o u p t o t h e w e s t .
T h e introduction of corals to the C i n c i n n a t i region d u r i n g R i c h m o n -
dian t i m e possible w a s related t o p r o g r e s s i v e s h a l l o w i n g o f t h e r e g i o n dur-
i n g the C i n c i n n a t i a n that c u l m i n a t e d i n t h e R i c h m o n d i a n ( A n s t e y a n d
F o w l e r 1969; H a y 1998). C o r a l b e d s d e v e l o p e d o n a shallow p l a t f o r m that
was f l a n k e d b y d e e p e r water toward t h e w e s t a n d n o r t h ( E l i a s 1982). Pres-
ent-day c o r a l reefs d e v e l o p a l o n g s i m i l a r p l a t f o r m m a r g i n s w h e r e a b r e a k
i n s l o p e separates shallow from d e e p e r water. Prior t o t h e R i c h m o n d i a n ,
the C i n c i n n a t i a n p l a t f o r m m a y h a v e b e e n d e e p e r a n d w i t h o u t a b r e a k i n
slope toward t h e west that h a v e favored c o r a l d e v e l o p m e n t .

T h e c o n u l a r i i d s arc a m i n o r g r o u p y e t are a m o n g t h e m o s t p r o b l e m a t i c Conulariids

fossils t o b e f o u n d i n t h e C i n c i n n a t i a n . T h e y are u s u a l l y f o u n d i n a c o m -
pressed c o n d i t i o n i n s h a l e s a n d siltstones o f M a v s v i l l i a n a n d R i c h m o n d i a n
formations. As compressed s p e c i m e n s , conulariids appear to have a high
t r i a n g u l a r s h a p e , w i t h f i n e l y striated m a r k i n g s i n a c h e v r o n - l i k e p a t t e r n o n
a very thin i n t e g u m e n t , b r o w n o r b l a c k i n c o l o r ( F i g u r e 6.8). T h e i n t e g u -
m e n t i s c a l c i u m - p h o s p h a t i c i n c o m p o s i t i o n . U n c o m p r e s s e d s p e c i m e n s are
found e l s e w h e r e that s h o w t h e o r i g i n a l s h a p e t o b e p y r a m i d a l a n d four-
sided, and s o m e h a v e t r i a n g u l a r f l a p s e x t e n d i n g f r o m t h e sides a t t h e w i d e
end, suggesting a means of closure. Cross-sections of uncompressed speci-
m e n s reveal a b i f u r c a t i n g s e p t u m o r i g i n a t i n g f r o m e a c h o f t h e f o u r sides.
E v i d e n c e for a t t a c h m e n t at t h e a p i c a l end is o c c a s i o n a l l y f o u n d , b u t is of-
ten l a c k i n g . Possibly c o n u l a r i i d s lived p a r t of their lives a t t a c h e d a n d later
b e c a m e free-living.

Poriferans and Cnidarians 81

 T h e z o o l o g i c a l affinities o f c o n u l a r i i d s h a v e b e e n d e b a t e d for a l o n g
t i m e . C h i e f l y o n t h e basis o f their four-part s t r u c t u r e , c o n u l a r i i d s h a v e
b e e n classified w i t h t h e 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 a v e a t e t r a m e r a l
(four-fold) b o d y p l a n . T h e r e are l i v i n g s c y p h o z o a n s w i t h a c h i t i n o u s t h e c a
a n d s o m e t h a t live a t t a c h e d b y m e a n s o f a stalk. S o m e h a v e a r g u e d that
conulariids should be r e c o g n i z e d as a distinct p h y l u m ( B a b c o c k 1996b;
Babcock and Feldmann 1986), b u t r e c e n t w o r k b y V a n Iten a n d others
( 1 9 9 6 ) , a n d H u g h e s a n d o t h e r s (2000), c o n f i r m s that the similarities b e -
t w e e n c o n u l a r i i d s a n d s c y p h o z o a n s are i n d i c a t i v e o f a c l o s e e v o l u t i o n a r y
r e l a t i o n s h i p . C o n u l a r i i d s arc f o u n d i n m a r i n e strata o f C a m b r i a n t h r o u g h
Triassic a g e . A s i n g l e s p e c i e s , Conularia formosa M i l l e r a n d D y e r , is re-
corded from the C i n c i n n a t i a n of the C i n c i n n a t i A r c h region.

82 A Sea without Fish

84 A Sea without Fish
 7
BRYOZOANS: "TWIGS"
AND "BONES"

The rocks in the C i n c i n n a t i region are l o a d e d with fossils. Visitors to the area
c o m m o n l y are struck by all the " t i l i n g s " in the rock that look like s m a l l t w i g s ,
or, with a stretch of the i m a g i n a t i o n , small p i e c e s of b o n e s ( F i g u r e 7.1A). T h e y
are the most c o m m o n fossils in the b e d r o c k of the area. I n d e e d , if y o u were
to pick up a fossil in the C i n c i n n a t i region at r a n d o m , c h a n c e s are that it
would be one of these objects. But they are neither twigs nor b o n e s . T h e y are,
in fact, the r e m a i n s of a g r o u p of o r g a n i s m s c a l l e d b r y o z o a n s (Plates 3 D , E ) .
Figure 7.1. A. Frag-
I f y o u l o o k a t a n u n b r o k e n s u r f a c e o f y o u r b r y o z o a n fossil w i t h y o u r
ments of bryozoan colo-
trust} h a n d - l e n s , y o u see that it is r e p l e t e w i t h tiny h o l e s ( f i g u r e 7.1 B). If nies are the most abun-
y o u shift y o u r field of v i e w to a b r o k e n s u r f a c e , t h e tiny h o l e s are r e v e a l e d dant fossils in the
to be minute tubes. E a c h o n e of those m i n u t e tubes was o n c e h o m e to an type-Cincinnatian. Par-
e q u a l l y m i n u t e a n i m a l . T h u s , t h e fossil i n y o u r h a n d w a s c o n s t r u c t e d b y a vohallopora ramosa
c o l o n y of tiny c r e a t u r e s . A b r y o z o a n c o l o n y is r e m i n i s c e n t of a p i e c e of (d'Orbigny), CMC IP
27957, Bellevue Lime-
coral found on a present-day b e a c h in that c o r a l reefs a l s o arc m a d e by
stone, Cincinnati, Ohio.
myriads o f i n d i v i d u a l a n i m a l s . D e s p i t e t h e s u p e r f i c i a l r e s e m b l a n c e o f b r y o -
Scale in mm. B. Surface
z o a n c o l o n i e s a n d coral c o l o n i e s t o o n e a n o t h e r , t h e a n i m a l s i n v o l v e d arc
of bryozoan colony
very different, i n d e e d . C o r a l s are m e m b e r s o f p h y l u m C n i d a r i a , c o m m o n l y
showing minute open-
called coelenterates. E a c h coral a n i m a l is basically s a c - s h a p e d with a single ings (zooecia) on the left
aperture serving as both m o u t h and anus. and a cross-section
A b r y o z o a n a n i m a l i s m o r e c o m p l e x l y o r g a n i z e d ( F i g u r e 7.2). T h e r e through a broken surface
is an a l i m e n t a r y c a n a l , w i t h a d i s t i n c t m o u t h on o n e e n d a n d a d i s t i n c t on the right. Each of the
a n u s on the other. S u r r o u n d i n g t h e m o u t h is a r i n g of t e n t a c l e s c a l l e d a openings leads to a tube

l o p h o p h o r e . T h e l o p h o p h o r e serves as a f o o d - g a t h e r i n g s t r u c t u r e a n d for that was home to a tiny,

individual animal. Trepos-
gas e x c h a n g e b e t w e e n the a n i m a l a n d t h e s u r r o u n d i n g w a t e r (in o t h e r
tome bryozoan, Mon-
w o r d s , it is also a respiratory s t r u c t u r e ) . T h e a n u s is l o c a t e d o u t s i d e of t h e
ticulipora mammulata
lophophore. This i s w h a t gives t h e t a x o n o f t h e b r y o z o a n s its t e c h n i c a l
d'Orbigny, CMC IP 51107,
n a m e p h y l u m Ectoprocta. The " p r o c t a " part o f t h e w o r d m e a n s " o p e n - Bellevue Limestone. Cin-
i n g , " a n d t h e " e c t o " m e a n s " o u t s i d e of." cinnati, Ohio. Diameter
Frankly, most p e o p l e d o not call t h e s e a n i m a l s e c t o p r o c t s . I n f o r m e r of individual openings
years, w h e n the c r e a t u r e s w e r e less well u n d e r s t o o d t h a n t h e y are t o d a y , (zooecia) about 0.2 mm.
p e o p l e spoke o f p h y l u m B r y o z o a . E v e n t u a l l y i t w a s r e c o g n i z e d that so-
c a l l e d p h y l u m B r y o z o a l u m p e d t o g e t h e r a n i m a l s that are not-at-all c l o s e l y
related t o o n e a n o t h e r . H e n c e , t h e n a m e " B r y o z o a " s h o u l d b e a b a n d o n e d ,
in the t e c h n i c a l s e n s e . H o w e v e r , like t h e u s e of t h e t e r m " g l a s s e s " to refer
t o items m a d e o f plastic, the t e r m " b r y o z o a n s " persists i n c o m m o n par-
l a n c e . B r y o z o a n s s o m e t i m e s are said t o c o m p r i s e p h y l u m P o l y z o a , e s p e -
cially i n G r e a t Britain. I n s o m e r e s p e c t s , " P o l y z o a " i s a n a p p r o p r i a t e n a m e
f o r the g r o u p . " P o l y " m e a n s " m a n y , " a n d " z o a , " " a n i m a l s . " T h e w o r d " B r y o -
z o a " literally m e a n s " m o s s a n i m a l s , " p r e s u m a b l y b e c a u s e a l i v i n g b r y o z o a n

85
Figure 7.2. Living b r y o - c o l o n y , w i t h its s u r f a c e o f m a n y m i n u t e a n i m a l s , m i g h t b e t h o u g h t t o re-
zoan, showing one zooid s e m b l e a r o c k c o a t e d w i t h m a n y o f t h e tiny plants w e c a l l m o s s . Potential
with tentacles extended c o n f u s i o n c a n e n s u e , h o w e v e r , i f o n e forgets the fact that b r y o z o a n s are
in feeding position (left)
d e c i d e d l y a n i m a l s , w h e r e a s m o s s e s are just a s d e c i d e d l y plants. R e g a r d l e s s
and the other partly re-
o f t h e t e c h n i c a l t e r m s a n d t h e r e a s o n i n g b e h i n d t h e m , m o s t p e o p l e refer
tracted (right). Drawing
to fossil e c t o p r o c t s c o l l o q u i a l l y as b r y o z o a n s , a n d t h e y h a v e d o n e so for
by Kevina Vulinec.
g e n e r a t i o n s . W e will f o l l o w that h o a r y tradition h e r e .
B r y o z o a n s are a n i m a l s . A l l a n i m a l s d e r i v e the e n e r g y t h e y n e e d t o
g r o w , r e p r o d u c e , a n d , i n d e e d , t o l i v e , b y c o n s u m i n g o t h e r o r g a n i s m s , or,
at least, o r g a n i c m a t t e r p r o d u c e d by l i v i n g o r g a n i s m s . A parasite, for ex-
a m p l e , a tape-worm living within the alimentary canal of another animal,
m a y a b s o r b o r g a n i c - r i c h fluids f r o m w i t h i n its host. A m o s q u i t o eats its
v i c t i m o n e tiny d r o p of b l o o d at a t i m e . But b r y o z o a n s arc n e i t h e r parasites
nor m o s q u i t o - l i k e . S o w h a t d o b r y o z o a n s c a t , a n d h o w d o t h e y eat it?

86 A Sea without Fish

 B r y o z o a n s arc a q u a t i c . D e p e n d i n g o n t h e k i n d , s o m e b r y o z o a n s live i n Figure 7.3. Cincinnatian
fresh water, but most live in salt water. In either c a s e , the b r y o z o a n subsists bryozoans. A. Colonies

o n m i n u t e o r g a n i s m s ( p r o t o z o a n s a n d s o on) a n d tiny bits o f o r g a n i c m a t t e r of the bryozoan genus

Spatiopora characteristi-
s u s p e n d e d in the water. T h e b r y o z o a n d o e s n o t just wait passively for s u c h
cally form a thin coating
food to fall into its m o u t h ; it literally filters the food f r o m the water. B r y o z o -
on shells of orthoconic
ans are active filter feeders. The i n d i v i d u a l a n i m a l spreads the tentacles of
cephalopods, MUGM
its l o p h o p h o r e into a f u n n e l - , b o w l - , or vase-like c o n f i g u r a t i o n (Plate 3 E ; uncatalogued, Cincinna-
F i g u r e 7.2), a n d cilia that line t h e tentacles m o v e in s u c h a w a y that f o o d tian, scale in mm. Note
particles are carried d o w n t o t h e m o u t h . N o t o n l y d o e s t h e i n d i v i d u a l p o l y p - that raised lumps on col-
ide generate f e e d i n g currents for itself, but the c o l o n i e s of at least s o m e kinds ony surface (monticules)
are elongated and
of b r y o z o a n s generate currents that e n h a n c e f e e d i n g in the c o l o n y as a w h o l e .
aligned with the axis of
In at least s o m e kinds of b r y o z o a n s there are p a r t i c u l a r areas on the c o l o n y
host nautiloid shell. B,
that have polypides that direct c u r r e n t s a w a y from the colony. T h e s e excur-
C. Ctenostome bryo-
rent chimney's (Plate 3 F ) carry water that a l r e a d y has b e e n filtered by t h e zoan, Ropalonaria
lophophores a w a y from t h e z o a r i u m , a n d t h e r e b y d r a w u n f i l t e r e d , nutrient- venosa Ulrich. B. Col-
laden water across p o l y p i d e s e l s e w h e r e in the c o l o n y ( M c K i n n e y a n d Jackson ony on shell of brachio-
1989). M o r e o v e r , the very topology of the c o l o n y may facilitate t h e passage pod Rafinesquina, CMC
of nutrient-filled water t h r o u g h t h e colony and across its p o l y p i d e - l i n e d , a n d , IP 40061, Waynesville
Formation, Butler Co.,
h e n c e , l o p h o p h o r e - l i n e d , a n d , h e n c e , f o o d - g a t h e r i n g surfaces.
Ohio, x 9. Arrow indi-
Fossil b r y o z o a n c o l o n i c s c o m e i n a w i d e variety o f sizes a n d s h a p e s
cates sac zooid. C.
( F i g u r e s 7.3, 7.4). some are l u m p s the size a n d s h a p e of a g u m - d r o p . S o m e
Scanning electron micro-
are stony masses larger t h a n y o u r fist. S o m e g r o w up from a shell or shell graph of polyester cast of
f r a g m e n t on the sea floor as a s m a l l b l a d e or d e l i c a t e l y b r a n c h e d s t r u c t u r e cavities excavated by
r e s e m b l i n g a m i n i a t u r e version of a present-day "stag-horn coral." S u c h c o l o - zooids into host shell,
nies c a n e x c e e d the size o f a b a s k e t b a l l , a l t h o u g h w h a t w e m o s t l y see are BMNH D.52264, x 22. B

twig shaped f r a g m e n t s . M a n y b r y o z o a n s g r o w as thin crusts on b r a c h i o p o d and C from Pohowsky

(1978, plate 1, figures 5,
shells or m o l l u s c shells (see F i g u r e 9 . 2 D ) . A s m a l l n u m b e r e v e n grow w i t h i n
7) and reprinted by per-
the shell matter of the b r a c h i o p o d or m o l l u s c , f o r m i n g tiny d e n d r i t i c or
mission of the Paleonto-
a n a s t o m o s i n g c a n a l s ( F i g u r e s 7.3B, C ) . W h a t e v e r its size a n d s h a p e , t h e hard
logical Research Institu-
parts of a bryozoan colony c o m p r i s e w h a t is c a l l e d a z o a r i u m . tion. D. Cystoporid
So zoaria e x h i b i t a t r e m e n d o u s variety in overall s h a p e . T h e y a l s o offer bryozoan, Constellaria
a t r e m e n d o u s variety i n details. The surface of the c o l o n y m a y be s m o o t h . florida Ulrich, CMC IP
Many, however, bear regularly spaced b u m p s , termed monticules, or regu- 51108, Fairview Forma-
tion, Kenton Co., Ken-
larly s p a c e d d e p r e s s i o n s , c a l l e d m a c u l a e . M o n t i c u l e s m a y b e e q u i d i m e n -
tucky. Inset, enlargement
sional i n m a p - v i e w , o r t h e y m a y b e e l o n g a t e d , e v e n r i d g e - l i k e . T h e y m a y
showing characteristic
be d i s p o s e d in a s e e m i n g l y r a n d o m array, or they m a y h a v e a d i s t i n c t pat-
star-shaped monticules,
tern. For e x a m p l e , s p e c i m e n s o f t h e aptly n a m e d g e n u s Constellaria are scale in mm. E. Cryp-
veritable c o n s t e l l a t i o n s o f star-shaped b u n c h e s o f m o n t i c u l e s ( F i g u r e 7.3D). tostome bryozoan, Esch-
S o m e s t u d e n t s o f b r y o z o a n s h a v e c o n c l u d e d that m o n t i c u l e s a n d m a c u l a e aropora sp., CMC IP
are simply m a n i f e s t a t i o n s o f t h e s a m e p h e n o m e n o n , s o d o n o t d i f f e r e n t i a t e 51110, Fairview Forma-
the t w o f r o m o n e a n o t h e r ; t h e y t h e r e f o r e c a l l all o f t h e m , w h e t h e r b u m p s tion, Boone Co., Ken-
tucky. Escharopora has
o r depressions, m a c u l a e ( M c K i n n e y a n d Jackson 1989). R e g a r d l e s s o f w h a t
zooecia on both sides of
o n e calls t h e s e e l e v a t i o n s a n d d e p r e s s i o n s , t h e y a p p a r e n t l y w e r e t h e p l a c e s
the thin, bladelike zo-
o n the c o l o n y w h e r e t h e e x c u r r e n t c h i m n e y s w e r e g e n e r a t e d .
arium. Inset, enlargement
All this variety w o u l d s e e m to offer a fertile field for t h e t a x o n o m i s t a showing zooecia, scale in
new n a m e for e a c h m o r p h o l o g i c v a r i a n t . I n d e e d , a w h o l e slew o f b r y o z o a n mm. F. Basal surface of
genera and species have b e e n r e c o g n i z e d by a whole slew of paleontolo- trepostome bryozoan
gists. B u t there is a p r o b l e m . S o m e t i m e s , in a s i n g l e c o l o n y , t h e c h a r a c t e r -

Bryozoans 87
88 A Sea without Fish
istics o f o n e " s p e c i e s , " o r e v e n " g e n u s , " g i v e w a y t o t h o s e o f a n o t h e r a l l i n that has grown on the
the s p a c e of a c e n t i m e t e r or two. P r e s u m a b l y , e v e r y o n e in a s i n g l e c o l o n y pedicle valve of the bra-

is of the s a m e s p e c i e s . T h u s , t h e c o n c l u s i o n is i n e v i t a b l e : o v e r a l l c o l o n y chiopod Rafinesquina

and overgrown it, Rich-
s h a p e and details on its s u r f a c e m a y not a l w a y s be reliable i n d i c a t o r s as to
ard Arnold Davis collec-
w h o is related to w h o m . I n d e e d , t h e r e is a m p l e e v i d e n c e p r o v i d e d by pres-
tion, Bellevue Limestone,
ent-day b r y o z o a n s that e n v i r o n m e n t c a n play a s i g n i f i c a n t role in c o l o n y
Cincinnati, Ohio, scale in
s h a p e , at least in s o m e taxa. mm. G. Trepostome
W e l l , if t h e s h a p e of the z o a r i u m is n o t an i n c o n t r o v e r t i b l e taxonomic bryozoan colony that has
indicator, w h a t , it a n y t h i n g , is? Is t h e r e an " i n n e r t r u t h " in b r y o z o a n tax- grown on shell of pelecy-
o n o m y ? It t u r n s o u t that t h e r e is just that, n a m e l y , t h e i n t e r n a l s t r u c t u r e of pod Ambonychia and
overgrown it, Richard
the colony. E a c h z o a r i u m consists o f t h e hard parts o f all o f t h e a n i m a l s
Arnold Davis collection,
that c o m p r i s e the c o l o n y . A n i n d i v i d u a l b r y o z o a n a n i m a l i s c a l l e d a z o o i d ,
Cincinnatian, horizon and
and the h a u l parts o f that i n d i v i d u a l a n i m a l c o n s t i t u t e a z o o e c i u m ( F i g u r e
locality unknown, scale
7.2). If o n e e x a m i n e s t h e h o l e s in a z o a r i u m w i t h a h a n d - l e n s or l o w - p o w e r in mm. H. Ring-shaped
m i c r o s c o p e , o n e c o m m o n l y sees that t h e h o l e s are not i d e n t i c a l ( F i g u r e bryozoan zoarium ("We-
7.1B). T h e r e m a y b e size classes o f larger h o l e s a n d s m a l l e r h o l e s ; there m a y ichold donut") presum-
e v e n be spine-like p r o j e c t i o n s in a d d i t i o n to h o l e s . It w o u l d a p p e a r t h a t , in ably encrusting nautiloid,
a t least s o m e c o l o n i e s , n o t all t h e z o o i d s w e r e i d e n t i c a l . I n s o m e p r e s e n t - CMC IP 51109, Richmon-
dian, Hamilton Co., Ohio,
day c o l o n i a l a n i m a l s , t h e r e i s p o l y m o r p h i s m , w i t h d i f f e r e n t - s h a p e d o r
x 0.7. I. Cross-section
different-sized i n d i v i d u a l s p e r f o r m i n g different tasks, for t h e g o o d of the
of ring-shaped bryozoan
colony, the s p e c i e s , o r b o t h . T h a t s e e m s t o h a v e b e e n t h e c a s e a m o n g a t
zoarium ("Weichold do-
least s o m e n o w - e x t i n c t b r y o z o a n s . A s m e n t i o n e d p r e v i o u s l y , t h e s u r f a c e o f nut") encrusting nau-
a zoarium m a y be m a r k e d by r e g u l a r l y s p a c e d b u m p s or d e p r e s s i o n s ( Fig- tiloid, Richard Arnold
ures 7.3A, D). In g e n e r a l , t h e s e m o n t i c u l e s and m a c u l a e , respectively, c o n - Davis collection, Upper
sist o f z o o e c i a o f sizes a n d n a t u r e s different f r o m t h o s e b e t w e e n t h e m , a n d , Maysvillian-Lower Rich-

h e n c e , s e e m t o represent areas o f the colon i n w h i c h the z o o i d s p e r f o r m e d mondian, Butler Co.,

Ohio, scale in mm. Dark,
p a r t i c u l a r f u n c t i o n s for t h e colony as a w h o l e .
elliptical zone on right
As we noted previously, s o m e bryozoan c o l o n i e s e x c e e d the size of bas-
and left sides of ring are
ketballs. How do we know that? The a n s w e r is p a i n s t a k i n g l y s i m p l e , with the calcitic replacement of
emphasis on the word " p a i n s t a k i n g . " In g e n e r a l , b r y o z o a n s are f o u n d o n l y as nautiloid shell.
small fragments scattered t h r o u g h o u t the rock. V e r y o c c a s i o n a l l y , h o w e v e r ,
o n e finds that all the f r a g m e n t s of a c o l o n y are l y i n g together, as part of a
single layer of rock. T h a t ' s the g o o d news: the c o m p l e t e colony is there! The
bad n e w s , however: a l t h o u g h the colony m a y be c o m p l e t e , it is not w h o l e .
T h a t ' s w h e r e t h e " p a i n s t a k i n g " c o m e s into the p i c t u r e . O n e m u s t oh-so-
carefully c o l l e c t e a c h f r a g m e n t , p a y i n g m e t i c u l o u s attention t o just w h e r e
the fragment was in the rock and adjacent to what o t h e r f r a g m e n t s . T h e n
o n e m u s t play t h r e e - d i m e n s i o n a l b r y o z o a n jigsaw p u z z l e a n d g l u e t h e tens,
or h u n d r e d s , or t h o u s a n d s of p i e c e s e a c h in its p r o p e r p l a c e .
A c c o r d i n g to the old nursery rhyme, "All the King's horses and all the
King's m e n c o u l d n ' t put H u m p t y together a g a i n " ( O p i e and O p i e 1955). A s
it h a p p e n s , s o m e of the local fossil c o l l e c t o r s a n d o t h e r p a l e o n t o l o g i s t s h a v e
b e e n m o r e clever, or at least m o r e persistent. For e x a m p l e , a m a t e u r p a l e o n -
tologists Jerry Rush a n d , m o r e recently, R o n F i n e h a v e spent c o u n t l e s s h o u r s
c o l l e c t i n g and r e c o n s t r u c t i n g b r y o z o a n c o l o n i e s from t h e t y p e - C i n c i n n a t i a n
rocks. It is to the p e r s e v e r a n c e of s u c h folk that we know that local bryozoan
c o l o n i e s did, in fact, s o m e t i m e s e x c e e d the size of basketballs. I n d e e d , o n e
colony re-assembled by Mr. F i n e is s o m e 66 cm by 35 cm by 15 cm ( F i g u r e

Bryozoans 89
Figure 7.4. Large bryozoan colonies from the type-Cincinnatian. A. Intact colony of a branching trepos-
tome bryozoan, Parvohallopora ramosa d'Orbigny, University of Cincinnati collections, Corryville Mem-
ber of Grant Lake Limestone, Hamilton Co., Ohio. Scale in mm. B. Intact colony of an unidentified
branching trepostome bryozoan. CMC IP uncatalogued, Cincinnatian, no horizon or locality data. Scale in
mm. C. Intact colony of trepostome bryozoan, Monticulipora mammulata d'Orbigny, CMC IP uncata-
logued, Cincinnatian, no horizon or locality data. Scale in mm. D. Trepostome bryozoan encrusted on
probable nautiloid shell that has disappeared, Monticulipora mammulata d'Orbigny, University of Cincin-
nati collections, Corryville Member of Grant Lake Limestone, Hamilton Co., Ohio. Scale 2 cm in length.
Note borings into bryozoan. E. Intact colony of trepostome bryozoan, Heterotrypa frondosa
d'Orbigny, CMC IP, Corryville Member of Grant Lake Limestone, Kenton Co., Kentucky. Colony about 65
cm in width. This colony was excavated and reassembled by Ron Fine. See Cuffey and Fine (2005).

90 A Sea without Fish

 Figure 7.5. Reconstruc-
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-
printed by permission of
Sigma Gamma Epsilon.

7 4 1 , ) . a w h o p p i n g 26 i n c h e s by 14 i n c h e s by 6 i n c h e s ( C u f f e y a n d F i n e 2005).
S i m i l a r efforts b y E r i c k s o n a n d W a u g h (2002), W a u g h a n d E r i c k s o n (2002),
and by W a u g h et al. (2004) p r o v i d e d new i n f o r m a t i o n a b o u t t h e f o r m a n d
patterns o f water f l o w t h r o u g h c o m p l e t e c o l o n i e s ( F i g u r e 7.5).
A n o t h e r s p e c t a c u l a r g r o w t h o f b r y o z o a n s w a s f o u n d i n a r o a d c u t sev-
eral m i l e s s o u t h o f M a y s v i l l e , K e n t u c k y . H e r e w e r e d i s c o v e r e d t w o b r o a d
mounds of bryozoans, each b e t w e e n three and three and a half meters
w i d e by a b o u t o n e - t h i r d m e t e r tall (10 feet by 1 foot). L i k e M r . F i n e ' s s p e c i -
m e n , t h e s e m o u n d s grew o n t h e sea floor, b u t , u n l i k e h i s , e a c h o f t h e
M a y s v i l l e m o u n d s consists o f n u m e r o u s i n d i v i d u a l c o l o n i e s a n d m a y h a v e
taken a t h o u s a n d years to g r o w to that s i z e ( C u f f e y 1998).
W h e n o r g a n i s m s p r o d u c e e l e v a t i o n s o n t h e sea f l o o r , s u c h c o n s t r u c t s
are technically t e r m e d b i o h e r m s ( " b i o " m e a n s "life," a n d " h e r m " m e a n s
" m o u n d " ) . T h e G r e a t Barrier R e e f i s t h e m o s t s p e c t a c u l a r e x a m p l e o f this
p h e n o m e n o n in today's o c e a n s . It must be a d m i t t e d that in c o m p a r i s o n to
the string of coral reefs that stretch m o r e t h a n 2000 km (1260 statute miles)
parallel to the east coast of A u s t r a l i a , t h e m o u n d s in N o r t h e r n K e n t u c k y are
less than miniscule. However, to the fossil c o l l e c t o r u s e d to b r y o z o a n frag-
m e n t s m u c h s m a l l e r than p i e c e s o f b l a c k b o a r d c h a l k , the b r y o z o a n m o u n d s
are g a r g a n t u a n . O n a j o c u l a r note, R o g e r C u f f e y , o n e o f t h e m o s t p r o d u c t i v e
b r y o z o a n workers alive today, l o n g has referred to t h e M a y s v i l l e m o u n d s a n d
such as " b r y o h e r m s . " A l a s ! T h i s p a r a g r a p h m u s t c l o s e on a sad note: s o m e -
t i m e d u r i n g the 1990s, road w i d e n i n g d e s t r o y e d the M a y s v i l l e m o u n d s .
There is, of c o u r s e , s o m e s o l a c e in the possibility that e v e n b i g g e r b r y o h e r m s
still may be buried s o m e w h e r e in t h e local rocks, a w a i t i n g discovery.

I n m o s t l o c a l i t i e s a n d i n most strata, b r y o z o a n s are t h e m o s t c o m m o n fos- Associations

sils o n e e n c o u n t e r s . I n s h e e r a b u n d a n c e a l o n e , t h e y m u s t b e r e c k o n e d t o

Bryozoans 91
 h a v e b e e n t r u l y i m p o r t a n t d e n i z e n s o f t h e C i n c i n n a t i a n sea floor. T h i s
i m p r e s s i o n i s n o t h i n g b u t e n h a n c e d w h e n o n e f o c u s e s i n o n t h e details o f
just w h e r e a n d w i t h w h o m t h e y 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 sea b o t t o m a p -
p e a r s t o h a v e b e e n soft m u d . M o s t o f t h e k i n d s o f a n i m a l s that w e g e n e r a l l y
f i n d p r e s e r v e d a s fossils d o n o t s e e m t o h a v e " l i k e d " soft, m u d d y b o t t o m s ,
p r e s u m a b l y b e c a u s e t h e m u d did n o t p r o v i d e solid f o o t i n g s u p o n w h i c h t o
b u i l d a stable life. It w a s all too e a s y to be e n g u l f e d by the o o z e . M o r e o v e r ,
t h e f i n e s e d i m e n t w a s t o o easily s w i r l e d u p into t h e w a t e r and c l o g g e d re-
spiratory a n d f o o d - g a t h e r i n g a p p a r a t i . l i v e n a c e n t i m e t e r or t w o a b o v e the
m u d w a s m o r e h o s p i t a b l e . B r y o z o a n s o r d i n a r i l y did n o t g r o w their c o l o n i e s
d i r e c t l y on t h e s u r f a c e of t h e m u d . B u t let a storm d r o p a few shells or frag-
m e n t s o f shells o n t o t h e g o o , a n d t h e sea floor w a s o p e n for c o l o n i z a t i o n .
So often, w h e n o n e is able to e x a m i n e the actual base of a colony w h e r e
g r o w t h c o m m e n c e d o n e d i s c o v e r s that the c o l o n y was f o u n d e d o n a frag-
m e n t of a shell of a b r a c h i o p o d or p e l e c y p o d , if n o t a c o m p l e t e or n e a r l y
c o m p l e t e shell ( F i g u r e s 7.3F, G , 7.4D). O n c e t h e sea floor w a s a bit stabi-
l i z e d , the b r y o z o a n s c o l o n i z e d a n d g r e w i n e a r n e s t .
O n c e e s t a b l i s h e d , t h e b r y o z o a n s t h e m s e l v e s a d d e d t o t h e stability o f
t h e sea floor in t h e i r i m m e d i a t e vicinity. First, as c o l o n i e s g r e w , a c e r t a i n
p r o p o r t i o n o f t h e m t o p p l e d over, a n d t h e i r skeletal m a t e r i a l b e c a m e incor-
p o r a t e d a s p a r t o f t h e sea f l o o r , t h u s i n c r e a s i n g t h e stability o f the b o t t o m
a n d m a k i n g i t m o r e h o s p i t a b l e for o t h e r c r e a t u r e s . M o r e o v e r , the little
" t h i c k e t s " o f b r y o z o a n c o l o n i e s p r o v i d e d p l a c e s for o t h e r o r g a n i s m s t o h i d e
f r o m p o t e n t i a l p r e d a t o r s or, i n t h e c a s e o f t h e p r e d a t o r s , p l a c e s f r o m w h i c h
t o o r c h e s t r a t e a m b u s h e s o f p o t e n t i a l prey. I n a d d i t i o n , s o m e a n i m a l s s c r a m -
b l e d u p t h e stalks a n d b r a n c h e s o f b r y o z o a n c o l o n i e s t o avoid t h e m u d d y
w a t e r i m m e d i a t e l y a d j a c e n t t o t h e sea floor. A n d l a r v a e that h a p p e n e d t o
a t t a c h u p i n a b r y o z o a n " t h i c k e t " w o u l d n o t o n l y h a v e e s c a p e d the worst
of the turbid water, but also m i g h t have had a better c h a n c e of latching
o n t o m i n u t e p a r t i c l e s o f f o o d s u s p e n d e d i n the water. M o r e o v e r , e v e n a s
trees a m e l i o r a t e t h e effects o f w i n d o n t h e l a n d , b r y o z o a n c o l o n i e s m u s t
h a v e m o d e r a t e d t h e c u r r e n t s o n t h e sea f l o o r . This w o u l d n o t o n l y h a v e
m a d e life easier for s o m e o r g a n i s m s , b u t w o u l d h a v e resulted i n e n t r a p p i n g
s e d i m e n t , t h e r e b y f u r t h e r h e l p i n g t o s t a b i l i z e t h e sea f l o o r .
O n e p a r t i c u l a r l y i n t r i g u i n g e x a m p l e o f how b r y o z o a n c o l o n i e s w e r e
used b y o t h e r c r e a t u r e s w a s d o c u m e n t e d b y D o u g l a s Shrake o f the O h i o
D i v i s i o n of G e o l o g i c a l S u r v e y in his master's thesis. Trilobites, like other
a r t h r o p o d s , are e n c l o s e d w i t h i n a hard exoskeleton. B e c a u s e this "suit of
a r m o r " c a n n o t e x p a n d a s the a n i m a l g r o w s , the trilobite p e r i o d i c a l l y m u s t
s h e d its e x o s k e l e t o n , e x p a n d in size, a n d h a r d e n up a n e w protective shield.
This is an e s p e c i a l l y t r y i n g t i m e for the trilobite, first b e c a u s e it c a n be dif-
f i c u l t t o w r i g g l e a n d s q u i r m o u t o f the old a r m o r , a n d , s e c o n d , b e c a u s e until
the n e w suit h a r d e n s , the a n i m a l is " n a k e d " a soft, t e m p t i n g morsel for any
p a s s i n g predator. S h r a k e f o u n d e v i d e n c e that i n d i v i d u a l s of the trilobite ge-
nus Primaspis resorted to l o w l y b r y o z o a n s to m a k e the t i m e of trial a bit less
t r y i n g ( S h r a k e 1987, 1989).

92 A Sea without Fish

 W h e n t h e t i m e for s h e d d i n g w a s a t h a n d , t h e t r i l o b i t e a p p a r e n t l y
c l i m b e d its w a y up into a s u i t a b l e part of a b r y o z o a n c o l o n y a n d w e d g e d
t h e p r o j e c t i o n s o f its e x o s k e l e t o n into t h e b r y o z o a n (see F i g u r e s 1 1 . 6 E , F )
T h i s e n a b l e d the trilobite t o pull itself o u t o f t h e old e x o s k e l e t o n a n d c o m -
m e n c e t h e h a r d e n i n g o f t h e n e w o n e , all t h e w h i l e b e i n g h i d d e n a m o n g s t
the b r y o z o a n fronds f r o m t h e e y e s o f w o u l d - b e p r e d a t o r s . A l t h o u g h this
m u s t h a v e b e e n a c o n v e n i e n t a r r a n g e m e n t for t h e trilobite, it may h a v e
b e e n less so for t h e b r y o z o a n s . S h r a k e f o u n d t h a t , in s o m e i n s t a n c e s , there-
w a s p a t h o l o g i c g r o w t h i n the c o l o n y a s t h e b r y o z o a n s grew u p a n d a r o u n d
the trilobite e x o s k e l e t o n t h e y h a d n o w a y t o d i s l o d g e .
T h e t r i l o b i t e / b r y o z o a n association d e s c r i b e d b y D o u g S h r a k e i s just o n e
of a host of e x a m p l e s of the interactions of b r y o z o a n s and m y r i a d o t h e r crea-
tures. O n the o n e h a n d , a b r y o z o a n larva w o u l d attach t o a l m o s t a n y o n e ,
given suitable c i r c u m s t a n c e s , and a c o l o n y w o u l d sprout. Bryozoan c o l o n i e s
have b e e n d o c u m e n t e d as a t t a c h e d to, e n c r u s t i n g , o v e r g r o w i n g , or e t c h e d
into articulate b r a c h i o p o d s , inarticulate b r a c h i o p o d s , c e p h a l o p o d s , c o r a l s ,
cornulitids, c r i n o i d s , foraminifers, h y d r o z o a n s , m o n o p l a c o p h o r a n m o l l u s c s ,
p e l e c y p o d s , trilobites, a n d , o f c o u r s e , o t h e r b r y o z o a n s , b o t h o f the s a m e a n d
of different species (see chapter 16, T a b l e 3). On the o t h e r h a n d , a n u m b e r of
other organisms have b e e n found a t t a c h e d to or b o r e d into b r y o z o a n s : c o r a l s ,
articulate brachiopods, inarticulate b r a c h i o p o d s , c o r n u l i t i d s , and p e l e c y p o d s ,
a l o n g with a n u m b e r of o r g a n i s m s of u n c e r t a i n affinities, i n c l u d i n g Catel-
locaula. Sanctum, Sphenothallus, a n d Trypanites (see c h a p t e r 16, Table 3).
In s o m e c a s e s , it is o b v i o u s that b o t h t h e " g u e s t " a n d t h e " h o s t " w e r e
alive a t t h e t i m e o f the a s s o c i a t i o n . I n o t h e r c a s e s , t h e " g u e s t " w a s merely
u s i n g a d e a d s h e l l , e x o s k e l e t o n , or w h a t e v e r as a h a n d y site for a t t a c h m e n t
on the sea floor. In o t h e r w o r d s , it c o m m o n l y is a t o u g h task to u n r a v e l
in-life a s s o c i a t i o n f r o m p o s t - m o r t e m h a p p e n s t a n c e . N o n e t h e l e s s , it is
a b u n d a n t l y o b v i o u s that the C i n c i n n a t i a n sea floor o f t h e a n c i e n t past, a n d ,
h e n c e , the C i n c i n n a t i a n rocks a n d fossils w e find today w o u l d h a v e b e e n
drastically different w i t h o u t the bryozoans.

O c c a s i o n a l l y a lucky c o l l e c t o r will find in the rocks of t h e C i n c i n n a t i a r e a Ordovician

a stone o b j e c t that looks rather like a d o u g h n u t ( F i g u r e s 7 . 3 H , I). C l o s e r Doughnuts
e x a m i n a t i o n reveals that this toroid fossil consists o f b r y o z o a n z o o e c i a ;
i n d e e d , it is a r i n g - s h a p e d z o a r i u m .
The b r y o z o a n rings have b e e n k n o w n for a l o n g time. Years a g o , w h e n
K e n n e t h F. Caster, the e m i n e n t paleontologist at the University of C i n c i n -
nati, was s h o w n o n e of t h e m by a local fossil collector, he q u i p p e d , " A h ! Yes!
A W e i c h o l d D o u g h n u t . " He then went on to e x p l a i n that W e i c h o l d was o n e
of the old-time collectors in the C i n c i n n a t i r e g i o n , a n d that these u n u s u a l
fossils had b e e n d u b b e d " W e i c h o l d d o u g h n u t s " or " W e i c h o l d rings," a l t h o u g h
h e did not know the specific c o n n e c t i o n b e t w e e n W e i c h o l d a n d t h e rings.
So why w o u l d a bryozoan z o a r i u m g r o w in the s h a p e of a ring? W e i c h -
old d o u g h n u t s tend to be s o m e 5 or 6 cm in d i a m e t e r (22 1/2 in). As it h a p -
p e n s , that d i a m e t e r is c o m p a r a b l e to that of the shells of s o m e of t h e o r t h o c -
o n i c c e p h a l o p o d s i n t h e l o c a l rocks. C o u l d there b e a c o n n e c t i o n ? W i t h i n

Bryozoans 93
 s o m e of the W e i c h o l d d o u g h n u t s , there is a r i n g of w h a t m i g h t be recrystal-
l i z e d c e p h a l o p o d shell ( F i g u r e 7.31). Perhaps the apertural part of the t u b e
that c o m p r i s e s the shell of an o r t h o c o n i c nautiloid c e p h a l o p o d broke off and
c a m e t o rest o n t h e sea floor. T h e n , o n e o r m o r e b r y o z o a n larvae settled o n
this hard o b j e c t p r o t r u d i n g a b o v e t h e o o z e . A s the z o a r i u m grew, i t a s s u m e d
t h e r i n g - s h a p e o f the " s e g m e n t " o f c e p h a l o p o d shell.
S u c h rings of cephalopod shells h a v e b e e n d e s c r i b e d and figured in the
scientific literature ( T e e t e r 1978), a n d s i m i l a r t h i n g s h a v e b e e n found in the
l o c a l rocks. H o w e v e r , t h e story m a y n o t b e q u i t e s o straightforward. T h e
p r o b l e m is that s o m e of the W e i c h o l d d o u g h n u t s s e e m to be b r y o z o a n hard
parts all the w a y t h r o u g h w i t h n o o b v i o u s r e m n a n t s o f c e p h a l o p o d shell.
F r a n k M c K i n n e y , the w e l l - k n o w n b r y o z o a n worker, has seen a ring-
s h a p e d c o l o n y of t h e b r y o z o a n g e n u s Constellaria, w i t h m u d in the center.
His interpretation was that t h e c o l o n y h a d slowed the water a n d c a u s e d m u d
to precipitate to s u c h an e x t e n t that g r o w t h of the c o l o n y was able to p r o c e e d
o n l y a t t h e p e r i p h e r y ( M c K i n n e y , pers. c o m m . ) . H o w e v e r , this c o l o n y was
s o m e 8 to 10 i n c h e s across ( 2 0 - 2 5 c m ) m o r e than t w i c e as b i g as t h e largest
W e i c h o l d rings. O b v i o u s l y , the p h e n o m e n o n n e e d s s o m e serious scientific
study.

Ordovician As m e n t i o n e d above, m a n y b r y o z o a n c o l o n i c s in the rocks of the C i n c i n -

Hitch-Hikers nati r e g i o n o r i g i n a l l y g r e w o n shells o n t h e sea floor. I n s o m e c a s e s , t h e

shell no l o n g e r s h e l t e r e d its m a k e r , b u t w a s m e r e l y a lifeless, h a r d o b j e c t
l y i n g o n t h e m u d . I n o t h e r c a s e s , b o t h the b r y o z o a n s a n d the o r g a n i s m s o n
w h i c h t h e z o a r i u m g r e w w e r e a l i v e . I n t h e s e i n s t a n c e s , t h e a t t a c h e r s are
c a l l e d e p i z o a , a n d t h e a t t a c h e e i s t h e h o s t ( D a v i s e t al. 1999).
S o m e b r y o z o a n s carried e p i z o i s m to a h i g h e r level.
N o t t o o u n c o m m o n l y , an o b s e r v a n t fossil c o l l e c t o r will find a fossil
that has t h e s i z e a n d s h a p e o f a n o r t h o c o n i c c e p h a l o p o d . H o w e v e r , u n l i k e
t h e c a s e o f a n o r d i n a r y n a u t i l o i d , t h e s u r f a c e b e a r s t h e tell-tale a p e r t u r e s
o f z o o e c i a ( F i g u r e 7 . 3 A ) a n d l o o k s are n o t d e c e i v i n g . T h e s p e c i m e n i s a n
o r t h o c o n i c n a u t i l o i d , b u t o n e that b e a r s a t h i n c o a t i n g that consists of a
b r y o z o a n colony. It is o b v i o u s t h a t t h e cephalopod shell w a s n o t just l y i n g
a r o u n d on t h e sea floor d e a d a n d e m p t y , b e c a u s e its entire exterior is cov-
e r e d b y t h e e n c r u s t e r w i t h n e i t h e r g a p s nor s e a m s . M o r e o v e r , t h e p i c t u r e
i s e n h a n c e d b y t h e s u r f a c e f e a t u r e s o f t h e z o a r i u m . Instead o f b e i n g e q u i -
d i m e n s i o n a l b u m p s , t h e m o n t i c u l e s are d e c i d e d l y e l o n g a t e , and their lon-
gest d i m e n s i o n is a l i g n e d w i t h the l e n g t h of t h e o r t h o c o n i c s h e l l . It is al-
m o s t a s t h o u g h the b r y o z o a n c o l o n y w a s c a r r i e d t h r o u g h t h e C i n c i n n a t i a n
sea o n t h e s w i m m i n g cephalopod. The w h o l e c e p h a l o p o d / b r y o z o a n as-
s e m b l a g e l o o k s s o s t r e a m l i n e d that e v e n the m o n t i c u l e s are d i s p o s e d s o a s
t o m i n i m i z e t h e f r i c t i o n o f s l i p p i n g t h r o u g h the water.
This b r y o z o a n / c e p h a l o p o d a s s o c i a t i o n h a s b e e n k n o w n for w e l l over
a c e n t u r y ( U l r i c h 1883). In fact, at least o n e taxon of b r y o z o a n s , Spatiopora,
s e e m s t o b e k n o w n o n l y a s e n c r u s t a t i o n s o n c e p h a l o p o d s (Baird e t al.
1989). E a r l y o n , t h e a s s o c i a t i o n w a s i n t e r p r e t e d as a parasite/host relation-
s h i p . H o w e v e r , in p a r a s i t i s m , t h e parasite c o n s u m e s part of the host. In the

94 A Sea without Fish

 Figure 7.6. A. Detailed
studies of bryozoans re-
quire carefully oriented
thin-sections or acetate-
peels. Diagram shows
orientation of sections
and 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
section of same. Both
approx. x 10.

b r y o z o a n / c e p h a l o p o d a s s o c i a t i o n , it is n o t likely that t h e b r y o z o a n s w e r e
" e a t i n g " the c e p h a l o p o d . It is possible that t h e y w e r e d e r i v i n g n u t r i t i o n
from leftovers a n d o r g a n i c d e b r i s g e n e r a t e d w h e n the c e p h a l o p o d , itself,
fed. It c o u l d be that the b r y o z o a n s p i c k e d up s u s p e n d e d m a t t e r from the
sea water a s the c e p h a l o p o d s w a m f r o m p l a c e t o p l a c e .
At first g l a n c e , o n e might worry that the weight of a "stony b r y o z o a n "
would have i m p e d e d significantly the s w i m m i n g of the c e p h a l o p o d . However,
the b r y o z o a n c o l o n y is just o n e z o o e c i u m thick and w o u l d have b e e n mostly
soft parts. Moreover, like present-day Nautilus, the O r d o v i c i a n c e p h a l o p o d
may have been able to c o m p e n s a t e for the extra weight of the bryozoans by

Bryozoans 95
 m e a n s of the gas in its c a m e r a e (see chapter 9). In addition, the b r y o z o a n coat-
i n g m i g h t have increased the hydrodynamic drag on the c e p h a l o p o d .
T h e r e even may have been sonic advantages to h a v n i n g a coating of
b r y o z o a n s . P r e s e n t - d a y " d e c o r a t o r c r a b s " arc c a m o u f l a g e d by the load of
a n e m o n e s a n d s u c h like that t h e y carry. I n d e e d , t h e c r a b s d e l i b e r a t e l y
" p l a n t " o t h e r c r e a t u r e s o n their dorsal s u r f a c e s . Perhaps the b r y o z o a n epi-
z o a h e l p e d c o n c e a l t h e C i n c i n n a t i a n c e p h a l o p o d s that b o r e t h e m . (Of
c o u r s e , t h e z o a r i u m w o u l d h a v e c o v e r e d , a n d t h e r e b y m a d e visually use-
less, a n y c o l o r p a t t e r n s that t h e c e p h a l o p o d s h a d ; but that is a story that
d o e s n o t b e l o n g i n t h e c h a p t e r a b o u t p h y l u m Bryozoa.)

Studying Bryozoans Most b r y o z o a n s p e c i m e n s c a n be identified o n l y on the basis of internal

s t r u c t u r e s , at least d e f i n i t i v e l y so. This m e a n s that, e x c e p t for S u p e r m a n ,
it is n e c e s s a r y to c u t t h e m o p e n n o s m a l l feat for t h e o r d i n a r y fossil c o l -
lector. It is n e c e s s a r y to u s e a r o c k saw to m a k e p r e c i s e l y o r i e n t e d c u t s
t h r o u g h a n i n d i v i d u a l s p e c i m e n . B e c a u s e the b r y o z o a n c o l o n y i s preserved
as p a r t of t h e r o c k , it w i l l n o t let e n o u g h light t h r o u g h to sec internal d e -
tails. This c a n b e o v e r c o m e i n t w o w a y s . T h e z e a l o u s a n d w e l l - e q u i p p e d
p a l e o n t o l o g i s t c a n c u t a n d g r i n d t h e s p e c i m e n into slices s o t h i n that t h e y
b e c o m e transparent. T h e s e are c a l l e d t h i n - s e c t i o n s , a n d t h e y arc w h a t
g e n e r a l l y arc u s e d i n s t u d y i n g b r y o z o a n s .
D e p e n d i n g on the n a t u r e of t h e s p e c i m e n , alternatively, it may be
p o s s i b l e to u s e w h a t are c a l l e d a c e t a t e p e e l s . Like a t h i n - s e c t i o n , a p e e l
starts w i t h a c a r e f u l l y o r i e n t e d c u t t h r o u g h the s p e c i m e n . T h e cut s u r f a c e
is t h e n c a r e f u l l y g r o u n d flat a n d t h e n e t c h e d in an a p p r o p r i a t e acid of ap-
p r o p r i a t e " s t r e n g t h . " II the s p e c i m e n is suitably p r e s e r v e d , c e r t a i n of its
features will be a bit m o r e resistant to d i s s o l u t i o n by the a c i d . H e n c e , t h e y
w i l l s t a n d o u t slightly from t h e s u r f a c e . If o n e takes a t h i n sheet of plastic
(the acetate) a n d uses a c e t o n e to allow the a c e t a t e to a d h e r e to the e t c h e d
s u r f a c e , it m a y be p o s s i b l e to p u l l a w a y the s h e e t , a l o n g w i t h e n o u g h of the
s p e c i m e n , t o reveal i n t e r n a l d e t a i l s .
T h e u s e o f t h i n - s e c t i o n s i s g e n e r a l l y c o n s i d e r e d t o b e " i n d u s t r y stan-
d a r d , " b u t , r e g a r d l e s s of w h i c h s t u d y t e c h n i q u e is u s e d , b o t h r e q u i r e e q u i p -
m e n t that m a y b e b e y o n d t h e b u d g e t , o r d e s i r e , o f the o r d i n a r y fossil c o l -
lector. Moreover, both require definite safety p r e c a u t i o n s a n d safety
e q u i p m e n t ; for e x a m p l e , a c i d s c a n e t c h m o r e t h a n just rock, a n d a c e t o n e
not only is flammable, but its v a p o r is u n h e a l t h y to b r e a t h e .
As indicated, both thin-sections and peels must be carefully oriented
within the zoarium. This i s t o m a x i m i z e t h e i n f o r m a t i o n that m a y b e d e -
rived a b o u t the internal s t r u c t u r e o f the c o l o n y . B e c a u s e there are different
k i n d s of z o o i d s in a c o l o n y , it is i m p o r t a n t to be a b l e to study t h e different
sizes, s h a p e s , a n d n a t u r e s of t h e z o o e c i a as v i e w e d in a p l a n e p e r p e n d i c u l a r
to t h e i n d i v i d u a l t u b e s . This is best d o n e in a t a n g e n t i a l s e c t i o n , w h i c h is
c u t p a r a l l e l to the s u r f a c e of the c o l o n y a n d n e a r its s u r f a c e (as o p p o s e d to
n e a r its c e n t e r , o r axis) ( f i g u r e 7.6). O n t h e o t h e r h a n d , a l o n g i t u d i n a l
set lion is cut p a r a l l e l to t h e l e n g t h of t h e i n d i v i d u a l t u b e s a n d c a n p r o v i d e
i m p o r t a n t i n f o r m a t i o n o n b o t h t h e g r o w t h o f the z o a r i u m a n d o f t h e z o o e -

96 A Sea without Fish

cia of w h i c h it consists. A t r a n s v e r s e s e c t i o n is o r i e n t e d at right a n g l e s to
the o t h e r t w o , for e x a m p l e , across a b r a n c h of a g i v e n c o l o n y . In s h o r t , o n e
n e e d s l o n g i t u d i n a l , t a n g e n t i a l , a n d transverse s e c t i o n s of a c o l o n y to g e t a
c o m p l e t e p i c t u r e o f its i n t e r n a l s t r u c t u r e , a n d this c o m p l e t e p i c t u r e i s es-
sential t o a n u n d e r s t a n d i n g o f just w h a t k i n d o f b r y o z o a n i s a t h a n d a n d
h o w it grew and was c o n s t i t u t e d .
After t h e p r o p e r l y o r i e n t e d t h i n - s e c t i o n s o r p e e l s are m a d e , t h e o n l y
way they can be studied adequately is under the m i c r o s c o p e . T h i s , again,
is a p i e c e of e q u i p m e n t that m a y c h a l l e n g e one's b u d g e t .

Bryozoans 97
98 A Sea without Fish
BRACHIOPODS: THE OTHER BIVALVES

B r a c h i o p o d s a r e a m o n g the m o s t c o m m o n fossils i n the O r d o v i c i a n r o c k s o f

t h e C i n c i n n a t i area. O n l y fossils o f b r y o z o a n s are m o r e n u m e r o u s t o t h e
n a k e d eye. In a s t u d y of t y p e - C i n c i n n a t i a n l i m e s t o n e s , M a r t i n ( 1 9 7 5 ) re-
ported that b r a c h i o p o d s a n d b r y o z o a n s t o g e t h e r c o n s t i t u t e a b o u t 6 0 p e r c e n t
of the fossil fragments c o m p r i s i n g the l i m e s t o n e s . There e v e n are s o m e lay-
ers, for e x a m p l e , in the B e l l e v u e Limestone, in w h i c h the rock is a veritable
c o q u i n a , i n this c a s e c o n s i s t i n g o f c o m p l e t e a n d n e a r l y c o m p l e t e shells o f
large, f l a t b r a c h i o p o d s o f a s i n g l e g e n u s . T h e s e aptly n a m e d " s h i n g l e d
Rafinesquina b e d s " c o m m o n l y are t h o u g h t of as r e m a i n s o f v e r y shallow-
water deposits r e m i n i s c e n t o f the s h i n g l e d b e a c h e s o f today. A l t h o u g h t h e y
have b e e n living o n E a r t h s i n c e the C a m b r i a n Period, b r a c h i o p o d s are n o t
w e l l - k n o w n a n i m a l s to m o s t of us. In fact, many folks c o n f u s e t h e m w i t h that
g r o u p o f m o l l u s c s that i n c l u d e s t h e c l a m s . M e m b e r s o f t h e p h y l u m B r a c h i o -
p o d a and those o f the m o l l u s c a n class P e l e c y p o d a are bivalved a n i m a l s , that Figure 8 . 1 . Comparison of
is, e a c h has a shell that consists of t w o valves. B u t there the r e s e m b l a n c e e n d s . a brachiopod with a
The b r a c h i o p o d s and p e l e c y p o d s are o t h e r w i s e strikingly different a n i m a l s . pelecypod. A and B,
First, t h e o r i e n t a t i o n s o f p e l e c y p o d s a n d b r a c h i o p o d s a r c d i f f e r e n t pelecypod. C and D,
Platystrophia ponde-
( F i g u r e 8.1). The t w o valves of a c l a m are a n a t o m i c a l l y left a n d r i g h t in
rosa showing sulcus and
p o s i t i o n , w i t h the h i n g e c o n n e c t i n g the v a l v e s l o c a t e d a t t h e top o f t h e a n i -
fold. Drawings from
m a l ( t e c h n i c a l l y c a l l e d d o r s a l ) . H o w e v e r , the t w o v a l v e s o f a b r a c h i o p o d
Meek (1873), courtesy of
are dorsal a n d v e n t r a l , r e s p e c t i v e l y , a n d t h e h i n g e is at t h e rear of t h e a n i -
the Ohio Department of
mal (posterior). T h u s , a l t h o u g h b o t h p e l e c y p o d s a n d b r a c h i o p o d s are bi- Natural Resources Divi-
laterally s y m m e t r i c a l a n i m a l s , the p l a n e s of s y m m e t r y of t h e t w o are at a sion of Geological
right a n g l e t o o n e a n o t h e r ( F i g u r e 8.1). W e c a n c o n c l u d e f r o m t h i s t h a t , Survey.
although animals of both groups each have two valves, "bivalvedness" in
e a c h g r o u p e v o l v e d i n d e p e n d e n t l y ; t h e t w o g r o u p s are n o t a t all closely
related, and n e i t h e r e v o l v e d from t h e o t h e r .
F r o m a p r a c t i c a l point of view, h o w e v e r , it h a p p e n s that t h e d i f f e r e n c e
in o r i e n t a t i o n g e n e r a l l y p r o v i d e s a c o n v e n i e n t w a y to tell fossil b r a c h i o p o d s
from fossil p e l e c y p o d s . E a c h v a l v e o f m o s t b r a c h i o p o d s c a n b e d i v i d e d into
two h a l v e s that are m i r r o r i m a g e s o f o n e a n o t h e r ( F i g u r e 8 . 1 C ) . O n t h e
other h a n d , it is t h e t w o v a l v e s of m o s t fossil p e l e c y p o d s t h a t are t h e m i r r o r
i m a g e s o f o n e a n o t h e r ( F i g u r e 8.1 B). I n o t h e r w o r d s , the p l a n e o f s y m m e t r y
of an ordinary p e l e c y p o d is b e t w e e n the valves; in an ordinary b r a c h i o p o d ,
it is d o w n the m i d d l e of e a c h v a l v e .
M o r e o v e r , i n g e n e r a l , the t w o v a l v e s o f a b r a c h i o p o d shell are N O T
mirror i m a g e s o f o n e a n o t h e r ( f i g u r e 8 . 1 D ) . For e x a m p l e , o n e v a l v e may-
be decidedly d e e p e r than its o p p o s i t e . In a d d i t i o n (or i n s t e a d ) , o n e v a l v e
may h a v e a p o r t i o n a l o n g t h e m i d l i n e that is d i s t i n c t l y c o n v e x t o w a r d the

99
Figure 8.2. A. Cross- o u t s i d e of t h e a n i m a l (this s t r u c t u r e is c a l l e d a fold), a n d the o t h e r valve
section of an articulate m a y h a v e a d i s t i n c t c o n c a v i t y in t h e s a m e p o s i t i o n (called a sulcus).
brachiopod. B. Interior
A s the overall s y m m e t r y o f p e l e c y p o d s and b r a c h i o p o d s differs, s o too
views of pedicle valve
d o e s the o p e r a t i o n of the shells. In a p e l e c y p o d , the two valves arc joined at
(left) and brachial valve
the h i n g e by an elastic pad or l i g a m e n t . W h e n the shell is held c l o s e d , the
(right) of the Cincinnatian
l i g a m e n t is s t r e t c h e d , so that w h e n t h e a n i m a l relaxes the t w o valves g a p e
orthid brachiopod He-
bertella. Drawings by apart from o n e a n o t h e r . T o close t h e shell, the p e l e c y p o d a n i m a l m u s t c o n -

Kevina Vulinec. tract o n e o r t w o a d d u c t o r m u s c l e s (the n u m b e r d e p e n d s o n the t y p e o f

clam).
I n b r a c h i o p o d s , h o w e v e r , t h e r e i s n o l i g a m e n t . The a n i m a l m u s t c o n -
tract w h a t are c a l l e d d i d u c t o r m u s c l e s t o o p e n t h e shell ( F i g u r e 8.2A). I t
uses a d d u c t o r m u s c l e s t o c l o s e t h e s h e l l ; t h e s e a d d u c t o r s arc s t r e t c h e d
w h e n t h e shell is o p e n .

700 A Sea without Fish

 T h u s , w h e n a p e l e c y p o d r e l a x e s , its shell o p e n s . b y w a y o f c o n t r a s t ,
w h e n a b r a c h i o p o d r e l a x e s , its shell t e n d s to c l o s e . This has i m p o r t a n t
i m p l i c a t i o n s for how o n e finds p e l e c y p o d s a n d b r a c h i o p o d s as fossils. U p o n
d e a t h (the u l t i m a t e r e l a x a t i o n ) , t h e i n d i v i d u a l v a l v e s of a p e l e c y p o d tend
t o get separated f r o m o n e a n o t h e r , b e c a u s e t h e y g a p e a p a r t , a l l o w i n g c m -
rents, for e x a m p l e , t o tear t h e m a s u n d e r . O n t h e o t h e r h a n d , t h e t w o v a l v e s
o f a b r a c h i o p o d shell m o r e c o m m o n l y r e m a i n t o g e t h e r , a n d are f o u n d that
w a y by the intrepid fossil c o l l e c t o r .
B r a c h i o p o d shells t e n d t o b e better p r e s e r v e d t h a n are p e l e c y p o d shells
for a n o t h e r r e a s o n , too. M o s t b r a c h i o p o d shells c o n s i s t o f c a l c i u m c a r b o n -
ate, b u t s o d o p e l e c y p o d shells. H o w ever, c a l c i u m c a r b o n a t e exists i n m o r e
than o n e f o r m . M o s t b r a c h i o p o d shells arc o f t h e m i n e r a l c a l c i t e , w h e r e a s
p e l e c y p o d s consist o f o r i n c l u d e a r a g o n i t e . The atoms of c a l c i u m , c a r b o n ,
and o x y g e n are a r r a n g e d differently i n a r a g o n i t e a n d c a l c i t e , a n d the t w o
s u b s t a n c e s have different p r o p e r t i e s . B e c a u s e o f this, p e l e c y p o d s t e n d n o t
to be p r e s e r v e d as w e l l as b r a c h i o p o d s . The p r a c t i c a l result is that you may
f i n d m a i n w e l l - p r e s e r v e d b r a c h i o p o d s I n the r o c k s o f t h e C i n c i n n a t i a r e a ,
b u t p e l e c y p o d s , w i t h few e x c e p t i o n s , are p r e s e r v e d a s i n t e r n a l m o l d s .
B r a c h i o p o d s are f i l t e r feeders. T h e y e x t r a c t s m a l l p a r t i c l e s o f o r g a n i c
matter from t h e sea water. T h e s e p a r t i c l e s are c a p t u r e d by a c i l i a t e d s t r u c -
ture c a l l e d the l o p h o p h o r e (Plate 3F; F i g u r e 8.2A). ' T h e l o p h o p h o r e o c -
c u p i e s m u c h o f the s p a c e b e t w e e n the v a l v e s , f o r m i n g a p a i r o f t u b u l a r
" a r m s " that e x t e n d on e a c h side of t h e p l a n e of s y m m e t r y . A gutter-like food
g r o o v e runs a l o n g t h e a r m s from w h i c h c i l i a t e d t e n t a c l e s e x t e n d to f o r m a
filter (Plate 3F; F i g u r e 8.2A). T h e b e a t i n g of t h e c i l i a c a u s e s w a t e r to flow
into the shell, a l o n g o r t h r o u g h t h e t e n t a c l e s , a n d t h e n o u t o f t h e shell
a g a i n . The food p a r t i c l e s stick to t h e c i l i a a n d are t r a n s p o r t e d by t h e m to
the food g r o o v e a n d m o u t h , w h i c h i s l o c a t e d o n t h e c e n t e r l i n e o f t h e a n i -
m a l . As with all a n i m a l s , food is m e t a b o l i z e d , a n d w a s t e is e x p e l l e d .
In at least s o m e b r a c h i o p o d s , the l o p h o p h o r e s arc s u p p o r t e d by projec-
tions from the interior s u r f a c e of o n e valve. This s o - c a l l e d brachial valve is
the o n e that is a n a t o m i c a l l y dorsal in p o s i t i o n . In s o m e i n s t a n c e s , e a c h
branch of the l o p h o p h o r e is c o m p l e x l y c o i l e d ; in s u c h c a s e s , the l o p h o p h o r e
support m a y be c o i l e d , too. It is t h e t w o " a r m s " of t h e l o p h o p h o r e that give
the b r a c h i o p o d s their n a m e ; t h e a n c i e n t C r e e k w o r d " b r a c h i o n " m e a n s
"arm." The " p o d " part o f the n a m e c o m e s from " p o d o s , " o n e o f the parts o f
s p e e c h of the ancient Greek word "pous." w h i c h m e a n s "toot"; it recalls a
time w h e n b r a c h i o p o d s w e r e t h o u g h t t o b e c l o s e relatives o f t h e m o l l u s c s ,
w h i c h i n c l u d e the gastropods, p e l e c y p o d s , a n d c e p h a l o p o d s , a m o n g others.

Before p r o c e e d i n g further, it m u s t be a d m i t t e d that the p i c t u r e of b r a c h i o - Inarticulates vs.

pods painted a b o v e is a bit over-simplified. The b r a c h i o p o d s do n o t c o m p r i s e Articulates
a single, h o m o g e n e o u s l i n e a g e of a n i m a l s . The b r a c h i o p o d s portrayed a b o v e
mostly fall into a g r o u p c a l l e d a r t i c u l a t e b r a c h i o p o d s . T h e y are c a l l e d a r t i c u -
lates, b e c a u s e the t w o valves o f t h e shell are a r t i c u l a t e d t h e y are c o n n e c t e d
together by way of a w e l l - d e v e l o p e d h i n g e ( F i g u r e S.2B). A l o n g the h i n g e of
o n e valve arc projections, c a l l e d t e e t h , that fit into sockets in the h i n g e area

Brachiopods 101
 o f the o p p o s i n g valve. B e c a u s e o f the i n t e r l o c k i n g teeth and sockets, the so-
c a l l e d d e n t i t i o n , it w o u l d be difficult for a w o u l d - b e d e v o u r e r of b r a c h i o p o d
flesh to twist the valves apart to get at supper.
A n i m a l s o f t h e o t h e r m a j o r g r o u p o f b r a c h i o p o d s , not surprisingly, are
t e r m e d i n a r t i c u l a t e s . I n t h e s e a n i m a l s , t h e r e are n e i t h e r teeth nor sockets.
N o t having a real h i n g e , t h e task of k e e p i n g t h e valves t o g e t h e r is a greater
c h a l l e n g e for a n i n a r t i c u l a t e . T h e m u s c u l a t u r e i s a g o o d d e a l m o r e c o m -
p l i c a t e d in i n a r t i c u l a t e s t h a n in a r t i c u l a t e s t o k e e p the t w o valves from
b e i n g twisted apart from o n e another.
In an a r t i c u l a t e , t h e r e is a h i n g e , w h i c h serves as a f u l c r u m . The d i d u c -
f o r m u s c l e s a n d t h e a d d u c t o r m u s c l e s , i n order t o o p e n a n d c l o s e the shell,
o p e r a t e a g a i n s t o n e a n o t h e r a b o u t t h e f u l c r u m ( F i g u r e 8.2A). B u t a n inar-
t i c u l a t e has n o s u c h f u l c r u m . T h e a n i m a l o p e n s its shell, n o t b y c o n t r a c t -
i n g d i d u c t o r s , b u t b y p u l l i n g t h e b o d y b a c k toward the rear o f t h e shell,
thereby c a u s i n g the v a l v e s t o g a p e s u f f i c i e n t l y f o r the a n i m a l t o f e e d , re-
spire, a n d p e r f o r m o t h e r n e c e s s a r y a c t i v i t i e s .
A n o t h e r c o m m o n d i f f e r e n c e b e t w e e n articulates and inarticulates in-
volves the c o m p o s i t i o n of t h e shell. In m o s t inarticulates the shell consists,
not o f c a l c i u m c a r b o n a t e , b u t , rather, o f c a l c i u m p h o s p h a t e . ( N o t e , h o w e v e r ,
that this is n o t a universal r u l e , for the shells of s o m e of the a n i m a l s tradition-
alh c a l l e d inarticulates are- c a l c i u m c a r b o n a t e , like those of the articulates.)
S t u d i e s of p r e s e n t - d a y forms h a v e b e e n taken to s u g g e s t that t h e ar-
ticulate and inarticulate b r a c h i o p o d s may, in fact, not be particularly
c l o s e l y related. A n i n a r t i c u l a t e b r a c h i o p o d h a s b o t h m o u t h a n d a n u s ,
w h e r e a s a n a r t i c u l a t e has n o a n u s . T h e early life histories o f the a n i m a l s
o f the t w o g r o u p s a r e d i f f e r e n t , t o o ; for e x a m p l e , t h e p e d i c l e o f a n i n a r t i c u -
late has a d i f f e r e n t o r i g i n t h a n d o e s t h e p e d i c l e o f a n a r t i c u l a t e . D e t a i l e d
s t u d i e s o f t h e g e n e t i c s o f present-clay a n i m a l s are b e g i n n i n g t o t h r o w light
o n the issue o f t h e r e l a t i o n s h i p s o f t h e v a r i o u s g r o u p s o f b r a c h i o p o d s ( C o -
h e n a n d G a w t h r o p 1 9 9 7 ) . H o w e v e r , p e n d i n g t h e a m a s s i n g o f f u r t h e r in-
f o r m a t i o n , w e w i l l follow t h e u s u a l tradition o f c o n s i d e r i n g t h e A r t i c u l a t a
and the Inarticulata to be two subphyla of the phylum Brachiopoda. O n e
a l t e r n a t i v e s c h e m e w o u l d b e t o r e c o g n i z e t h o s e t w o g r o u p s a s separate
p h y l a . O n t h e third h a n d , s o m e e x p e r t s o n b r a c h i o p o d s prefer t o d o a w a y
w i t h the f o r m a l taxa A r t i c u l a t a a n d I n a r t i c u l a t a a l t o g e t h e r a n d r e c o g n i z e
instead t h r e e s u b p h y l a ( W i l l i a m s e t al. 2000); f o l l o w e r s o f that s c h e m e
retain t h e c o n c e p t s o f a r t i c u l a t e d b r a c h i o p o d s a n d i n a r t i c u l a t e d b r a c h i o -
p o d s , but only as descriptive terms.

Brachiopod R e g a r d l e s s of s u c h t a x o n o m i c issues, t h e b r a c h i o p o d s as a w h o l e are b e n -

Life Habits t h i c c r e a t u r e s . S o m e o f t h e m s i m p l y lie o n t h e sea floor, w h e r e a s o t h e r s

e a c h are p h y s i c a l l y a t t a c h e d to t h e sea floor by m e a n s of a fleshy stalk c a l l e d
a p e d i c l e . In 1972 Peter R i c h a r d s p u b l i s h e d a s t u d y that e x p l o r e d t h e rela-
t i o n s h i p b e t w e e n shell f o r m a n d life habits o f m a n y c h a r a c t e r i s t i c C i n c i n -
n a t i a n b r a c h i o p o d s . His r e s e a r c h d e m o n s t r a t e d that C i n c i n n a t i a n b r a c h i o -
p o d s h a v e a variety of life habits a n d that c a r e f u l a t t e n t i o n to m o r p h o l o g i c a l
features of the shell as w e l l as p r e s e r v a l i o n a l e v i d e n c e helps to e x p l a i n the

102 A Sea without Fish

diversity s e e n in this i m p o r t a n t g r o u p . His work has g u i d e d a n d inspired
m u c h s u b s e q u e n t r e s e a r c h a n d t h e d i s c u s s i o n that f o l l o w s

Inarticulates

S o m e p r e s e n t - d a y i n a r t i c u l a t e s ( g e n u s L i n g u l a a n d its relatives) s p e n d
m u c h o f their t i m e i n a b u r r o w . W h e n i t c o m e s t i m e t o f e e d , t h e p e d i c l e
e x t e n d s so that at least the a n t e r i o r part of t h e shell p r o j e c t s up into t h e
water. In some b r a c h i o p o d s , t h e r e is an o p e n i n g in t h e shell t h r o u g h w h i c h
the p e d i c l e e x t e n d s . In g e n e r a l , this p e d i c l e f o r a m e n is in t h e a n a t o m i c a l l y
l o w e r valve, w h i c h , h e n c e , i s c a l l e d t h e p e d i c l e v a l v e . T h e p e d i c l e o f s o m e
b r a c h i o p o d s may be a t t a c h e d to a n o t h e r shell on the sea floor; this m a y
result i n p e d i c l e a t t a c h m e n t scars w h i c h c o n s i s t o f t i n y , c h a r a c t e r i s t i c pits
in the other shell. In an i n a r t i c u l a t e b r a c h i o p o d , as in you, there is a m o u t h ,
esophagus, stomach, intestine, and anus. In an articulate b r a c h i o p o d , how-
ever, there i s n o a n u s ( f i g u r e 8.2A). W h a t m i g h t h a v e b e e n a c o m p l e t e
digestive tract is, in fact, a c u l de sac. T h u s , t h e o n l y e g r e s s for w a s t e m a t e -
rial i s b a c k o u t t h e m o u t h . I n p r e s e n t - d a y a n i m a l s that h a v e b e e n s t u d i e d ,
solid waste is r e g u r g i t a t e d as s m a l l p e l l e t s ; t h e s e are t h e n e x p e l l e d from t h e
shell by rapid s n a p p i n g of the v a l v e s .
A l t h o u g h the a r t i c u l a t e s are m o r e readily n o t i c e d , i n a r t i c u l a t e s are n o t
rare. M o s t o f t h e m , h o w e v e r , grew o n the shells o f o t h e r a n i m a l s a n d ,
h e n c e , are relatively s m a l l a n d easily o v e r l o o k e d ( F i g u r e 8 . 3 ) . P r e s u m a b l y ,
the other shells p r o v i d e d a solid substrate to w h i c h to a t t a c h ; it c o u l d w e l l
b e that the little " h a n g e r s - o n " ( t e c h n i c a l l y c a l l e d e p i z o a ) m a d e their l i v i n g
b y c o n s u m i n g the waste m a t t e r e x p e l l e d b y their " h o s t s . " I n a n y c a s e , t h e
shells of t h e s e i n a r t i c u l a t e s c o m m o n l y l o o k a bit like s c a b s , blisters, or little
v o l c a n o e s o n the shells o f a r t i c u l a t e b r a c h i o p o d s . S o m e o f t h e m g r e w s o
tightly affixed to their larger c o u s i n s that t h e r i b b i n g of t h e shells of t h e
latter d e f o r m or show t h r o u g h t h e shells of t h e i n a r t i c u l a t e s .
A n o t h e r kind of inarticulate may be f o u n d in the rocks of the C i n c i n n a t i
region by the s h a r p - e y e d c o l l e c t o r . G i v e n the m o n i k e r " i n a r t i c u l a t e , " it is
ironic that this o t h e r g r o u p c o m p r i s e s a n i m a l s that h a v e t o n g u e - s h a p e d
shells. Luckily for paleontologists, t h e s e s o - c a l l e d l i n g u l i d e s are represented
i n present-day o c e a n s , s o that w e readily c a n see t h e m . A n i n d i v i d u a l m e m -
ber of g e n u s Lingula (Latin for " t o n g u e " ) , f r o m w h i c h t h e g r o u p gets its
n a m e , has a way o f life r e m i n i s c e n t o f that o f s o m e b u r r o w i n g w o r m s . T h e
a n i m a l a n c h o r s the distal e n d of its l o n g p e d i c l e to the sea floor a n d uses its
shell to d i g vertically d o w n into the s e d i m e n t front e n d foremost. In d u e
course, the burrower veers to the h o r i z o n t a l a n d then b a c k up to the sea floor.
M o s t o f the U - s h a p e d burrow c o l l a p s e s from b e h i n d , s o that o n l y t h e o n e
vertical tube r e m a i n s , with the o p e n i n g of the shell at or n e a r the sea floor
and the p e d i c l e p o i n t i n g d o w n into the s e d i m e n t . W h e n n e c e s s a r y (or d e -
sired?), the g a p e b e t w e e n the valves c a n be raised a b o v e t h e sea floor by ex-
tension of the p e d i c l e , and the a n i m a l c a n f e e d , or whatever. In t i m e s of
danger, the a n i m a l c a n retreat into its burrow by c o n t r a c t i n g t h e p e d i c l e .
L i n g u l i d e s a r e not p a r t i c u l a r l y c o m m o n i n t h e t y p e - C i n c i n n a t i a n .
Very o c c a s i o n a l l y a s p e c i m e n is f o u n d w i t h its shell o r i e n t e d p e r p e n d i c u l a r

Brachiopods 103
104 A Sea without Fish
to the stratification, in w h a t a p p e a r s to be its life p o s i t i o n , t h a t is, t h e a n i - Figure 8.3. A. Pseudo-
mal's o r i e n t a t i o n in s p a c e d u r i n g its life. T h e i n d i v i d u a l in F i g u r e 8.3A is lingula sp., CMC IP

a case in p o i n t . 51994, Cincinnatian, ho-

rizon and locality un-
It is o n l y by great g o o d f o r t u n e that that p a r t i c u l a r i n d i v i d u a l m a d e it
known, x 1.9. Specimen
into a m u s e u m . R a l p h D u r y was a C i n c i n n a t i e n t o m o l o g i s t o f s o m e c o n -
oriented perpendicular to
siderable repute. However, his a c t u a l l i v e l i h o o d w a s in real estate. A c c o r d -
bedding, presumably in
i n g t o C h a r l e s D u r y , C h a r l e s ' s son a n d d i r e c t o r o f t h e C i n c i n n a t i M u s e u m
life position with beak
o f N a t u r a l History for m o r e t h a n f i v e d e c a d e s , the s p e c i m e n i n f i g u r e 8.3A downward. B. Trema-
started its m u s e u m c a r e e r i n a real estate p r o j e c t . C h a r l e s D u r y w a s h a v i n g tis millepunctata Hall,
a stone w a l l built. B e i n g a m e t i c u l o u s fellow, he visited his sites on a r e g u l a r CMC PT 585, brachial
and f r e q u e n t basis. H e h a p p e n e d t o m a k e o n e o f t h e s e visits t o t h e b u i l d i n g valve interior, Cincinna-
site shortly after a load of s t o n e h a d b e e n d e l i v e r e d . In t h e c o u r s e of e x a m - tian, horizon and locality

i n i n g the stone to be c e r t a i n that it w a s up to his s t a n d a r d s , he saw t h e unknown, x 3. C. Pet-

rocrania scabiosa (Hall)
c h u n k with the l i n g u l i d e i n its life p o s i t i o n . S o , instead o f e n d i n g u p a s
encrusting on Heber-
part o f a w a l l , t h e p i e c e o f rock e n d e d u p a s p a r t o f t h e c o l l e c t i o n s o f t h e
tellasp., MUGM 29461,
C i n c i n n a t i M u s e u m o f N a t u r a l H i s t o r y , a l o n g w i t h Charles D u r y ' s i n s e c t
Arnheim, Oxford, Ohio,
c o l l e c t i o n b u t that i s a n o t h e r s t o r y ( V u l i n e c a n d D a v i s 1984). x 1.3. D. Petrocrania
scabiosa (Hall) encrust-
ing on brachial valve of
Articulates
Rafinesquina sp., Bruce
T h e vast majority o f the b r a c h i o p o d s that o n e sees i n the O r d o v i c i a n rocks and Charlotte Gibson
of the C i n c i n n a t i area are a r t i c u l a t e s . T h e r e is a t r e m e n d o u s variety of t h e m Collection, no. 1042,

( f i g u r e s 8.4-8.9). S o m e h a v e a n external s h a p e that m a k e s t h e m easy t o Richmondian, Franklin

Co., Indiana, approx. x
identify t o g e n u s or, e v e n , s p e c i e s . O n t h e o t h e r h a n d , there are s o m e that
1.5. E. Philhedra lae-
are beastly difficult to tell apart. T h e p r o b l e m is that brachiopods that are
lia (Hall), MUGM 26219,
only distantly related may look alike externally ( F i g u r e 8.6). It is o n l y w h e n
two specimens encrusted
o n e c a r e f u l l y studies the various features o n t h e i n n e r s u r f a c e o f e a c h valve
on Rafinesquina bra-
that their true relationships may b e c o m e a p p a r e n t . U s e f u l features i n c l u d e chial valve exterior, Lib-
the scars w h e r e m u s c l e s a t t a c h e d to t h e v a l v e s , t h e s t r u c t u r e s that s u p p o r t e d erty Formation, Preble
the l o p h o p h o r e s , and s o o n . O n e e v e n m a y n e e d t o e x a m i n e the internal Co., Ohio, x 3. F.
structure of the shell material that m a k e s up t h e valves. It s e e m s that t h e Schizocrania filosa Hall,
e n v i r o n m e n t in w h i c h the brachiopods lived led different l i n e a g e s to e v o l v e CMC IP 36, encrusted on

similar external shapes. The p h e n o m e n o n is c a l l e d c o n v e r g e n t e v o l u t i o n , Rafinesquina pedicle

valve exterior, Corryville
and the result i s h o m e o m o r p h y t h e e x i s t e n c e o f t w o o r m o r e k i n d s o f a n i -
Formation, Warren Co.,
mals that are not closely related b u t that n o n e t h e l e s s look alike. T h i s is a
Ohio, x 1.8.
c o m m o n e n o u g h o c c u r r e n c e that it is an old saving: "Homeomorphy is rife
a m o n g s t the brachiopods." This, of c o u r s e , presents a p r o b l e m to t h e c o l l e c -
for of fossil brachiopods. It m e a n s that he or she m u s t m a k e a real effort to
obtain loose valves that have b e e n naturally c l e a n e d over t i m e so as to reveal
their internal features, h a i l i n g that, the c o l l e c t o r m u s t b r e a k , c u t , or g r i n d
s p e c i m e n s o p e n and c l e a n t h e m m e t i c u l o u s l y t o reveal t h e i n n e r truth.
S o m e o f the a r t i c u l a t e b r a c h i o p o d s m u s t h a v e s p e n t m u c h o f t h e i r
lives a n c h o r e d b y their p e d i c l e s t o s h e l l s , h a r d g r o u n d s , o r o t h e r solid o b -
jects o n t h e sea floor ( F i g u r e 8.9E). They w e r e n o t f r o z e n i n p o s i t i o n ,
t h o u g h , b e c a u s e t h e y a p p a r e n t l y had adjustor m u s c l e s that a l l o w e d t h e
individual a n i m a l t o m o v e its shell w i t h r e s p e c t t o t h e p e d i c l e . W e k n o w
this b e c a u s e present-day a r t i c u l a t e s h a v e s u c h adjustor m u s c l e s , a n d w h e r e
t h e s e adjusters a t t a c h to the insides of t h e v a l v e s , t h e r e are m u s c l e scars

Brachiopods 105
Figure 8.4. A, B. Plectorthis neglecta (James), MUGM uncatalogued. Fairview Formation, Hamilton
Co., Ohio. A. Brachial valve exterior. B. Pedicle valve exterior, both x 1.8. C, D. Dalmanella emac-
erata (Hall), MUGM 24362, Kope Formation, Hamilton Co., Ohio. C. Pedicle valve exterior. D. Brachial
valve exterior, both x 2.4. E, F. Sowerbyella rugosa (Meek), MUGM 24564, Kope Formation, Cincin-
nati, Ohio. E. Pedicle valve exterior. F. Brachial valve exterior, both x 2.2. G. Slab covered with Sower-
byella rugosa, Kope Formation. Courtesy of Paul E. Potter.

106 A Sea without Fish

Figure 8.5. A-D. Hebertella occidentalis (Hall), MUGM 11205, Waynesville Formation, Franklin Co.,
Indiana. A. Brachial valve exterior. B. Pedicle valve exterior. C. Pedicle valve interior, showing muscle
scars and triangular pedicle opening. D. Posterior view of articulated valves and triangular pedicle open-
ing, all x 1.2. E-H. Glyptorthis insculpta (Hall); E-F, MUGM 22450, Liberty Formation, Preble Co.,
Ohio. E. Brachial valve exterior. F. Pedicle valve exterior, x 2.8. G - H , MUGM 29458, Arnheim Forma-
tion, Butler Co., Ohio. G. Pedicle valve interior, showing muscle scars, triangular pedicle opening, and
encrusting cyclostome bryozoans, x 2.2. H. Brachial valve interior, showing muscle scars and cardinal
processes, x 1.8.

Brachiopods 107
Figure 8.6. A-C. Plaesiomys subquadrata (Hall), MUGM 22933 Liberty Formation, Preble Co.,
Ohio. A. Brachial valve exterior, inarticulate brachiopod Philhedra laelia on beak, x 2.1. B. Pedicle
valve exterior, x.2.1. C. Pedicle valve interior, showing muscle scars, x 2.4. D-F. Retrorsirostra carleyi
(Hall), MUGM 23037, Arnheim Formation, Butler Co., Ohio. D.
Brachial valve exterior. E. Pedicle valve exterior, note triangular pedicle opening and flanking inter-
area. F. Pedicle valve interior, showing muscle scars, x 2.

108 A Sea without Fish

Figure 8.7. Three of the many described species of Platystrophia. A-D. Platystrophia ponderosa Foer-
ste, MUGM 24060, Maysvillian, Campbell Co., Kentucky A. Pedicle valve exterior, x 7.5. B. Brachial
valve exterior, x 7.5. C. Pedicle valve interior, showing pedicle opening and deep muscle scar, x
1.0. D. Brachial valve interior, showing muscle scars, x 1.3. Early collectors of Cincinnatian fossils referred
to this large, robust brachiopod as the "double-headed Dutchman." E, F. Platystrophia laticosta
(Meek), MUGM 11315, Maysvillian, Cincinnati, Ohio. E. Brachial valve exterior. F. Pedicle valve exterior,
both x 7.5. G, H Platystrophia acutilirata (Conrad), MUGM 23360. G. Pedicle valve exte-
rior., H. Brachial valve exterior, Whitewater Formation, Preble Co., Ohio, both x 1.4.

Brachiopods 109
Figure 8.8. The many faces of Rafinesquina: Rafinesquina alternata (Conrad). A. University of Cincin-
nati collections, left, pedicle valve up, with encrusting edriasteroids and cyclostome bryozoans, right, bra-
chial valve up, with encrusting edrioasteroid Streptaster vorticellatus and crinoid locrinus subcrassus
wedged beneath lower left edge, Corryville Formation, Boone Co., Kentucky, scale in mm. B. Pedicle
valve interior, CMC IP 51111, showing muscle scars, scale in mm. C. Brachial valve interior, University of
Cincinnati collections, showing muscle scars and pair of cardinal processes along hinge at top, Corryville
Formation, Boone Co., Kentucky, size about same as in B. D. Rafinesquina pavement, with pedicle valves
up, Fairview Formation, Kenton Co., Kentucky. E. Shingled Rafinesquina bed, with valves perpendicular
to bedding, Fairview Formation, Kenton Co., Kentucky.

110 A Sea without Fish

Figure 8.9. A, B. Hiscobeccus capax (Conrad), MUGM 25490, Liberty Formation, Preble Co.,
Ohio. A. Brachial valve exterior. B. Posterior view of articulated valves showing small pedicle opening,
x 2.2. C, D. Rhynchotrema dentatum (Hall), MUGM 25933, Whitewater Formation, Preble Co.,
Ohio. C. Pedicle valve exterior. D. Anterior view of articulated valves showing pronounced sulcus, x
3.4. E. Zygospira modesta (Say), CMC IP 51112, Corryville Formation, Boone Co., Kentucky, attached in
life position to bryozoan Parvohallopora sp., x 1.9. F. Catazyga schuchertana (Ulrich), MUGM 7614,
Waynesville Formation, Jefferson Co., Indiana, brachial valve exterior, x 3.4.

Brachiopods in
Figure 8.10. Environmen-
tal distribution of bra-
chiopods in the Cincin-
natian Series. Shoreface
environments are equiva-
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
University Press. Re-
that o c c u r i n p a r t i c u l a r p o s i t i o n s . M a n y fossil b r a c h i o p o d s h a v e c o m p a -
printed with permission
of the publisher. rable m u s c l e scars.
O n t h e o t h e r h a n d , s o m e k i n d s o f fossil a r t i c u l a t e s h a v e n o p e d i c l e
f o r a m e n , t h e o p e n i n g t h r o u g h w h i c h the p e d i c l e e x t e n d s b e y o n d the h i n g e
a r e a o f t h e s h e l l . T h e r e has b e e n a g r e a t d e a l o f d i s c u s s i o n a s t o h o w t h e s e
c r e a t u r e s s u r v i v e d i n a r e a s w h e r e t h e sea floor w a s soft m u d .
B r a c h i o p o d s of g e n u s Rafinesquina ( f i g u r e 8.8) are, p e r h a p s , the m o s t
c o m m o n o f the larger a r t i c u l a t e s i n t h e rocks o f t h e C i n c i n n a t i area. T h e
overall s h a p e of t h e shell is d e s c r i b e d as " c o n c a v o - c o n v e x " ; the brachial valve
( a n a t o m i c a l l y dorsal) is c o n c a v e to the exterior, and the other valve (anatomi-
cally ventral) is c o n v e x to the outside. T h e w h o l e shell, t h e n , is saucer- or
b o w l - s h a p e d . T h e adult a n i m a l d o e s not s e e m t o h a v e h a d a p e d i c l e n o
p e d i c l e f o r a m e n . Hence, it m u s t h a v e b e e n free on the sea floor. But i m a g i n e
a s a u c e r - s h a p e d shell in a c u r r e n t ; all too easily, it w o u l d h a v e b e e n flipped
so that the c o n v e x side w a s u p p e r m o s t . The result w o u l d have b e e n that the
c o m m i s s u r e , the o p e n i n g b e t w e e n the valves, w o u l d have b e e n against the
sea floor. If that sea floor w e r e soft m u d , the a n i m a l w o u l d have had consider-
able difficulty g e n e r a t i n g sufficient c u r r e n t s with the cilia of its l o p h o p h o r e
so as to b r i n g l i f e - g i v i n g nutrients and oxygen b e t w e e n the valves. T h u s ,
t h e r e s e e m s to be a c o n f l i c t b e t w e e n h y d r o d y n a m i c s a n d biology.
The bleak p i c t u r e of the b r a c h i o p o d . u p s i d e d o w n with its o p e n i n g in
t h e m u d , may be m i s l e a d i n g . It portrays t h e a n i m a l as an i m m o b i l e l u m p
u n a b l e t o r i g h t itself. T r u e , t h e r e w a s n o p e d i c l e o n w h i c h the a n i m a l , us-
i n g a d j u s t e r m u s c l e s , c o u l d twist a n d t u r n itself b a c k into a v i a b l e p o s i t i o n .
B u t , w h a t if the a n i m a l w e r e less like an i n a n i m a t e s a u c e r a n d m o r e like a
l i v i n g s c a l l o p of t o d a y ? A s c a l l o p is a k i n d of b i v a l v e d m o l l u s c . T h e "scal-

112 A Sea without Fish

l o p " y o u enjoy in y o u r favorite s e a f o o d r e s t a u r a n t is an a d d u c t o r m u s c l e of
one of those p e l e c y p o d s . The adductor of a scallop is powerful e n o u g h , in
life, to s n a p the a n i m a l ' s valves t o g e t h e r so swiftly that t h e c r e a t u r e c a n be
lifted a b o v e the sea floor. S o m e s c a l l o p s c a n e v e n s w i m for s o m e d i s t a n c e ,
a l t h o u g h rather jerkily a n d i n d e c i d e d l y i r r e g u l a r trajectories.
D a t t i l o (2004) has f o u n d e v i d e n c e that o n e C i n c i n n a t i a n b r a c h i o p o d
with a c o n c a v o - c o n v e x shell, Sowerbyella, w a s c a p a b l e of e s c a p i n g f r o m
burial b e n e a t h s e d i m e n t s stirred up by storms, p r e s u m a b l e by s n a p p i n g its
valves. Individuals of Rafinesquina m a y h a v e h a d s i m i l a r c a p a b i l i t i e s , b e -
c a u s e c o n v e x - u p s p e c i m e n s are f o u n d w i t h a moat-like furrow a r o u n d the
c o m m i s s u r e that f o r m e d w h i l e the b r a c h i o p o d w a s alive ( M e y e r 2006).
T h e s e recent findings suggest that these b r a c h i o p o d s w i t h o u t p e d i c l e a t t a c h -
m e n t m i g h t have led m u c h m o r e a c t i v e lives than previously r e a l i z e d .

D e s p i t e the h i g h diversity o f t y p e - C i n c i n n a t i a n b r a c h i o p o d s , t h e d i s t r i b u - Distribution of

tion o f s p e c i e s i s not u n i f o r m t h r o u g h o u t t h e s t r a t i g r a p h i c s u c c e s s i o n . Type Cincinnatian
There are distinct a s s o c i a t i o n s o f s p e c i e s a n d shell t y p e s t h a t c h a r a c t e r i z e
Brachiopods in
different stratigraphic intervals a n d e v e n i n d i v i d u a l b e d s . R e c e n t r e s e a r c h
Time and Space
In Steven Holland, in c o l l a b o r a t i o n with A r n o l d M i l l e r , D a v i d M e y e r , a n d
B e n j a m i n D a t t i l o ( H o l l a n d e t al. 2 0 0 1 ) s h o w e d that t h e relative a b u n d a n c e
o f b r a c h i o p o d s and o t h e r fossils c h a n g e s w i t h i n t h e K o p e F o r m a t i o n , a unit
g e n e r a l l y regarded as having a u n i f o r m shaley lithology. In the l o w e r K o p e ,
fossil a s s e m b l a g e s a s f o u n d o n c h a r a c t e r i s t i c l i m e s t o n e b e d d i n g s u r f a c e s
are d o m i n a t e d by the s m a l l , t h i n - s h e l l e d Sowerbyella, a l o n g w i t h b r a n c h -
ing b r y o z o a n s , s m a l l , s l e n d e r c r i n o i d s like Cincinnaticrinus a n d Ectenocri-
mis, a n d t h e trilobite Cryptolithus. H i g h e r in t h e K o p e , a n o t h e r t h i n -
shelled b u t larger b r a c h i o p o d , Dalmanella, b e c o m e s m o r e a b u n d a n t . I n
the highest sections of the Kope, the large thin-shelled brachiopods
Rafinesquina and Strophomena a n d t h e l a r g e , rather t h i c k - s h e l l e d Plat-
ystrophia b e c o m e the d o m i n a n t b r a c h i o p o d s . I n c o n j u n c t i o n w i t h c h a n g e s
in the lithology, b e d d i n g , a n d o t h e r c h a r a c t e r i s t i c fossils, Holland. M i l l e r ,
M e y e r , a n d D a t t i l o (2001) i n t e r p r e t e d the c h a n g e s i n t h e K o p e F o r m a t i o n
as a p a l e o b a t h y m e t r i c g r a d i e n t r e f l e c t i n g t r a n s i t i o n f r o m d e e p e r to s h a l -
l o w e r water.
T h r o u g h o u t all t y p e - C i n c i n n a t i a n d e p o s i t i o n a l s e q u e n c e s t h e n a t u r e o f
the b r a c h i o p o d a s s e m b l a g e s provides o n e o f t h e m o s t reliable a n d a b u n d a n t
indicators of p a l e o e n v i r o n m e n t a l c o n d i t i o n s s u c h as d e p t h a n d level of water
m o v e m e n t e n e r g y (see c h a p t e r 15). The relationship b e t w e e n b r a c h i o p o d
m o r p h o l o g y and d e p t h reflects b o t h t h e t y p e o f s u b s t r a t u m a n d t h e level o f
water m o v e m e n t energy. In the d e e p e r water, m u d d y e n v i r o n m e n t s , brachiopods with s m a l l , t h i n , flat shells a c t e d like s n o w s h o e s
In shallower water, larger, c o n c a v o - c o n v e x brachiopods like Rafinesquina
and Hebertella were better a d a p t e d to stronger w a v e energy. O t h e r larger
brachiopods like Platystrophia, with thicker shells a n d w e l l - d e v e l o p e d plica-
tions (ribs radiating from the b e a k ) , c h a r a c t e r i z e s o m e o f t h e s h a l l o w e s t ,
highest w a v e - d i s t u r b e d e n v i r o n m e n t s . F i g u r e 8.10 s h o w s the e n v i r o n m e n t a l
distribution o f other b r a c h i o p o d s w i t h i n the t y p e - C i n c i n n a t i a n .

Brachiopods 113
 E v e n on a s m a l l e r s c a l e , b r a c h i o p o d s reveal s o m e very basic a s p e c t s of
life o n t h e L a t e O r d o v i c i a n sea f l o o r . V e r y d e n s e l y p o p u l a t e d l i m e s t o n e
beds, featuring brachiopods like Rafinesquina, Strophomena, and Dal-
manella, are v e r y c o m m o n t h r o u g h o u t t h e t y p e - C i n c i n n a t i a n a n d are
c a l l e d s h e l l p a v e m e n t s (also c a l l e d s h i n g l e d b e d s a s n o t e d above). I n s u c h
shell p a v e m e n t s , t h e b r a c h i o p o d s a r e u s u a l l y p r e s e r v e d w i t h c o n v e x valves
upward, s o m e t i m e s covering the entire bed surface (Figures 8.4G, 8.8D,
8.8E). S h e l l p a v e m e n t s c a n be as t h i n as a s i n g l e layer of shells, or t h i c k e r ,
w i t h t h e e n t i r e t h i c k n e s s u p t o a f e w tens o f c e n t i m e t e r s c o n s i s t i n g o f
s t a c k e d b r a c h i o p o d s . In s o m e c a s e s , t h e valves are vertical or tilted at vari-
o u s a n g l e s a n d p a c k e d c l o s e l y t o g e t h e r i n a n e d g e w i s e shell b e d . T h e e d g e -
w i s e shell p a v e m e n t s are g o o d e v i d e n c e o f w a t e r m o v e m e n t i n the form o f
w a v e o s c i l l a t i o n s b e c a u s e t h e r e a r e p r e s e n t - d a y e x a m p l e s o f e d g e w i s e shell
b e d s a n d s h a l e f r a g m e n t s f o r m e d i n very shallow water b y w a v e oscillations.
II the hingelines or b e a k s of the b r a c h i o p o d s w e r e a l w a y s d i r e c t e d d o w n -
ward in an e d g e w i s e b e d , it m i g h t be possible that t h e d e n s e l y p a c k e d shells
a c t u a l l y h a d lived in a m a n n e r s i m i l a r to an oyster b e d . However, analysis
o f v a l v e o r i e n t a t i o n w i t h i n e d g e w i s e shell b e d s s h o w s that valves d o not
show such a pattern and even can be predominantly hingeline-upward
( S e i l a c h e r 1 9 7 3 , pers. c o m m . ) .
T h e r e has b e e n c o n s i d e r a b l e d e b a t e as to how shell p a v e m e n t s c o u l d
h a v e f o r m e d . M a n y w o r k e r s felt that the t h i n , c o n c a v o - c o n v e x shells o f the
characteristic brachiopods like Rafinesquina and Strophomena initially
l i v e d o n a soft, m u d d y sea f l o o r , w i t h t h e c o n v e x v a l v e d o w n w a r d . D u r i n g
a storm, the fine-grained m u d s c o u l d h a v e b e e n s w e p t away, l e a v i n g a
c o n c e n t r a t i o n of shells ( c a l l e d a l a g d e p o s i t ) to form the shell p a v e m e n t .
Possibly t h e shells w e r e e v e n c a r r i e d s o m e d i s t a n c e b y storm c u r r e n t s t o b e
d e p o s i t e d later. I n c a s e s w h e r e p a v e m e n t s f o r m e d g e w i s e b e d s , storms
c o u l d v e r y w e l l h a v e b e e n i n v o l v e d , b u t n o t all p a v e m e n t s are e d g e w i s e .
It is also p o s s i b l e that shell p a v e m e n t s a c c u m u l a t e d by a b u n d a n t pro-
d u c t i o n of shells of a s i n g l e s p e c i e s over s o m e t i m e span in o n e p l a c e . In
o n e shell p a v e m e n t f r o m t h e C o r r y v i l l e M e m b e r o f the G r a n t L a k e F o r m a -
tion in n o r t h e r n K e n t u c k y , m o s t l y c o n v e x - u p w a r d shells of Rafinesquina
form a k i n d of i m b r i c a t e d or s h i n g l e d b e d , but the s p a c e s b e t w e e n the
shells a r e m u d - f i l l e d , f o r m i n g a t y p e o f l i m e s t o n e k n o w n a s p a c k s t o n e
( M e y e r 1990). I f t h e m u d h a d b e e n r e m o v e d b y a s t o r m , t h e r e m a i n i n g
shell b e d c o u l d h a v e f o r m e d a g r a i n s t o n e . S h e l l s i n t h e u p p e r s u r f a c e o f
the b e d a r e a m i x t u r e o f a r t i c u l a t e d shells w i t h g o o d p r e s e r v a t i o n o f f i n e
s u r f a c e features a n d d i s a r t i c u l a t e d shells that are a b r a d e d a n d b r o k e n . B o t h
a b r a d e d a n d u n a b r a d e d shells are e n c r u s t e d w i t h b r y o z o a n s a n d edrioast-
eroid e c h i n o d e r m s . S o m e a r t i c u l a t e d b r a c h i o p o d s h a v e the m o a t - l i k e fea-
ture m e n t i o n e d a b o v e that s u g g e s t s a c t i v i t y o f the l i v i n g b r a c h i o p o d . A l l
t h e s e f e a t u r e s a r e e v i d e n c e that t h e shell b e d a c c u m u l a t e d g r a d u a l l y w i t h -
out significant transportation. Ultimately the entire bed was smothered by
an influx of m u d , probably p r o d u c e d by a s t o r m .
C h a r a c t e r i s t i c a l l y , a stratigraphic s e c t i o n w i t h the t y p e of shell pave-
m e n t d e s c r i b e d a b o v e will c o n t a i n repetitions o f thin s h e l l - p a v e m e n t s s m o t h -
e r e d b y s h a l e s . Harris a n d M a r t i n (1979) d e s c r i b e d this pattern i n t h e

114 A Sea without Fish

W a y n e s v i l l e F o r m a t i o n as a form of p a l e o e c o l o g i c s u c c e s s i o n (see F i g u r e
4.S). In present-day settings, e c o l o g i c s u c c e s s i o n o c c u r s w h e n o n e a s s e m -
blage of a n i m a l s or plants alters the habitat IN s u c h a way that o t h e r species
c a n replace the s o - c a l l e d p i o n e e r s p e c i e s . Harris a n d M a r t i n (1979) sug-
gested that thin-shelled brachiopods w e r e p i o n e e r s p e c i e s that first c o l o n i z e d
soft m u d d y p a t c h e s of the sea floor a n d provided a p a v e m e n t on w h i c h e n -
c r u s t i n g a n i m a l s like b r y o z o a n s and i n a r t i c u l a t e b r a c h i o p o d s c o u l d settle.
E v e n t u a l l y other species c o u l d take a d v a n t a g e o f the shell p a v e m e n t a n d
thickets of b r y o z o a n s , so that the diversity of the a s s e m b l a g e i n c r e a s e d u p -
ward from the b o t t o m of a p a v e m e n t b e d . S t o r m s frequently s m o t h e r e d t h e
shelly patches with m u d , thus i n t e r r u p t i n g the s u c c e s s i o n until b r a c h i o p o d
larvae o n c e again c o u l d c o l o n i z e the barren m u d s . s o m e p a l e o e c o l o g i s t s
have questioned w h e t h e r p a l e o e c o l o g i c s u c c e s s i o n c o m p a r a b l e t o present-
day succession c a n be d e t e c t e d in the fossil record b e c a u s e m o s t stratigraphic
c h a n g e s in fossil a s s e m b l a g e s represent a m u c h l o n g e r t i m e scale t h a n the
scale of years to d e c a d e s over w h i c h present-day s u c c e s s i o n o c c u r s . A l t h o u g h
we still do not know how m u c h t i m e was r e q u i r e d for the f o r m a t i o n of char-
acteristic, thin t y p e - C i n c i n n a t i a n shell p a v e m e n t s , it is possible that they
formed over a short t i m e scale. It also s e e m s c o r r e c t to v i e w the brachiopods
as h a v i n g a pivotal role in p r o v i d i n g a hard s u b s t r a t u m o n t o w h i c h e n c r u s t -
ing a n i m a l s c o u l d settle, thus altering the habitat in the m a n n e r of s u c c e s -
sional pioneers. C l e a r l y , s u c c e s s i o n at the scale of i n d i v i d u a l shell p a v e m e n t s
was an important act on the stage of the C i n c i n n a t i a n sea floor, a n d brachiopods played a major role in that e v o l u t i o n a r y play.

Brachiopods 115
116 A Sea without Fish
 9
MOLLUSCS: HARD, BUT
WITH A SOFT CENTER

Everyone k n o w s m o l l u s c s t h e o h - s o - f a m i l i a r snails a n d s l u g s , t h e c l a m s ,
m u s s e l s , s c a l l o p s , a n d oysters, t h e o c t o p u s , a n d t h e s q u i d . B u t the m o l l u s c
story is not a s i m p l e o n e . T h e r e are m o r e k i n d s of m o l l u s c s t h a n of a n y
other g r o u p o f a n i m a l s , save t h e a r t h r o p o d s . S o w h a t l i n k s all the m o l l u s c s
together?
T h e word " m o l l u s c " is derived from the Latin word " m o l l u s c u s , "
m e a n i n g "soft." This refers to t h e fact that e v e r y m o l l u s c h a s a soft, fleshy
b o d y . But that, o f c o u r s e , i s not t h e i m a g e c o n j u r e d u p i n t h e m i n d ' s e y e
a t the m e n t i o n o f snails, c l a m s , a n d oysters. I n m o s t o f t h e m o l l u s c s , the
soft parts are e n c l o s e d w i t h i n a h a r d s h e l l . A n d it is on t h e basis of differ-
e n c e s i n the shells that t h e m o l l u s c s o f t h e t y p e - C i n c i n n a t i a n are d i f f e r e n -
tiated from o n e a n o t h e r .
O n e m i g h t b e t e m p t e d t o sort t h e s h e l l e d m o l l u s c s into t h r e e g r o u p s ,
a t least with respect t o their s h e l l s . T h e a n i m a l s o f o n e g r o u p h a v e b a s i c a l l y
a single shell; take, for e x a m p l e , t h e c o i l e d c o n e of m o s t snails or that of
the pearly n a u t i l u s . In c o n t r a s t to t h e s e u n i v a l v e d m o l l u s c s are t h o s e in
w h i c h the soft parts a r e e n c l o s e d b e t w e e n t w o " s h e l l s " (strictly s p e a k i n g ,
e a c h of the t w o is c a l l e d a " v a l v e " ) . The b i v a l v e d m o l l u s c s that l e a p m o s t
Figure 9.1. Gastropod
readily to m i n d are the c l a m s , m u s s e l s , oysters, a n d t h e i r k i n . In a third
mollusc, showing internal
g r o u p , and a relatively s m a l l o n e at that, t h e shell c o n s i s t s of a n u m b e r of
features. Drawing by
plates a r r a n g e d so that the a n i m a l , at first g l a n c e , a p p e a r s to be s e g m e n t e d . Kevina Vulinec.
(In this c a s e , l o o k s a r e d e c e i v i n g , b e c a u s e , u n l i k e t h e a n n e l i d w o r m s a n d
the a r t h r o p o d s , m o l l u s c s a r e not truly s e g m e n t e d . ) The p r e s e n t - d a y c h i t o n s
e x e m p l i f y the p o l y p l a c o p h o r a n g r o u p (literally, " m a n y plate b e a r i n g " ) .
Regardless of w h e t h e r the a n i m a l h a s a shell t h a t is a s i n g l e v a l v e or
o n e c o n s i s t i n g o f t w o valves o r m a n y p l a t e s , t h e m a t e r i a l o f t h e shell i s
p r i m a r i l y c a l c i u m c a r b o n a t e i n the m i n e r a l f o r m o f e i t h e r c a l c i t e o r a r a g o -
nite. B e c a u s e a r a g o n i t e is less stable t h a n c a l c i t e , a r a g o n i t i c shells u s u a l l y
dissolve after d e a t h , l e a v i n g just e x t e r n a l a n d i n t e r n a l m o l d s t o r e c o r d
w h e r e the shell o n c e h a d b e e n (see c h a p t e r 5). This, of c o u r s e , c a n g r e a t l y
affect w h a t we find as fossils.
As a m o l l u s c progresses t h r o u g h its life, it g e n e r a l l y a d d s shell m a t e r i a l
at the periphery of its shell or of e a c h valve of its shell. Thus, the life history
of that shell or valve is r e c o r d e d in a series of c o n c e n t r i c g r o w t h - l i n e s visible
on the exterior of the shell. This a l l o w s us to d e c i p h e r the c h a n g e s in t h e size
and shape that the shell or valve went t h r o u g h d u r i n g the life of t h e indi-
vidual a n i m a l . ( T h i s stands in s t r o n g contrast to a n i m a l s that s h e d their
exoskeletons as they grow. An individual adult trilobite, for e x a m p l e , d o e s n o t
present to e v e n the careful o b s e r v e r its life history in its hard parts.)

117
 O f c o u r s e , e v e n i n m o l l u s c s that h a v e shells, the hard parts are m e r e l y
a part of the w h o l e a n i m a l . In g e n e r a l , t h e b o d y of a m o l l u s c i n c o r p o r a t e s
f i v e o f w h a t c o m m o n l y are c a l l e d b o d y r e g i o n s : t h e h e a d , the foot, the
visceral mass, the m a n t l e - c o m p l e x , and the gills (technically termed
c t e n i d i a , f r o m their c o m b - l i k e shape). The m a n t l e , the s h e l l , a n d t h e m a n -
tle-cavity together c o m p r i s e the m a n t l e - c o m p l e x . The m a n t l e is a sheet of
tissue that h a n g s d o w n o n e a c h side o f t h e b u l k o f the a n i m a l o r o t h e r w i s e
e n c l o s e s it. The s h e l l , if p r e s e n t , is a t t a c h e d to the o u t s i d e of the m a n t l e
and is s e c r e t e d by it. To the inside of the m a n t l e is the m a n t l e - c a v i t y , in
w h i c h a r e l o c a t e d t h e g i l l s , i f t h e y are p r e s e n t .
T h e last p h r a s e o f t h e p r e v i o u s p a r a g r a p h i s a n i m p o r t a n t tip-off. N o t
all k i n d s o f m o l l u s c s h a v e all f i v e b o d y r e g i o n s d e v e l o p e d t o t h e s a m e ex-
tent. S n a i l s , for e x a m p l e , e a c h h a v e a n o b v i o u s h e a d , w h e r e a s c l a m s d o not.
S q u i d s h a v e w e l l - d e v e l o p e d gills; h o w e v e r , in terrestrial snails, there are no
c t e n i d i a , a n d t h e m a n t l e - c a v i t y serves, in e f f e c t , as a l u n g .
It is o n l y fair to a d m i t that t h e r e is so m u c h m o r p h o l o g i c a n d a n a t o m i -
cal v a r i a t i o n a m o n g t h e m o l l u s c s as a w h o l e that it is difficult to p o i n t to
a n y trait that o c c u r s in all m o l l u s c s . S o m e h a v e shells, a n d s o m e do not. In
m o s t , the shell is e x t e r n a l , but in some it is i n t e r n a l . S o m e h a v e g i l l s , a n d
s o m e d o not. S o m e h a v e h e a d s , a n d s o m e d o not. A n d s o o n .
R e g a r d l e s s o f w h e t h e r o n e i s c o n v i n c e d that form follows f u n c t i o n , o r
v i c e versa, the t r e m e n d o u s m o r p h o l o g i c a l s p e c t r u m e x h i b i t e d b y t h e m o l -
luscs as a w h o l e is c o i n c i d e n t w i t h t r e m e n d o u s e c o l o g i c a l and b e h a v i o r a l
s p e c t r a . S a v e for the fact that m o l l u s c s h a v e not m a s t e r e d in-air flight,
v i r t u a l l y a n y terrestrial or a q u a t i c e n v i r o n m e n t on E a r t h will h a v e a repre-
sentative suite o f m o l l u s c s .
This t r e m e n d o u s array of s i z e s , s h a p e s , b e h a v i o r s , and w a y s of life
a m o n g t h e m o l l u s c s i s t h e result o f m i l l i o n s o f c e n t u r i e s o f o r g a n i c e v o l u -
tion. I n c l u d e d in this a l m o s t i n c r e d i b l e diversity of a n i m a l s are, to b r a g a
bit, n o t o n l y t h e largest o f all i n v e r t e b r a t e s , but a l s o t h e m o s t i n t e l l i g e n t o f
all invertebrates ( b o t h , it h a p p e n s , b e i n g c e p h a l o p o d s ) . M o r e o v e r , t h e larg-
est i n d i v i d u a l n e r v e c e l l s are said to o c c u r in m o l l u s c s ( a g a i n , c e p h a l o p o d s
are t h e c h a m p i o n s ) .

Who's W
Whhoo aammoonngg In t h e i n t r o d u c t i o n to this chapter, m o l l u s c s w e r e separated into u n i v a l v e d
tt h
h ee Molluscs
Molluscs m o l l u s c s , bivalved m o l l u s c s , a n d polyplacophoran m o l l u s c s . A l t h o u g h per-
haps c o n v e n i e n t t o i n t r o d u c e the n o t i o n o f m o r p h o l o g i c a l variation a m o n g
the m o l l u s c s as a phylum, those three " g r o u p s " are a gross over-simplification.
M a l a c o l o g i s t s , t h o s e w h o study m o l l u s c s , use the s h a p e of the shell to sort
m o l l u s c s into true b i o l o g i c a l g r o u p s i n t o g r o u p s w h o s e m e m b e r s are e v o l u -
t i o n a r y related t o o n e a n o t h e r . H o w e v e r , u n l i k e c o n c h o l o g i s t s , those w h o
study shells, m a l a c o l o g i s t s u s e all sorts of traits. In a d d i t i o n to t h o s e of the
shell. Of c o u r s e , in most fossils, o n l y the hard parts are preserved. N o n e t h e -
less, it is the goal of the p a l e o n t o l o g i s t to p u t the l i v i n g a n i m a l b a c k into its
shell a n d to r e c o g n i z e the real, b i o l o g i c a l g r o u p s of fossils.
I n the O r d o v i c i a n r o c k s o f t h e C i n c i n n a t i r e g i o n , f o s s i l s o f the follow-
i n g b i o l o g i c a l g r o u p s of phylum M o l l u s c a are f o u n d (recall that a p h y l u m

118 A Sea without Fish

consists o f a n u m b e r o f s m a l l e r g r o u p s o f o r g a n i s m s c a l l e d classes): class
C e p h a l o p o d a , class G a s t r o p o d a , class M o n o p l a c o p h o r a , class P e l e c y p o d a ,
class R o s t r o c o n c h i a , class P o l y p l a c o p h o r a , a n d class S c a p h o p o d a . T h e r e
are s o m e g r o u p s o f m o l l u s c s o f w h i c h n o s p e c i m e n s are k n o w n f r o m t h e
local rocks. For e x a m p l e , t h e class A p l a c o p h o r a i n c l u d e s c e r t a i n p r e s e n t -
day a n i m a l s that a r e d e v o i d o f p r e s e r v a b l e h a r d parts; h e n c e , their n a m e ,
w h i c h m e a n s " n o plate b e a r i n g . "

S n a i l s , w i t h their c h a r a c t e r i s t i c c o i l e d s h e l l s , are p r o b a b l y t h e e a s i e s t to Class G a s t r o p o d a

Class
r e c o g n i z e o f the m o l l u s c s in t h e r o c k s o f t h e C i n c i n n a t i r e g i o n . S n a i l s are T hhee Snails
T Snails
very c o m m o n i n m a i n e n v i r o n m e n t s o f today's w o r l d , a l o n g w i t h their rela-
tives, t h e s o - c a l l e d slugs a n d sea-slugs. ( A n i m a l s o f t h e latter t w o g r o u p s
either h a v e no shell at all, or a s m a l l , i n t e r n a l o n e ; t h u s , they are u n l i k e l y
to he p r e s e r v e d as fossils.)
Snails are the q u i n t e s s e n t i a l u n i v a l v e d m o l l u s c ( F i g u r e 9.1). Each snail
has a single p r o m i n e n t v a l v e . In m o s t s n a i l s , this is a l o n g , n a r r o w , c o n i c a l
t u b e that is c o i l e d off to o n e side, so that it r e s e m b l e s a screw. Of c o u r s e ,
b y n o w , y o u h a v e c o m e t o e x p e c t that t h e story i s b o u n d t o b e far m o r e
complicated.
First, in s o m e snails, the shell is so flared o p e n t h a t it r e s e m b l e s a c a p
or a s h i e l d , rather than a screw; take, for e x a m p l e , t h e l i m p e t s a n d t h e a b a -
lones of today. T h e n , there are the snails in w h i c h t h e c o i l i n g is n o t to t h e
side; rather, e a c h w h o r l lies d i r e c t l y in l i n e w i t h t h e o n e n e x t to it, so t h a t
the c o i l is m o r e like a g a r d e n h o s e c o i l e d flat on t h e g r o u n d (this is c a l l e d
planispiral c o i l i n g ) .
A l t h o u g h basically u n i v a l v e d , s o m e snails h a v e a h a r d s t r u c t u r e that
serves t o b l o c k t h e a p e r t u r e o f t h e shell w h e n t h e a n i m a l w i t h d r a w s for
protection. D e p e n d i n g on the kind of snail, this o p e r c u l u m , as it is called,
m a y b e m a d e o f c a l c i u m c a r b o n a t e , like t h e s h e l l , o r o f a s u b s t a n c e c a l l e d
c o n c h i o l i n , w h i c h i s rather like t h e c h i t i n o f a n insect's e x o s k e l e t o n . N o t
all snails h a v e o p e r c u l a , a n d n o o p e r c u l a t e g a s t r o p o d s h a v e b e e n r e p o r t e d
from the rocks o f t h e C i n c i n n a t i r e g i o n .
The n a m e " g a s t r o p o d " literally m e a n s " s t o m a c h foot," a n d t h e b o d y
region of a g a s t r o p o d c a l l e d t h e " f o o t " g e n e r a l l y is a b r o a d , flat, c r e e p i n g
o r g a n ( F i g u r e 9.1). T h e l i v i n g a n i m a l has a p r o m i n e n t h e a d , g e n e r a l l y
c o m p l e t e with o n e o r t w o pairs o f stalks o r t e n t a c l e s .
Y o u may well have e n c o u n t e r e d a snail or a slug in y o u r g a r d e n , but m o s t
gastropods are aquatic. The snail or s l u g in y o u r g a r d e n probably was (or h a d
b e e n ) d e v o u r i n g y o u r plants; many g a s t r o p o d s are herbivores. G a s t r o p o d s
have a structure associated with the m o u t h that looks rather like a carpenter's
rasp. T h e radula, as it is c a l l e d , works like a rasp, too; t h e a n i m a l p r o t r u d e s
the radula from its m o u t h and scrapes bits of plant matter into the m o u t h .
But not all snails are h e r b i v o r e s . I n d e e d , s o m e subsist on t h e flesh of
others. An e s p e c i a l l y s t r i k i n g e x a m p l e f r o m t h e l o c a l r o c k s is p r o v i d e d by
b r a c h i o p o d shells or o t h e r shells that e a c h h a v e a tidy, c i r c u l a r h o l e . In s u c h
c a s e s , a snail u s e d its r a d u l a to drill a h o l e in t h e shell of a n o t h e r a n i m a l -
object: dinner.

Molluscs 119
Figure 9.2. Cincinnatian M o s t o f t h e snail fossils i n t h e r o c k s o f the C i n c i n n a t i r e g i o n consist
monoplacophorans and a of m o l d s . T h e a c t u a l shell m a t t e r has b e e n dissolved away, a n d all that is
bellerophontid gastro- left is t h e s e d i m e n t that o r i g i n a l l y filled or that s u r r o u n d e d t h e b u r i e d
pod. A. Archinacella
s h e l l , or b o t h . E x c e p t i o n s to this g e n e r a l i t y are t h e snails of the g e n u s Cy-
area Wahlman, USNM
clonema; h e r e , shell m a t t e r , n o t u n c o m m o n l y , is present.
40615, a monoplacoph-
oran, Waynesville
Formation, Waynesville, Gastropods of the type-Cincinnatian
Ohio, a, dorsal, b, lateral,
c, anterior, all x 1.6. From S n a i l s ( F i g u r e s 9.2, 9.3) are very c o m m o n fossils t h r o u g h o u t t h e C i n c i n -

Wahlman (1992, plate 3, n a t i a n Series in its t y p e a r e a , b u t , b e c a u s e t h e y g e n e r a l l y are p r e s e r v e d o n l y

figures 7, 9, 10). B. Hel- a s i n t e r n a l m o l d s , then- c a n b e o v e r l o o k e d , a n d p r e c i s e i d e n t i f i c a t i o n c a n
cionopsis striata Ulrich be difficult. N e v e r t h e l e s s , s n a i l s u n d o u b t e d l y p l a c e d an i m p o r t a n t role in
and Scofield, USNM the e c o l o g y of the C i n c i n n a t i a n sea, especially in some e n v i r o n m e n t s
45827, a monoplacoph- w h e r e t h e y w e r e very a b u n d a n t . H o l l a n d ' s c o m p i l a t i o n o f C i n c i n n a t i a n
oran, Richmondian, Mar-
fossils lists sixty-three s p e c i e s of snails in t w e n t y - t h r e e g e n e r a as o c c u r r i n g
ion Co., Kentucky, a, dor-
a b o v e t h e b a s e o f t h e K o p e F o r m a t i o n ( H o l l a n d 2005). O f t h e s e , s i x t e e n
sal, b, oblique-lateral, c,
s p e c i e s i n n i n e g e n e r a b e l o n g t o the p l a n i s p i r a l l y c o i l e d b e l l e r o p h o n t i d s
lateral, all x 1.7. From
that w e r e revised t a x o n o m i c a l l y b y W a h l m a n (1992). o f the o t h e r f o u r t e e n
Wahlman (1992, plate 2,
figures 1, 2, 3). C. Cyr- g e n e r a a n d f o r t y - s e v e n s p e c i e s , the g e n u s Cyclonema a c c o u n t s for e l e v e n
tolites ornatus Conrad, s p e c i e s , b a s e d o n t h e 1970 s t u d y b y T h o m p s o n (1970). S p e c i m e n s o f a n
USNM 265906, compos- a d d i t i o n a l p l a t y c e r a t i d , w h i c h b e l o n g in Naticonema, o c c u r in the t y p e -
ite mold of a monopla- C i n c i n n a t i a n , but t h e s e h a v e not b e e n d e s c r i b e d in the scientific literature
cophoran, Corryville For- ( F e l t o n , pers. c o m m . ; B o w s h e r 1955, plate 1). T h e r e m a i n i n g t h i r t e e n g e n -
mation, Cincinnati, Ohio,
era, i n c l u d i n g thirty-six s p e c i e s , h a v e not b e e n t h o r o u g h l y revised i n r e c e n t
a, lateral, b, dorsal, x 1.7.
y e a r s , a n d , c o n s e q u e n t l y , a c o n c l u s i v e s t a t e m e n t a b o u t the total t a x o n o m i c
From Wahlman (1992,
diversity o f C i n c i n n a t i a n g a s t r o p o d s w o u l d b e p r e m a t u r e . S o m e o f the
plate 6, figures 6,
m o s t c o m m o n g a s t r o p o d s are illustrated i n F i g u r e 9.3.
7). D. Cyrtolites orna-
tus, MUGM 18120, two S p e c i m e n s of Cyclonema are by far the best-preserved a n d m o s t easily
individuals somehow pre- r e c o g n i z e d C i n c i n n a t i a n s n a i l s b e c a u s e t h e i r shells w e r e c a l c i t i c rather
served with opposing t h a n a r a g o n i t i c in c o m p o s i t i o n ( F i g u r e s 9 . 3 A - F ) . I n d i v i d u a l s of Cyclonema
apertures, each encrusted are f o u n d i n e v e r y f o r m a t i o n o f t h e t y p e - C i n c i n n a t i a n . S o m e s p e c i e s , s u c h
by bryozoan Leptotrypa as C. humerosum, r a n g e f r o m t h e F a i r v i e w t h r o u g h the W a y n e s v i l l e For-
clavacoidea. Cincinna-
m a t i o n s ( T h o m p s o n 1970), b u t o t h e r s are m o r e restricted w i t h i n that r a n g e
tian. x 1.7. E. Sinuites
(Felton, pers. c o m m . ) T h o m p s o n d i f f e r e n t i a t e d s p e c i e s o f Cyclonema o n
cancellatus (Hall), CMC
t h e basis o f shell s h a p e , n a t u r e o f t h e a p e r t u r e , a n d o r n a m e n t (for e x a m p l e ,
IP 44304, a calcitic speci-
up to t h r e e sets of spiral ridges). G r o w t h - l i n e s p a r a l l e l to t h e a p e r t u r a l
men of a monoplacoph-
oran. Cincinnatian, Cincin- m a r g i n c u t across t h e spiral o r n a m e n t to c r e a t e a r e t i c u l a t e p a t t e r n . A

nati, Ohio, a, anterior, b, q u a n t i t a t i v e s t u d y o f v a r i a t i o n i n shell form a n d o r n a m e n t w i t h i n and

anterolateral, x 1.5. From a m o n g t h e m a n y s p e c i e s a n d f o r m s o f Cyclonema w o u l d h e l p t o clarify the
Wahlman (1992, plate 11, r e c o g n i t i o n of s p e c i e s a n d to d e t e r m i n e how v a r i a t i o n is related to d e p o s i -
figures 7, 9). F. Salpin- tional e n v i r o n m e n t .
gostoma richmondensis
S p e c i m e n s of Cyclonema are s o m e t i m e s f o u n d a t t a c h e d to the u p p e r
Ulrich, USNM 45983, an
s u r f a c e o r t e g m e n o f c r i n o i d c a l y c e s . This a s s o c i a t i o n b e t w e e n g a s t r o p o d s
internal mold of a bellero-
and crinoids is o n e of the best-known cases of interaction between species
phontid gastropod.
in t h e fossil r e c o r d . In t h e t y p e - C i n c i n n a t i a n , s p e c i m e n s of b o t h Cy-
Whitewater Formation,
Richmond, Indiana, x 1.4. clonema a n d Naticonema are f o u n d a t t a c h e d to the t e g m e n s of c r i n o i d s ,

From Wahlman, 1992, u s u a l l y of Glyptocrinus a n d Pycnocrinus (see c h a p t e r 12). B e c a u s e , in m o s t

plate 26, figure 2. s p e c i m e n s , t h e snail i s p o s i t i o n e d d i r e c t l y o v e r the a n a l o p e n i n g o f the

120 A Sea without Fish

c r i n o i d , m o s t workers h a v e c o n c l u d e d that the snail led u p o n the partly
digested feces of the c r i n o i d . B o w s h e r i n t e r p r e t e d this a s s o c i a t i o n as o n e of
c o p r o p h a g y a n d illustrated e x a m p l e s from t h e C i n c i n n a t i a n ( B o w s h e r
1955). S i m i l a r a s s o c i a t i o n s b e t w e e n related snails a n d c r i n o i d s are f o u n d
from y o u n g e r P a l e o z o i c strata t h r o u g h t h e P e r m i a n , b u t t h o s e f r o m t h e

Molluscs 121
722 A Sea without Fish
C i n c i n n a t i a n a r e a m o n g the o l d e s t - k n o w n c a s e s . S p e c i m e n s o f Cyclonema Figure 9.3. Cincinnatian
varicosum a t t a c h e d to Pycnocrinus are k n o w n from the L e x i n g t o n L i m e - gastropods. A. Cy-

stone, just below t h e C i n c i n n a t i a n ( F e l t o n , p e r s . c o m m . ) . W h e t h e r all clonema varicosum

Hall, CMC IP 51118, Cyn-
cases o f snails a t t a c h e d t o P a l e o z o i c c r i n o i d s are i n s t a n c e s o f c o p r o p h a g y
thiana Formation, Lexing-
has b e e n d e b a t e d ; o t h e r possibilities i n c l u d e p a r a s i t i s m , p r e d a t i o n , a n d
ton, Kentucky, x 1.7. B.
commensalism ( B a u m i l l e r 1990; M o r r i s a n d F e l t o n 1993). (In c o m m e n s a l -
Cyclonema humero-
ism, individuals of the associated species gain s o m e advantage w h i l e the sum, Steven Felton col-
host i s u n a f f e c t e d . G i v e n the size a n d l o c a t i o n o f t h e s n a i l s o n t h e c r i n o i d lection, showing aper-
h o s t , it is difficult to see how the host c o u l d r e m a i n u n a f f e c t e d . ) A m o r e ture. Grant Lake
c o m p l e x association b e t w e e n C i n c i n n a t i a n g a s t r o p o d s , c r i n o i d s , a n d t u b e s Formation, Brown Co.,

of Cornulites e n c r u s t i n g t h e snails w a s i n v e s t i g a t e d by M o r r i s a n d F e l t o n Ohio, x 1.2. C. Cy-

clonema humerosum
(1993, 2003). These a u t h o r s s u g g e s t e d that Cornulites g a i n e d an a d v a n t a g e
Ulrich, CMC IP 51117,
b y e n c r u s t i n g the snails that lived o n the e l e v a t e d c r o w n s o f t h e c r i n o i d s ,
Maysvillian, Cincinnati,
either by s h a r i n g in the fecal feast or simply by v i r t u e of t h e e l e v a t e d p o s i -
Ohio, x 1.6. D. Cy-
tion provided by the c r i n o i d . clonema bilix lata (Con-
Most i n d i v i d u a l s of Cyclonema that are f o u n d are n o t a t t a c h e d to cri- rad), CMC IP 51116, Arn-
noids, s u g g e s t i n g that t h e a s s o c i a t i o n w i t h c r i n o i d s w a s not o b l i g a t e , a n d heim Formation,
that s p e c i m e n s of Cyclonema w e r e a b l e to m a k e their l i v i n g in o t h e r ways. Cincinnati, Ohio, x
1.7. E. Cyclonema
W e lack direct e v i d e n c e for t h e f e e d i n g habits o f f r e e - l i v i n g s p e c i m e n s o f
sublaeve Ulrich, CMC IP
Cyclonema, b u t the f e e d i n g habits o f p r e s e n t - d a y g a s t r o p o d s m a y p r o v i d e
51114, Fairview Forma-
s o m e c l u e s . Cyclonema. a l o n g with the m a j o r i t y o f taxa o f C i n c i n n a t i a n
tion, Cincinnati, Ohio, x
gastropods, is placed in the order A r c h a e o g a s t r o p o d a . L i v i n g archaeogas- 2.1. F. Cyclonema
t r o p o d s i n c l u d e m a n y f a m i l i a r f o r m s , for e x a m p l e , l i m p e t s , a b a l o n e s , a n d gracile striatulum Ul-
p e r i w i n k l e s . M o s t p r e s e n t - d a y a r c h a e o g a s t r o p o d s are h e r b i v o r e s t h a t live rich, CMC IP 51113, Kope
b y g r a z i n g o n a l g a l o r b a c t e r i a l c o a t i n g s o n t h e s u b s t r a t u m , b u t s o m e are Formation, Cincinnati,

predatory- o n s p o n g e s o r o n m i c r o o r g a n i s m s a n d o r g a n i c d e t r i t u s e i t h e r b y Ohio, x 3.2. G. Subu-

lites (Fusispira) Ulrich,
d e p o s i t f e e d i n g o r s u s p e n s i o n f e e d i n g ( W a h l m a n 1992). T h e r e i s n o rea-
CMC IP 51115, internal
son t o s u p p o s e that t y p e - C i n c i n n a t i a n a r c h a e o g a s t r o p o d s c o u l d not h a v e
mold, Kope Formation,
had a s i m i l a r r a n g e of f e e d i n g habits.
Kenton Co., Kentucky, x
O n e s u b g e n u s o f C i n c i n n a t i a n s n a i l s , Subulites (Fusispira), b e l o n g s t o 1.3. H. Paupospira
the order N e o g a s t r o p o d a ; i n d i v i d u a l s o f this t a x o n are n o t a b l e e a c h for moorei (Safford),
h a v i n g a h i g h spire and large size ( f i g u r e 9.3G). P r e s e n t - d a y n e o g a s t r o p o d s MUGM 23292, encrusted
i n c l u d e m a n y f a m i l i a r m a r i n e s n a i l s , for e x a m p l e , the w h e l k s , t h e s p i n y by tabulate coral Pro-
taraea richmondensis,
m u r e x e s , and the v e n o m o u s c o n e s . B e c a u s e m o s t n e o g a s t r o p o d s are p r e d a -
Whitewater Formation,
tory, it is possible that the snails of Subulites (Fusispira) w e r e p r e d a t o r s .
Preble Co., Ohio, x 1.5.
A d d i t i o n a l e v i d e n c e that s o m e t y p e - C i n c i n n a t i a n snails w e r e p r e d a -
Note slit in aperture.
tory c o m e s from c i r c u l a r h o l e s f o u n d in brachiopods, as w e l l as o t h e r fos-
sils. M a n y present-day predatory snails attack their prey by " d r i l l i n g "
t h r o u g h the shell of the prey (usually a c l a m ) a n d t h e n i n s e r t i n g a p r o b o s c i s
that secretes d i g e s t i v e e n z y m e s . T h e d r i l l i n g is a c o m b i n a t i o n of d i s s o l u -
tion of the shell by a c i d i c s e c r e t i o n s a n d r a s p i n g by t h e r a d u l a . T h e result-
ing holes generally are p e r f e c t l y c i r c u l a r , c o m m o n l y w i t h a b e v e l e d e d g e
s l o p i n g i n w a r d , but t h e r e i s m u c h v a r i a t i o n i n size a n d f o r m o f t h e h o l e s .
C i r c u l a r h o l e s in t y p e - C i n c i n n a t i a n shells, c h i e f l y brachiopods, h a v e at-
tracted the a t t e n t i o n o f m a n y w o r k e r s o v e r the y e a r s , F e n t o n a n d F e n t o n
(1931) first favored the i n t e r p r e t a t i o n that p r e d a t o r y snails w e r e r e s p o n s i b l e
for the b o r i n g s . B u c h e r (1938) p o i n t e d o u t that b o r e d brachiopods a n d p e l e -
c y p o d s are very rare in the t y p e - C i n c i n n a t i a n , b u t d e s c r i b e d a t h i n b e d

Molluscs 123
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.

rich in i n d i v i d u a l s of t h e b r a c h i o p o d Onniella from t h e R i c h m o n d i a n in

w h i c h t h e f r e q u e n c y o f b o r e d shells i s c l o s e r t o t h e f r e q u e n c i e s o f b o r e d
s h e l l s i n p r e s e n t - d a y s e t t i n g s . C a r r i k e r a n d Y o c h e l s o n (1968) c o m p a r e d
b o r i n g s i n M i d d l e a n d U p p e r O r d o v i c i a n b r a c h i o p o d s from K e n t u c k y with
present-day borings m a d e by gastropods. A l t h o u g h they found m a n y simi-
larities b e t w e e n O r d o v i c i a n a n d p r e s e n t - d a y b o r i n g s , t h e y c o n c l u d e d that
g a s t r o p o d s w e r e not n e c e s s a r i l y r e s p o n s i b l e for the O r d o v i c i a n b o r i n g s .
K a p l a n a n d B a u m i l l e r (2000) s u g g e s t e d that t h e v a r i a t i o n s i n b o r i n g s f o u n d
in O r d o v i c i a n shells c o u l d result f r o m many different borers. Their analysis
of the Onniella shell b e d r e v e a l e d a p r e f e r e n c e in b o r i n g for t h e c o n v e x
p e d i c l e v a l v e . H o w e v e r , c o n t r o v e r s y c o n t i n u e s t o rage o v e r t h e interpreta-
t i o n o f t h e s e b o r i n g s . W i l s o n a n d P a l m e r (2001) p o i n t e d o u t that, i n the
s a m e Onniella shell b e d , s o m e b o r i n g s w e r e d r i l l e d o u t w a r d f r o m t h e in-
terior o f d i s a r t i c u l a t e d v a l v e s , a n d o t h e r s p e n e t r a t e not o n l y t h e m a r g i n s o f
v a l v e s b u t a l s o t h e a d j a c e n t s u b s t r a t u m . Their i n t e r p r e t a t i o n was that t h e s e
b o r i n g s are n o t p r e d a t o r y b o r i n g s at a l l , but rather w e r e drilled into the
shells c e m e n t e d into a h a r d g r o u n d s u r f a c e a s d w e l l i n g s o f u n k n o w n o r g a n -

124 A Sea without Fish

isms. In a reply, K a p l a n a n d B a u m i l l e r (2001) stated that their s a m p l e s are Figure 9.5. Pelecypod
not from h a r d g r o u n d s , a n d t h e y c o n t i n u e d t o r e g a r d b o r i n g s a s predatory mollusc, showing internal

i n n a t u r e . T h e o n g o i n g a r g u m e n t s s h o w that b o r i n g s i n t h e t y p e - C i n c i n - features. Drawing by

Kevina Vulinec.
natian are p r o b a b l y the result o f m a n y t y p e s o f b e h a v i o r b y d i f f e r e n t o r g a n -
isms, a n d , a l t h o u g h not all c a n b e a s s u m e d t o b e predatory, n e v e r t h e l e s s
s o m e w e r e very likely p r o d u c e d by d r i l l i n g predation very s i m i l a r to g a s t r o -
p o d d r i l l i n g i n present-day e n v i r o n m e n t s .
C i n c i n n a t i a n g a s t r o p o d s s o m e t i m e s are f o u n d i n g r e a t a b u n d a n c e i n
t h i n b e d s a l m o s t t o t h e e x c l u s i o n o f o t h e r fossils. A l t h o u g h s u c h b e d s
o c c u r in many units in the section, two o c c u r r e n c e s deserve special note.
O n e is within the M i a m i t o w n Shale, w h i c h is Maysvillian in age, and the
other i s the M a r b l e Hill B e d ( R i c h m o n d i a n ) ( F i g u r e 9.4). T h e M i a m i t o w n
S h a l e is a t h i n , s h a l e - d o m i n a t e d f o r m a t i o n that a t t a i n s a m a x i m u m thick-
ness o f f i v e m e t e r s a t M i a m i t o w n , n e a r t h e G r e a t M i a m i River, a n d t h i n s
t o less than o n e m e t e r n e a r C i n c i n n a t i ( D a t t i l o 1996). W i t h i n t h e M i a n i -
itown is a layer, n i c k n a m e d the " g a s t r o p o d s h a l e , " a b o u t 1.5-2 m e t e r s t h i c k ,
characterized by abundant molluscs, i n c l u d i n g p e l e c y p o d s , the m o n o p l a -
cophoran Cyrtolites, and g a s t r o p o d s ( F i g u r e 9.4A). N e a r t h e top of this
interval is a t h i n l i m e s t o n e that is p a c k e d w i t h g a s t r o p o d s , m o s t l y of
Paupospira bowdeni (referred to Loxoplocus in o l d e r literature), p r e s e r v e d
a s internal m o l d s . T h e s o - c a l l e d " g a s t r o p o d s h a l e , " a l o n g w i t h t h e t h i n
l i m e s t o n e , c a n b e traced from M i a m i t o w n , i n O h i o , t o n o r t h e r n K e n t u c k y ,
a distance of 20 km. T h e p a l e o e n v i r o n m e n t a l significance of the snail-rich
M i a m i t o w n b e d i s not entirely clear. T h e o v e r l y i n g B e l l e v u e L i m e s t o n e ,
w i t h its t h i n , w a v y b e d s c o n t a i n i n g large b r a c h i o p o d s a n d b r y o z o a n s , i n d i -
cates very shallow water, s h o a l i n g c o n d i t i o n s . T h u s t h e M i a m i t o w n c o u l d
represent a slightly d e e p e r e n v i r o n m e n t , p e r h a p s l a g o o n s s h e l t e r e d b e -
t w e e n shell-rich shoals o f t h e B e l l e v u e . O r g a n i c - r i c h m u d s t h a t a c c u n i u -

Molluscs 125
Figure 9.6. Cincinnatian pelecypods. A. Ambonychia sp., MUGM De549A, right valve, Cincinnatian, no
locality information, x 0.6. B. Ambonychia cultrata (Ulrich), MUGM 29546, right valve, Waynesville For-
mation, Butler Co., Ohio, x 0.8. C. Carotidens demissa (Conrad), MUGM 29431, left valve, Corryville For-
mation, Clermont Co., Ohio, x 1.1. D. Anomalodonta gigantea Miller, CMC IP 35898, internal mold of
right valve, Waynesville Formation, Warren Co., Ohio, x 0.8. E. Opisthoptera casei (Meek and Worthen),
MUGM 23392, right valve, Whitewater Formation, Preble Co., Ohio, x 0.9. F. Ischyrodonta truncata Ul-
rich, MUGM 15066, internal mold of right valve, Whitewater Formation, Butler Co., Ohio, x 1.1.

126 A Sea without Fish

Figure 9.7. Cincinnatian pelecypods and rostroconchs. A. Modiolopsis sp. cf. M. modiolaris (Conrad),
USNM 46230, "butterflied" specimen with black organic film and remnants of ligament, Fairview Forma-
tion, Covington, Kentucky, x 0.85. From Pojeta (1971, plate 15, figure 6). B. Modiolopsis modiolaris
(Conrad), USNM 46719, internal mold preserved at right angles to bedding and highly foreshortened,
Fairview Formation, Cincinnati, Ohio, x 2.4. From Pojeta et al. (1986, plate 17, figure5). C. Technopho-
rus faberi Miller, USNM 07219, left-lateral view, Kope Formation, Covington, Kentucky, x 3. From Pojeta
and Runnegar (1976, plate 11, figure 3). D. Technophorus milleri Pojeta and Runnegar, MUGM 6848,
right-lateral view, Whitewater Formation, Butler Co., Ohio, x 5.1. From Pojeta and Runnegar (1976, plate
14, figure 7).

lated in the l a g o o n s c o u l d h a v e p r o v i d e d a h a b i t a t f a v o r a b l e for s n a i l s , b u t

hostile to filter-feeding b r a c h i o p o d s a n d b r y o z o a n s .
The M a r b l e Hill Bed is a n o t h e r r e m a r k a b l e o c c u r r e n c e of g a s t r o p o d s ,
in w h i c h l i m e s t o n e lenses up to about o n e m e t e r thick are p a c k e d with s p e c i -
m e n s of three s p e c i e s of g a s t r o p o d s , Paupospira bowdeni, P. tropidophora
( M e e k ) , and P. moorei ( U l r i c h ) ( F e l t o n , pers. c o m m . ; f i g u r e 9.4B). This b e d
o c c u r s near the top o f the C i n c i n n a t i a n i n the R o w l a n d M e m b e r o f the
D r a k e s f o r m a t i o n (following stratigraphic n o m e n c l a t u r e a d o p t e d i n K e n -

Molluscs 127
Figure 9.8. A. Basal side of stromatoporoid with borings made by pelecypods, some of which are occu-
pied by the pelecypod Corallidomus scobina, OSU 8420, probably from the Waynesville Formation,
Brown Co., Ohio, x 0.6. B. Corallidomus scobina, USNM 70458, bryozoan colony with pelecypods and
inarticulate brachiopod, Trematis sp., in life position among bryozoan branches, Waynesville Formation,
Clinton Co., Ohio, x 0.7. From Pojeta (1971, plate 16, figure 5). C. Corallidomus scobina, USNM 70458,
left valve of specimen attached to bryozoan colony shown in B, x 2.8. D. Carotidens sp., USNM 162734,
left valve with edrioasteroid Isorophus cincinnatiensis and encrusting bryozoans, Corryville Formation,
Clermont Co., Ohio, x 1.4. From Pojeta (1971, plate 10, figure 15). A and C, courtesy of John Pojeta, Jr.

128 A Sea without Fish

tucky; this is a p p r o x i m a t e l y e q u i v a l e n t to the top of the Waynesville F o r m a - Figure 9.9. Life habits of
tion as used in other literature). T h e M a r b l e Hill B e d is e x p o s e d n e a r B e d - Late Ordovician pelecy-

ford, i n Trimble C o u n t y , K e n t u c k y , a n d a d j a c e n t s o u t h e a s t e r n I n d i a n a pods. From Pojeta (1971,

figure 9).
(Hattin et al. 1961; S w a d l e y 1979). L i k e the M i a m i t o w n snail b e d , the M a r b l e
Hill B e d lies in close stratigraphic p r o x i m i t y to rocks of very shallow water
d e p o s i t i o n a l e n v i r o n m e n t s . S w a d l e y (1979) interpreted the M a r b l e Hill B e d
as a c o m p l e x of offshore shoals a n d t i d a l - c h a n n e l deposits a l o n g the m a r g i n
of a s h a l l o w l a g o o n a l area a d j a c e n t to intertidal mudflats.

This is the g r o u p that, today, is r e p r e s e n t e d by c l a m s , m u s s e l s , s c a l l o p s , Pelecypods: a.k.a.,

c o c k l e s , oysters, a n d their relatives ( F i g u r e s 9.5,9.6). T h e s e are all b i v a l v e d The Bivalves
m o l l u s c s e a c h shell consists o f t w o v a l v e s . (For t h a t r e a s o n , s o m e folks
use a different n a m e for t h e c l a s s , viz., B i v a l v i a . A l t h o u g h s u c h u s a g e m a y
be l o g i c a l , it b e l i e s t h e fact that t h e r e are o t h e r b i v a l v e d a n i m a l s that are-
not a t all c l o s e l y related t o t h e m o l l u s c s . E x a m p l e s i n c l u d e t h e m e m b e r s
of p h y l u m Brachiopod a and the ostracods, of phylum Arthropoda.)
This, o f c o u r s e , b r i n g s u p t h e q u e s t i o n : h o w d o e s o n e tell the r e m a i n s
o f the several m a j o r g r o u p s o f b i v a l v e d o r g a n i s m s f r o m o n e a n o t h e r ? F r o m
our p e r s p e c t i v e , in t h e t y p e - C i n c i n n a t i a n , it is i m p o r t a n t to be a b l e to dif-
ferentiate p e l e c y p o d s from b r a c h i o p o d s . T h e s e latter are v e r y c o m m o n i n
the local r o c k s , a n d their shells fall into the s a m e s i z e r a n g e a s d o t h o s e o f
p e l e c y p o d s . O n e c l u e c o m m o n l y i s p r o v i d e d b y t h e state o f p r e s e r v a t i o n .
G e n e r a l l y , i n the C i n c i n n a t i r e g i o n , p e l e c y p o d s are m o r e p o o r l y p r e s e r v e d
t h a n are b r a c h i o p o d s (see c h a p t e r 8 ) a n d o c c u r a s m o l d s . O n e o f t h e not-
u n c o m m o n modes of preservation is o n e called a composite mold. Here,
the a r a g o n i t e shell i s g o n e , a n d t h e e x t e r n a l a n d i n t e r n a l m o l d s are super-

Molluscs 129
 i m p o s e d o n t o o n e a n o t h e r , so that t h e features of the exterior of a valve and
t h o s e of t h e interior are visible in e s s e n t i a l l y a s i n g l e s u r f a c e . The shell
m a t t e r o f t h e l i v i n g p e l e c y p o d s o r e p r e s e n t e d m u s t h a v e b e e n fairly h i g h
in organic material, b e c a u s e the composite mold c o m m o n l y includes a
c a r b o n f i l m w h e r e t h e shell s u b s t a n c e o n c e w a s ( F i g u r e 9.7A).
A l t h o u g h p r e s e r v a t i o n may provide a c l u e , the real way to identify the
p e l e c y p o d s is by r e f e r e n c e to t h e m o r p h o l o g y of t h e shell. The p l a n e of
s y m m e t r y in a t y p e - C i n c i n n a t i a n pelecypod is b e t w e e n t h e valves, w h e r e a s ,
i n a n o r d i n a r y a r t i c u l a t e b r a c h i o p o d , t h e p l a n e o f s y m m e t r y r u n s across
e a c h v a l v e p e r p e n d i c u l a r t o the h i n g e (see F i g u r e 8 . 1 ) .
O f c o u r s e , t h e fossils i n t h e r o c k s i n a n d n e a r C i n c i n n a t i h a v e b e e n
t h e r e for h u n d r e d s o f m i l l i o n s o f years. M a n y t h i n g s m i g h t h a v e h a p p e n e d
d u r i n g that span o f t i m e . S o m e t i m e s , for e x a m p l e , the d e a d shell w a s bur-
ied i n o t h e r t h a n a l i v i n g p o s i t i o n . A s soft s e d i m e n t s b u i l t u p ever d e e p e r
o n t h e sea floor, t h e y b e c a m e c o m p r e s s e d into rock. I n s o m e i n s t a n c e s , the
o r i g i n a l bilateral s y m m e t r y o f t h e o n c e - l i v i n g a n i m a l (and, h e n c e , o f the
shell) w a s distorted by t h e p r e s s u r e , so that the shell a p p e a r s s k e w e d . In
s u c h i n s t a n c e s , the o r i g i n a l s y m m e t r y m a y not b e i m m e d i a t e l y o b v i o u s .

P e l e c y p o d s of t h e Type Cincinnatian

B e c a u s e p e l e c y p o d s are c o m m o n l y p r e s e r v e d a s m o l d s i n the t y p e - C i n c i n -
n a t i a n , they are easily o v e r l o o k e d a n d c a n be difficult to identify. I n d e e d ,
m a n y s p e c i m e n s are n o t h i n g m o r e t h a n c l a m - s h a p e d blobs. O n e e x c e p t i o n
is i n d i v i d u a l s of t h e pterioid g e n u s Carotidens that are p r e s e r v e d as calcitic
shells t h r o u g h o u t t h e t y p e - C i n c i n n a t i a n ( F i g u r e 9 . 6 C ) . F o r m s preserved
a s m o l d s that are very c o m m o n are o f t h e g e n u s Ambonychia ( F i g u r e s 9.6A,
B) a n d t h e g e n u s Modiolopsis ( F i g u r e s 9.7A, B). O c c a s i o n a l l y , pelecypods
are p r e s e r v e d in life p o s i t i o n , as in t h e c a s e of t h e b u r r o w i n g Modiolopsis
( F i g u r e 9.7B), or Corallidomus, t h e e a r l i e s t - k n o w n c a s e of pelecypod bor-
i n g into a h a r d s u b s t r a t u m ( F i g u r e s 9.8A, B; Pojeta a n d P a l m e r 1976).
Epizoa a t t a c h e d to o n e valve of an a r t i c u l a t e d pelecypod may also i n d i c a t e
t h e p r e f e r r e d life o r i e n t a t i o n o f t h e p e l e c y p o d ( F i g u r e 9.8D).
I n c l u d i n g t h e c o m m o n l y e n c o u n t e r e d f o r m s , a c o n s i d e r a b l e diversity
o f p e l e c y p o d s has b e e n d o c u m e n t e d i n the t y p e - C i n c i n n a t i a n . H o l l a n d
(2005) listed 164 s p e c i e s of p e l e c y p o d s , in t h i r t y - e i g h t g e n e r a , for t h e O r d o -
vician rocks of the C i n c i n n a t i region. However, the originally described
s p e c i m e n s o f several g e n e r a a n d n u m e r o u s s p e c i e s are either poorly pie-
s e r v e d , o f u n c e r t a i n t a x o n o m i c status, o r b o t h , s o that the a c t u a l diversity
is surely l o w e r . R o b e r t Frey (1987a) traced the diversity a n d a b u n d a n c e of
pelecypods in t h e f o r m a t i o n s of t h e C i n c i n n a t i r e g i o n . In t h e s h a l e s , silt-
stones, and thin limestones of the Edenian ( K o p e Formation), he recorded
fourteen genera. In Maysvillian t h r o u g h early R i c h m o n d i a n shales and
l i m e s t o n e s , he f o u n d s i x t e e n g e n e r a . In o n e s h a l e b e d of the W a y n e s v i l l e
Formation he found twenty-two species of pelecypods. In Mavsvillian
l i m e s t o n e s , p e l e c y p o d s are g e n e r a l l y less a b u n d a n t t h a n b r a c h i o p o d s and
b r y o z o a n s , b u t i n t h e R i c h m o n d i a n r o c k s , t h e y o c c u r i n h i g h e r diversity.
I n R i c h m o n d i a n l i m e s t o n e s , Frey r e c o r d e d t w e n t y g e n e r a o f p e l e c y p o d s .

130 A Sea without Fish

 Thus. F r e y ' s work d o c u m e n t e d the spread o f p e l e c y p o d s f r o m m a i n l y c l a s -
tic (shale-rich) s e d i m e n t a r y e n v i r o n m e n t s i n early a n d m i d d l e C i n c i n n a -
tian t i m e into c a r b o n a t e e n v i r o n m e n t s in the late C i n c i n n a t i a n . The a b u n -
d a n c e and diversity o f p e l e c y p o d s i n t h e t y p e - C i n c i n n a t i a n d e m o n s t r a t e
that, e v e n e a r l y i n their e v o l u t i o n a r y history, p e l e c y p o d s o c c u p i e d m a r i n e
e n v i r o n m e n t s in a variety of offshore s e t t i n g s , h e r a l d i n g t h e i r s u b s e q u e n t
diversification a n d i n c r e a s i n g a b u n d a n c e i n t h e late P a l e o z o i c a n d M e s o -
z o i c (A. I. M i l l e r 1989).
B y Late O r d o v i c i a n t i m e , p e l e c y p o d s had e x p l o i t e d a l m o s t the full
range of living habits found in present-day forms ( F i g u r e 9.9). M o s t c o m -
monly, Late O r d o v i c i a n p e l e c y p o d s used byssal threads for a t t a c h m e n t to
objects on or w i t h i n the s e d i m e n t s i m i l a r to present-day mytilids (mussels).
Epibyssate forms, such as A m b o n y c h i a , a t t a c h e d at the s e d i m e n t surface or
nestled w i t h i n the b r a n c h e s of b r y o z o a n s . Endobyssate forms, s u c h as Modi-
olopsis and Pseudocolpomya, a t t a c h e d to shell f r a g m e n t s or s e d i m e n t gr ai n s
just b e l o w the s e d i m e n t surface a n d e x t e n d e d the shell for filter f e e d i n g .
F r e e - b u r r o w i n g forms i n c l u d e d deposit feeders (who fed on o r g a n i c particles
with in the sediment), such as Ctenodonta, and shallow infaunal filter feeders,
such as Ischyrodonta. As m e n t i o n e d a b o v e , the ability to b o r e into hard sub-
strata was first seen in the Late O r d o v i c i a n C o r a l l i d o m u s ( F i g u r e s 9 . 8 A - C ) .
O r d o v i c i a n p e l e c y p o d s had not yet d e v e l o p e d the e l o n g a t e siphons a n d m o r e
m u s c u l a r foot that e n a b l e d m e m b e r s of the class to exploit d e e p e r and m o r e
rapid b u r r o w i n g after the e n d of the P a l e o z o i c . Likewise, O r d o v i c i a n p e l e -
c y p o d s had not a c q u i r e d the habit of c e m e n t i n g t h e shell to a hard substrate,
as seen in present-day oysters a n d o t h e r p e l e c y p o d s that inhabit coral reels.
It is also interesting to note that O r d o v i c i a n p e l e c y p o d s l a c k e d the d e v e l o p -
m e n t of thick, c o r r u g a t e d shells and p r o j e c t i n g spines f o u n d in present-da)
forms such as the "giant c l a m " (of g e n u s Tridaena) or "spiny oysters" (of g e n u s
Spondylus). S e l e c t i v e pressure for t h e e v o l u t i o n of these k i n d s of protective
m o r p h o l o g i e s possibly was absent in the Ordov ician sea b e c a u s e s hell- cr us h-
ing ( d u r o p h a g o u s ) predators had not yet evolved.

T h e s q u i d s , c u t t l e f i s h , a n d o c t o p i are t h e m o s t f a m i l i a r c e p h a l o p o d s i n Cephalopods
today's w o r l d , a n d t h e y c e r t a i n l y are not h e a v i l y a r m o r e d . For e x a m p l e ,
m e m b e r s of g e n u s Octopus h a v e no shells at a l l , a n d those of t h e squid a n d
cuttlefish are internal. This is in s t r o n g c o n t r a s t to t h e fossil c e p h a l o p o d s
o f the C i n c i n n a t i r e g i o n a l l k n o w n k i n d s h a d their soft parts p r o t e c t e d b y
an e x t e r n a l shell. (It h a p p e n s that t h e r e is a s i n g l e g e n u s of p r e s e n t - d a y
c e p h a l o p o d s that e a c h has an e x t e r n a l s h e l l , n a m e l y , Nautilus, s h o w n in
Plate 4. This is t h e sole r e m n a n t of t h e vast h o r d e s of e x t e r n a l l y s h e l l e d
c e p h a l o p o d s that o n c e c o u r s e d t h r o u g h the seas o f o u r planet.)
T h e w o r d " c e p h a l o p o d " literally m e a n s " h e a d - f o o t , " w h i c h s e e m s sin-
gularly appropriate for a c r e a t u r e that is c h a r a c t e r i z e d by a p r o m i n e n t h e a d
and a n u m b e r of h i g h l y flexible t e n t a c l e s . A l t h o u g h s u c h soft parts h a v e
not b e e n f o u n d p r e s e r v e d i n t h e rocks o f t h e C i n c i n n a t i r e g i o n , t h e c e -
p h a l o p o d s probably had p r o m i n e n t h e a d s a n d t e n t a c l e s d u r i n g life. W h a t
r e m a i n of the local fossil c e p h a l o p o d s are their e x t e r n a l shells.

Molluscs 131
 In t h e majority, t h e shell is a l o n g , s t r a i g h t , c o n i c a l t u b e c l o s e d at the
narrow e n d . ( S t r a i g h t shells are said to be o r t h o c o n i c . ) There are, h o w e v e r ,
s o m e i n w h i c h t h e shell i s c u r v e d o r e v e n c o i l e d . I n m o s t o f t h e k i n d s o f
c e p h a l o p o d s p r e s e n t , t h e r e are p a r t i t i o n s that g o across the t u b e for part o f
its l e n g t h ; t h e s e are c a l l e d septa ( s i n g u l a r , s e p t u m ) . The c h a m b e r s that are
s e p a r a t e d b y t h e septa are c a l l e d c a m e r a e (singular, c a m e r a ) . A t t h e larger
e n d of t h e c o n i c a l t u b e t h a t c o m p r i s e s t h e shell is a p o r t i o n in w h i c h t h e r e
are n o septa. T h i s i s c a l l e d t h e b o d y c h a m b e r . The c a m e r a e are c o n n e c t e d
to o n e a n o t h e r by a t u b e t h a t r u n s t h r o u g h all the septa from the b o d y
c h a m b e r to t h e first c a m e r a at t h e tip of t h e c o n e ; this i n n e r t u b e is c a l l e d
the siphuncle. T h e l i n e a l o n g w h i c h a p a r t i c u l a r s e p t u m m e e t s t h e outer
w a l l of t h e shell is c a l l e d a s u t u r e .
A l l p r e s e n t - d a y c e p h a l o p o d s live i n t h e o c e a n s , a n d t h e r e i s e v e r y
r e a s o n t o c o n c l u d e that t h e fossil f o r m s w e r e m a r i n e c r e a t u r e s , too. D u r i n g
life, t h e a p t l y - n a m e d b o d y c h a m b e r was o c c u p i e d b y the b u l k o f the soft
parts o f t h e c e p h a l o p o d a n i m a l , a n d t h e o t h e r c a m e r a e a p p a r e n t l y c o n -
t a i n e d gas. (In p r e s e n t - d a y Nautilus, the gas is s i m i l a r in c o m p o s i t i o n to
air, b u t w i t h o u t t h e o x y g e n . ) S u c h a gas-filled shell w o u l d h a v e served as a
float, b u o y i n g the a n i m a l up in the water and, d e p e n d i n g on h o w m u c h
gas w a s in t h e c a m e r a e , a l l o w i n g the a n i m a l to stay at a p a r t i c u l a r water
d e p t h w i t h o u t t h e effort o f s w i m m i n g u p w a r d s o r d o w n w a r d s . A t great
d e p t h s , o f c o u r s e , t h e r e i s t r e m e n d o u s w a t e r p r e s s u r e s o m u c h pressure,
in f a c t , that s u c h a gas-filled shell w o u l d h a v e b e e n c r u s h e d . T h u s , e x t i n c t
c e p h a l o p o d s w i t h e x t e r n a l shells a l m o s t c e r t a i n l y lived i n w a t e r n o d e e p e r
t h a n a few h u n d r e d m e t e r s , a n d m o s t of t h e m p r o b a b l y s w a m in water a
g o o d d e a l s h a l l o w e r t h a n that.
M o s t p r e s e n t - d a y c e p h a l o p o d s are a c t i v e s w i m m e r s . T h e y take water
into t h e m a n t l e - c a v i t y , a n d t h e y e x p e l it t h r o u g h a s p o u t c a l l e d t h e h y p o -
n o i n e w h i c h i s l o c a t e d b e l o w t h e t e n t a c l e s a n d m o u t h (Plate 4 A ) . This
c a u s e s t h e a n i m a l t o m o v e i n t h e o p p o s i t e d i r e c t i o n ; the h y p o n o m e , h o w -
ever, is very flexible, so t h a t t h e a n i m a l c a n s w i m in a l m o s t any d i r e c t i o n .
W e a s s u m e that e x t i n c t c e p h a l o p o d s s w a m the s a m e way.
A l l c e p h a l o p o d s t o d a y are predators. There is no reason to s u p p o s e
that e x t i n c t o n e s w e r e a n y d i f f e r e n t . In fact, t h e r e are a few s p e c i m e n s
k n o w n i n w h i c h g u t c o n t e n t s are p r e s e r v e d ; t h e s e c o n f i r m that a t least
s o m e e x t i n c t c e p h a l o p o d s ate o t h e r o r g a n i s m s . (Alas! N o s u c h s p e c i m e n s
are k n o w n f r o m t h e r o c k s o f t h e C i n c i n n a t i region.)
A l l o f t h e c e p h a l o p o d s k n o w n from t h e r o c k s o f t h e area a r o u n d C i n -
c i n n a t i c o m m o n l y are l u m p e d t o g e t h e r a s " n a u t i l o i d s . " I n this s e n s e , the
a s s e m b l a g e of a n i m a l s so d e n o t e d is a " w a s t e b a s k e t " in that it i n c l u d e s a n i -
m a l s f r o m a n u m b e r of d i f f e r e n t taxa that do n o t s e e m to be c l o s e l y related
b i o l o g i c a l l y . (It is rather like l u m p i n g t o g e t h e r h o r s e s , c a t t l e , p i g s , deer,
r h i n o c e r o s e s , tapirs, a n d s o o n , into " h o o f e d m a m m a l s . " ) W h a t i s w o r s e ,
t h e r e is a f o r m a l t a x o n of c e p h a l o p o d s c a l l e d s u b c l a s s N a u t i l o i d e a . Part of
t h e p r o b l e m i s that t h e " n a u t i l o i d s , " i n t h e g e n e r a l s e n s e ( w h i c h i n c l u d e s
t h o s e t h a t a c t u a l l y b e l o n g i n t h e N a u t i l o i d e a ) , c a n l o o k rather s i m i l a r ex-
ternally. H o w e v e r , w h e n w e l l - p r e s e r v e d s p e c i m e n s are c a r e f u l l y b r o k e n o r
c u t o p e n , m e t i c u l o u s s t u d y r e v e a l s that a n u m b e r o f s t r i k i n g l y different

132 A Sea without Fish

 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 shift-
ing toward the midpoint
of the animal's length (G2

B ),2 permitting stability in

a horizontal orientation.
In the lower diagram, the
nautiloid has grown, with
additional cameral de-
posits formed closer to
the head, so that the
centers of gravity and
buoyancy (G ,B )
3 3 main-
kinds o f " n a u t i l o i d s " a r e r e c o g n i z a b l e . T h i s p r e s e n t s a d o u b l e d i l e m m a t o tain a stable horizontal
the fossil c o l l e c t o r . To c u t a s p e c i m e n o p e n a n d s t u d y it c a r e f u l l y n o t o n l y orientation. See text for
requires s p e c i a l i z e d e q u i p m e n t ( w h i c h i s e x p e n s i v e ) , b u t a l s o t r a i n i n g a n d further discussion. From

experience. Moreover, one needs well-preserved m a t e r i a l a n d not m a n y Flower (1957, figures

4-6) and reprinted by
folk w a n t t o c u t u p that p r i z e d , " p e r f e c t " s p e c i m e n . S o m e t i m e s t a p h o n o m y
permission of the Geo-
is k i n d , h o w e v e r , a n d a s p e c i m e n is f o u n d that just h a p p e n s to h a v e b e e n
logical Society of
broken or b e e n e r o d e d in s u c h a way that the i n n e r s e c r e t s are r e v e a l e d .
America.
M u c h of t h e t i m e , h o w e v e r , it is t o u g h to a s s i g n a g i v e n s p e c i m e n to a
h i g h e r taxon w i t h o u t d a m a g i n g it.
M a n y o f t h e internal d e t a i l s o f t h e " n a u t i l o i d s " s e e m t o h e related t o
hydrostatics, that is, t o t h e p o s i t i o n a n d o r i e n t a t i o n o f t h e l i v i n g a n i m a l i n
the water. R e c a l l that t h e shell o f m o s t " n a u t i l o i d s " i n t h e r o c k s o f t h e
C i n c i n n a t i r e g i o n is a l o n g , s t r a i g h t c o n e . T h e b u l k of t h e soft parts of t h e
a n i m a l o c c u p i e d t h e b o d y c h a m b e r , w h e r e a s t h e part o f t h e a n i m a l s u b d i -
vided into c a m e r a e ( t e c h n i c a l l y c a l l e d t h e p h r a g m o c o n e ) s e e m s m o s t l y t o
h a v e b e e n f i l l e d w i t h gas. T h u s , t h e part o f t h e shell t o w a r d t h e a p e x o f t h e
c o n e would have b e e n lighter than the other e n d , w h i c h c o n t a i n e d the
w e i g h t o f the b o d y . T h e result w o u l d h a v e b e e n t h a t t h e c o n e w o u l d h a v e
t e n d e d t o b e o r i e n t e d i n the water w i t h its b l u n t e n d , a n d , h e n c e , a p e r t u r e ,
d o w n a n d , potentially, the a n i m a l ' s f a c e i n t h e m u d o f t h e sea floor.

Molluscs 133
Figure 9 . 1 1 . Cincinnatian A p p a r e n t l y , f r o m t h e p o i n t o f v i e w o f o r g a n i c e v o l u t i o n , this w a s not a
orthoconic nautiloids. b e n e f i c i a l s i t u a t i o n . At least in a n u m b e r of e v o l u t i o n a r y l i n e a g e s , c h a n g e s
A-C from Davis and
in t h e shell o c c u r r e d that solved this p r o b l e m in o n e w a y or a n o t h e r , to a
Mapes (1996), courtesy
g r e a t e r or less e x t e n t .
of the Ohio Department
B e f o r e p r o c e e d i n g f u r t h e r on this t a c k , we n e e d to talk a bit a b o u t
of Natural Resources Divi-
sion of Geological Sur- o r g a n i c e v o l u t i o n , per s e l e s t w e g o astray into t h i n k i n g that, b e c a u s e the

vey. A, B. Treptoceras l i v i n g cephalopod did n o t " l i k e " its f a c e in the m u d , it d e l i b e r a t e l y e v o l v e d

duseri (Hall and Whit- its shell m o r p h o l o g y a n d s t r u c t u r e to avoid that c o n d i t i o n .
field), Cincinnatian, scale As far as is k n o w n , the volition of o r g a n i s m s has no effect on the organic
bars = 1 cm. A. OSU e v o l u t i o n of a l i n e a g e . T h u s , no c r e a t u r e c a n will the o r g a n i c evolution of its
47422, internal mold of
d e s c e n d e n t s to go into a n y p a r t i c u l a r d i r e c t i o n , no matter how beneficial (or
part of phragmocone
enjoyable?) that direction m i g h t prove to be. Rather, it a particular direction
and living chamber.
m e a n s that m o r e creatures will survive to pass their g e n e s on to future genera-
B. OSU 47417, part of
tions, t h e n t h e o r g a n i c e v o l u t i o n of a l i n e a g e w ill tend in that direction. This
external shell, with dark
m e c h a n i s m is c a l l e d o r g a n i c e v o l u t i o n by natural s e l e c t i o n , or natural selec-
longitudinal lines which
are remains of what, in tion, for short. S o m e paleontologists and other biologists speak of evolutionary

life, were color bands. "strategics" w h e r e b y a l i n e a g e "solves" s o m e p a r t i c u l a r " p r o b l e m . " These are

C. Cameroceras in- figures of s p e e c h . They are, in tact, g e n e r a l i z a t i o n of what c h a n g e s actually

aequabile (Miller), inter- are seen in t h e fossil record of the p a r t i c u l a r l i n e a g e . they definitely do not
nal mold of portion of imply that o r g a n i s m s p e r c e i v e d a p r o b l e m and willfully evolved to solve it.
siphuncle, OSU 47420, L e t u s r e t u r n t o t h e t y p e - C i n c i n n a t i a n . O n e w a y t o b r i n g t h e shell o f
Cincinnatian, scale bar =
an o r t h o c o n i c cephalopod into a h o r i z o n t a l o r i e n t a t i o n w o u l d be to add
1 cm. D. Endoceras
w e i g h t to the p h r a g m o c o n e ( f i g u r e 9.10). This w a s d o n e in v a r i o u s w a y s in
sp., MUGM 29579, inter-
v a r i o u s l i n e a g e s ; t h e r e are t w o m a i n c a t e g o r i e s i n t h e " n a u t i l o i d s " :
nal mold of part of
phragmocone and sip-
1 . d e p o s i t i o n o f c a l c i u m c a r b o n a t e w i t h i n the c a m e r a e (result: c a m e r a l
huncle, Liberty Forma-
deposits);
tion, Butler Co., Ohio,
2 . d e p o s i t i o n o f c a l c i u m c a r b o n a t e w i t h i n t h e s i p h u n c l e (result: s i p h u n -
scale in mm. E. Gor-
byoceras duncanae c u l a r deposits).

Flower, CMC IP 31393,

Whether t h e s e d e p o s i t s are p r e s e n t , a n d , if so. w h e r e in the shell they
external shell with fine
o c c u r a n d w h a t f o r m t h e y take a r e very i m p o r t a n t i n d e t e r m i n i n g t h e
longitudinal lines, White-
p r o p e r t a x o n o m i c p o s i t i o n of a g i v e n fossil. S i p h u n c u l a r deposits o c c u r in
water Formation, x 1.4.
a n e s p e c i a l l y g r e a t v a r i e t y o f c o n f i g u r a t i o n s . I n the s u b c l a s s E n d o c e r a -
t o i d e a , for e x a m p l e , t h e s i p h u n c u l a r d e p o s i t s c o m p r i s e a S e r i e s o f c o n e s
s t a c k e d e a c h i n s i d e t h e n e x t , w i t h t h e larger e n d toward the a p e r t u r e o f the
s h e l l ; t h e s e are c a l l e d e n d o c o n e s .
A n o t h e r way to "solve" the orientation "problem" would be to bring
t h e c e n t e r o f b u o y a n c y ( r e s u l t i n g f r o m t h e gas i n the c a m e r a e ) a b o v e the
c e n t e r o f gravity ( r e s u l t i n g e s p e c i a l l y f r o m t h e m a s s o f the a n i m a l i n the
b o d y c h a m b e r ) . The m o s t o b v i o u s w a y t o d o this w o u l d b e t o c o i l the shell
s o that t h e b o d y c h a m b e r h a n g s b e n e a t h the gas-filled p h r a g m o c o n e ; t h e
p r e s e n t - d a y Nautilus is t h e p r o d u c t of s u c h an e v o l u t i o n a r y l i n e a g e .

C e p h a l o p o d s of the Type Cincinnatian

N a u t i l o i d c e p h a l o p o d s are c o m m o n fossils t h r o u g h o u t the t y p e - C i n c i n n a -

tian. they g e n e r a l l y o c c u r a s i n t e r n a l m o l d s o f partial p h r a g m o c o n e s .
B e c a u s e the o r i g i n a l a r a g o n i t i c shell m a t e r i a l w a s not p r e s e r v e d , the q u a l -

134 A Sea without Fish

Molluscs 135
 ity of n a u t i l o i d p r e s e r v a t i o n is often p o o r , m a k i n g i d e n t i f i c a t i o n difficult.
However, e x c e p t i o n a l p r e s e r v a t i o n of n a u t i l o i d s in a 1.5-meter-thick s h a l e
or c l a y s t o n e w i t h i n the W a y n e s v i l l e f o r m a t i o n f o r m e d the basis for a d e -
tailed s t u d y o f t y p e - C i n c i n n a t i a n n a u t i l o i d s b y R o b e r t C . Frey (1988,1989)
that p r o v i d e d u n i q u e insight into t h e p a l e o e c o l o g y o f C i n c i n n a t i a n n a u -
tiloids. This s h a l e , i n f o r m a l l y c a l l e d t h e "Treptoceras duseri s h a l e " for the
s p e c i e s o f the m o s t a b u n d a n t n a u t i l o i d s , o c c u r s i n W a r r e n a n d C l i n t o n
c o u n t i e s of O h i o . F r e y r e p o r t e d a total of t w e l v e s p e c i e s of n a u t i l o i d s , in
e i g h t g e n e r a , from this u n i t , of w h i c h t h r e e s p e c i e s of Treptoceras a c c o u n t e d

136 A Sea without Fish

for 7 6 p e r c e n t o f t h e 302 s p e c i m e n s f o u n d . S p e c i m e n s o f o t h e r s p e c i e s are Figure 9.12. Upper Cin-
m u c h less a b u n d a n t ; i n c l u d e d h e r e are t h e c y r t o c o n i c f o r m s ( h a v i n g cinnatian (Richmondian)

c u r v e d shells) Manitoulinoceras tenuiseptum a n d M. williamsae (8 p e r c e n t ) , nautiloids. A. Charac-

toceras baeri (Meek and
Oncoceras delicatum (4 p e r c e n t ) , Zittelloceras russelli, and Z. williamsae (<1
Worthen), MUGM 29591,
p e r c e n t ) , l o n g i c o n i c f o r m s ( h a v i n g l o n g , t a p e r i n g , s t r a i g h t shells) tenta-
Whitewater Formation
tively a s s i g n e d to g e n u s Isorthoceras (6 p e r c e n t ) , t h e e n d o c e r i d Camero-
Preble Co., Ohio, scale
ceras inaequabile (6 p e r c e n t , h a v i n g a w i d e s i p h u n c l e i n f i l l e d w i t h c o n i c a l bar = 2 cm. Compare to
deposits), the o r t h o c e r i d Gorbyoceras curvatum (<1 p e r c e n t , an o r t h o c o n i c present-day Nautilus,
form), a n d the a s c o c e r i d Schuchertoceras obscurum (<1 p e r c e n t , a f o r m that Plate 4. B. Beloito-
lost part o f t h e p h r a g m o c o n e d u r i n g g r o w t h t o m a i n t a i n stability). T h e ceras amoenum (Miller),
CMC IP 24415, internal
m a t e r i a l o f t h e outer w a l l o f e a c h shell w a s p r e s e r v e d o n l y w h e n e n c r u s t e d
mold of phragmocone of
b y b r y o z o a n s , b u t the septa a n d s i p h u n c u l a r s t r u c t u r e s o f t h e shell interiors
a cyrtoconic form, White-
w e r e r e p l a c e d b y c a l c i t e . B o d y c h a m b e r s a n d p h r a g m o c o n e s w e r e infilled
water Formation, Butler
with c l a y s t o n e , b u t , i n s o m e cases, t h e c a m e r a e r e m a i n e d e m p t y o r w e r e Co., Ohio, x 1. C. Di-
infilled w i t h c a l c i t e crystals. These preservational features led Frey to c o n - estoceras eos (Hall and
c l u d e that the n a u t i l o i d a s s e m b l a g e w a s b u r i e d i n situ a s c o m p l e t e , e m p t y Whitfield), MUGM 382,
shells. R e m o v a l o f e n c r u s t i n g b r y o z o a n s r e v e a l e d t h e r e m a r k a b l e p r e s e r v a - internal mold of partial
tion o f r e m n a n t s o f c o l o r p a t t e r n s o n t h e e x t e r i o r o f t h e s h e l l . I n c o n t r a s t phragmocone and body
chamber, Whitewater
to shells of t h e present-day Nautilus that are k n o w n to float after d e a t h a n d
Formation, Preble Co.,
drift with c u r r e n t s , the t h i n n e r shells a n d septal m o r p h o l o g y o f t h e s e O r -
Ohio, scale in mm.
d o v i c i a n n a u t i l o i d s suggest that d e a d a n i m a l s sank t o the b o t t o m .
The a s s e m b l a g e p r o b a b l y is a g o o d r e p r e s e n t a t i o n of t h e l i v i n g n a u -
tiloid c o m m u n i t y , i n w h i c h i n d i v i d u a l s o f d i f f e r e n t s p e c i e s h a d d i s t i n c t i v e ,
s p e c i a l i z e d m o d e s o f life. L o n g i c o n i c f o r m s p r o b a b l y w e r e a c t i v e s w i m -
mers c l o s e t o t h e sea floor, w h e r e a s c y r t o c o n i c f o r m s m a y h a v e lived c l o s e r
t o the s u b s t r a t u m , p e r h a p s c r a w l i n g o r s w i m m i n g i n short bursts. W i t h n o
f i s h o n the s c e n e d u r i n g t h e C i n c i n n a t i a n , n a u t i l o i d s w e r e u n d o u b t e d l y
the top predators. The a s s o c i a t i o n in t h e s o - c a l l e d Treptoceras duseri s h a l e
o f n a u t i l o i d s w i t h m a n y c o m p l e t e trilobites, s i m i l a r t o o t h e r t r i l o b i t e - n a u -
tiloid a s s e m b l a g e s in the O r d o v i c i a n of N o r t h A m e r i c a , s u g g e s t s a b i o t i c
relationship b e t w e e n t h e t w o g r o u p s ( F r e y 1989). I n d i v i d u a l s o f p r e s e n t - d a y
Nautilus e a c h possess jaws that c o n s i s t b o t h o f o r g a n i c a n d m i n e r a l i z e d
c o m p o n e n t s ; o n t h e o t h e r h a n d , t h e jaws o f t h e s q u i d s o f today's o c e a n s
consist e x c l u s i v e l y o f o r g a n i c m a t e r i a l , a n d , h e n c e , w o u l d not b e p r e s e r v e d
ordinarily. A l t h o u g h jaw s t r u c t u r e s i n t h e O r d o v i c i a n n a u t i l o i d s h a v e n o t
b e e n f o u n d , it is p o s s i b l e that t h e y p o s s e s s e d o r g a n i c jaws by w h i c h t h e y
c o u l d h a v e a t t a c k e d prey s u c h a s trilobites. I n t e r e s t i n g l y , a p p a r e n t w o u n d s
h a v e b e e n f o u n d i n trilobites, for e x a m p l e , o f g e n u s Isotelus, f r o m t h e " 7 .
duseri s h a l e " that c o u l d h a v e r e s u l t e d f r o m a t t a c k s by n a u t i l o i d p r e d a t o r s
(Frey 1989; B a b c o c k 2003). O t h e r i n s t a n c e s o f injuries possibly c a u s e d b y
bites o f n a u t i l o i d s o c c u r i n b r a c h i o p o d s ( A l e x a n d e r 1986), g a s t r o p o d s (Fel-
ton, pers. c o m m . ) , a n d c r i n o i d s ( D o n o v a n a n d S c h m i d t 2001).
T h e association of s p e c i m e n s of Treptoceras, of t h e e n d o c e r i d Camero-
ceras, and of Oncoceras is characteristic of M a v s v i l l l i a n - t o - e a r l y - R i c h m o n -
dian shales in the C i n c i n n a t i region and e l s e w h e r e in eastern N o r t h A m e r i c a
( F i g u r e 9.11; Frey 1989,1995). A nautiloid f a u n a of m u c h greater diversity is
well k n o w n from Late O r d o v i c i a n strata of the s a m e a g e f r o m a w i d e area of
N o r t h A m e r i c a to the west and n o r t h , closer to t h e p a l e o e q u a t o r , in an e n v i -

Molluscs 137
 r o n m e n t of p r e d o m i n a n t l y l i m e s t o n e d e p o s i t i o n (Frey 1989,1995). There are
t w o i n c u r s i o n s o f this "tropical f a u n a " into the C i n c i n n a t i A r c h region: the
first d u r i n g the Edenian, a n d the s e c o n d , a l o n g with a host of o t h e r species
o f invertebrates, d u r i n g middle-to-late R i c h m o n d i a n t i m e (Frey 1989). T h e
oldest R i c h m o n d i a n e l e m e n t s of this " t r o p i c a l " nautiloid a s s e m b l a g e are seen
in the W a y n e s v i l l e "Treptoceras duseri s h a l e " in the form of s p e c i m e n s of
Sehuehertoceras ohscurum a n d Gorbyoceras curvatum (Frey 1985,1995). Rich-
m o n d i a n l i m e s t o n e - r i c h u n i t s , s u c h a s the D r a k e s , S a l u d a , and W h i t e w a t e r
F o r m a t i o n s c o n t a i n a diverse nautiloid fauna that represents the peak of this
" R i c h m o n d i a n invasion." A c c o r d i n g to F l o w e r (1946) and Frey (1995), this
fauna totals sixty-five s p e c i e s in twenty-four g e n e r a , of w h i c h twenty species
are c o m m o n . S o m e characteristic forms are Narthecoceras dunni (Frey, 1981)
and Gharactoceras, Diestoceras, a n d Gorbyoceras, s h o w n in F i g u r e 9.12.

N a u t i l o i d T r a c e Fossils?

T h e late R o u s s e a u H. F l o w e r , o n e of t h e m o s t prolific researchers on P a l e o -

z o i c n a u t i l o i d c e p h a l o p o d s , was inspired b y t y p e - C i n c i n n a t i a n c e p h a l o p o d s
as the first d o c t o r a l s t u d e n t ( P h . D . , 1939) of K e n n e t h E. C a s t e r at the U n i -
versity of C i n c i n n a t i . In a 1955 p a p e r , F l o w e r drew attention to the tact that,
despite t h e a b u n d a n c e o f c e p h a l o p o d s i n the P a l e o z o i c fossil r e c o r d , there
w e r e v i r t u a l l y no reports of trace fossils attributed to activities of nautiloid
c e p h a l o p o d s . H e w e n t o n t o illustrate t w o t y p e - C i n c i n n a t i a n trace fossils
that h e c o n c l u d e d r e p r e s e n t e d i n t e r a c t i o n s o f n e k t o n i c n a u t i l o i d s w i t h the
s e d i m e n t s o f the sea floor. F l o w e r interpreted the c o m m o n " t u r k e y track,"
a t r o u g h - l i k e i m p r e s s i o n now c a l l e d Trichophycus (see c h a p t e r 14, F i g u r e
14.1E) to result f r o m a n a u t i l o i d p l o u g h i n g into t h e b o t t o m . In a rare c a s e ,
a n a u t i l o i d rested at t h e e n d of o n e s u c h t r o u g h . He s h o w e d a sketch of a
b e d d i n g s u r f a c e , t w o t o t h r e e m e t e r s across, o n w h i c h m a i n t r o u g h s were
o r i e n t e d radially a r o u n d a c o n c e n t r a t i o n of fossil debris, w h i c h s u g g e s t e d to
h i m a f e e d i n g p a t t e r n . F l o w e r illustrated a n o t h e r trace that consisted of a
h o r s e s h o e - s h a p e d pattern of shallow, c u r v e d g r o o v e s on the surface of a b e d .
T h e s e he i n t e r p r e t e d to represent i m p r e s s i o n s of tentacles m a d e as a n a u -
tiloid d u g into the b o t t o m for stability in h i g h c u r r e n t flow. S u b s e q u e n t l y ,
w h e n R i c h a r d G . O s g o o d (1970) c o n d u c t e d his analysis o f all k n o w n C i n -
c i n n a t i a n trace fossils. F l o w e r s n a u t i l o i d traces c a m e u n d e r r e n e w e d scru-
tiny. O s g o o d c o n c l u d e d that t h o s e traces c o u l d not b e attributed t o nautiloid
activity. T h e s u p p o s e d n a u t i l o i d i m p r e s s i o n s w e r e s h o w n t o b e basal por-
tions of burrow systems that e x t e n d e d u p w a r d into softer, u s u a l l y e r o d e d
shales. In d e b u n k i n g the n a u t i l o i d i n t e r p r e t a t i o n , O s g o o d (1970) displayed
n o t a b l e g o o d g r a c e i n n a m i n g t h e " t e n t a c u l a r i m p r e s s i o n " i n Flower's
h o n o r : Vascifodina floweri. The p o t e n t i a l for C i n c i n n a t i a n n a u t i l o i d traces
r e m a i n s ; in fact, O s g o o d (1970, plate 83, figure 3) illustrated o n e impression
w i t h raised e d g e s that he c o n s i d e r e d a possible n a u t i l o i d t o u c h d o w n .

The So-Called A l t h o u g h representatives of seven classes of the phylum M o l l u s c a o c c u r in

"Minor Molluscs t h e O r d o v i c i a n r o c k s o f t h e C i n c i n n a t i r e g i o n o r nearby, o n l y t h r e e are

138 A Sea without Fish

c o m m o n . T h e s e a r e the g a s t r o p o d s (snails), t h e p e l e c y p o d s ( c l a m s ) , a n d
the c e p h a l o p o d s (relatives of p r e s e n t - d a y Nautilus, s q u i d , a n d o c t o p u s ) .
Type Cincinnatian representatives o f e a c h o f t h e s e " m a j o r g r o u p s " a r e c o n -
sidered in s o m e d e t a i l a b o v e . In a d d i t i o n to t h e s e b e t t e r - k n o w n c l a s s e s ,
s p e c i m e n s of four less w e l l - k n o w n classes often referred to as t h e " m i n o r
m o l l u s c s " also o c c u r i n the O r d o v i c i a n r o c k s o f t h e C i n c i n n a t i a r e a . T h e s e
are t h e m o n o p l a c o p h o r a n s , t h e r o s t r o c o n c h s , t h e p o l y p l a c o p h o r a n s (chi-
tons), and the s c a p h o p o d s ("tusk shells").

Monoplacophorans

M o n o p l a c o p h o r a n s (there i s n o o t h e r " c o m m o n n a m e " ) are a c t u a l l y f a i r l y

diverse in the t y p e - C i n c i n n a t i a n , w i t h twenty-three s p e c i e s in e i g h t g e n e r a
( F i g u r e 9.2; W a h l m a n 1992; H o l l a n d 2005). M o n o p l a c o p h o r a n s are s i m i l a r
to the present-day g a s t r o p o d s c a l l e d l i m p e t s in that e a c h h a s a s i n g l e , c a p -
s h a p e d or p l a n i s p i r a l l y c o i l e d shell a n d a m u s c u l a r foot for l o c o m o t i o n .
Present-day forms like Neopilina g r a z e on m i c r o b i a l m a t s t h r o u g h u s e of
the r a d u l a . Cyrtolites i s the m o s t c o m m o n t y p e - C i n c i n n a t i a n f o r m ( f i g -
ures 9 . 2 C , D) p r e s e r v e d e i t h e r as a calcitic shell or as an i n t e r n a l m o l d , b u t
c o m m o n l y it is e n c r u s t e d w i t h b r y o z o a n s . The k e e l e d , p l a n i s p i r a l shell
with a d i a m o n d - s h a p e d a p e r t u r e is d i s t i n c t i v e .

Rostroconchs

R o s t r o c o n c h s h a v e a p s e u d o b i v a l v e d shell that is sharply folded a l o n g the

dorsal axis t o g i v e the a p p e a r a n c e o f b e i n g b i v a l v e d , a l t h o u g h c a l c i f i c a t i o n
is c o n t i n u o u s across t h e axis (Pojeta 1987). R o s t r o c o n c h s are restricted to
rocks o f the P a l e o z o i c Era a n d are t h o u g h t t o r e p r e s e n t a n i n t e r m e d i a t e
stage in the e v o l u t i o n from u n i v a l v e d to b i v a l v e d m o l l u s c s (Pojeta a n d
R u n n e g a r 1976). O n l y s p e c i m e n s o f Technophorus o c c u r i n t h e t y p e - C i n -
c i n n a t i a n , w i t h t w o s p e c i e s in the E d e n i a n a n d M a y s v i l l i a n , a n d a third
s p e c i e s i n the R i c h m o n d i a n ( f i g u r e s 9 . 7 C , D ) . R o s t r o c o n c h s p r o b a b l y
lived i n f a u n a l l y as deposit or s u s p e n s i o n f e e d e r s .

Polyplacophorans

Polyplacophorans, c o m m o n l y called "chitons," are familiar to those w h o

h a v e e x p l o r e d the r o c k y intertidal z o n e o f t o d a y s o c e a n s . C h i t o n s differ
from all o t h e r m o l l u s c s in h a v i n g a shell c o m p o s e d of e i g h t s e p a r a t e plates
(called valves) that o v e r l a p like s h i n g l e s to p r o v i d e a p r o t e c t i v e c o v e r i n g
that p e r m i t s f l e x i b i l i t y . P r e s e n t - d a y c h i t o n s c r e e p o v e r h a r d substrata a n d
g r a z e a l g a e with the radula. T h e earliest fossil c h i t o n s a r e f r o m the U p p e r
C a m b r i a n , but t h e y o c c u r m o r e c o m m o n l y i n the O r d o v i c i a n o f N o r t h
A m e r i c a ( H o a r e and Pojeta 2006). A l t h o u g h n o c h i t o n fossils h a v e b e e n
reported a s h a v i n g b e e n f o u n d i n t h e t y p e - C i n c i n n a t i a n , d i s a r t i c u l a t e d
valves o c c u r i n o l d e r U p p e r O r d o v i c i a n strata o f t h e K r o n e L i m e s t o n e o f
the H i g h B r i d g e G r o u p i n c e n t r a l K e n t u c k y .

Molluscs 139
 Scaphopods
S c a p h o p o d s s o m e t i m e s g o b y t h e c o m m o n n a m e " t u s k shells." T h e y are
another group of molluscs of w h i c h s p e c i m e n s potentially could o c c u r in
t h e t y p e - C i n c i n n a t i a n , b u t , a s y e t , n o n e h a v e b e e n d o c u m e n t e d i n the
s c i e n t i f i c literature a s h a v i n g c o n i c f r o m t h e local rocks. Present-day sca-
p h o p o d s e a c h h a v e a s m a l l , c u r v e d c o n i c a l shell o p e n a t b o t h e n d s , a n d
live p a r t l y b u r i e d i n t h e s e d i m e n t a s d e p o s i t feeders. T h e o l d e s t - k n o w n
fossil s c a p h o p o d , Rhytiodentalium kentuckyensis, w a s d e s c r i b e d from t h e
L e x i n g t o n L i m e s t o n e o f K e n t u c k y , the f o r m a t i o n just below the base o f the
C i n c i n n a t i a n (Pojeta a n d R u n n e g a r 1979). Possible s c a p h o p o d s p e c i m e n s
h a v e b e e n f o u n d i n t h e t y p e - C i n c i n n a t i a n ( F e l t o n , pers. c o m m . ) .

140 A Sea without Fish

 Figure 10.1. Cincinnatian worms and worm-like fossils. A. Tentaculites
richmondensis (Miller), CMC IP 17551, Waynesville Formation, Clinton Co.,
Ohio. Slab showing parallel alignment of shells. Scale in mm. B. Annelid
worm, Protoscolex ornatus Ulrich, CMC IP 37990, Kope Formation, Cov-
ington, Kentucky. This is a rare case in the Cincinnatian of soft-body preser-
vation, x 7.5. C. Tubes of Cornulites sp. attached to the column of the
crinoid locrinus subcrassus, University of Cincinnati collections, Corryville
Formation, Hamilton Co., Ohio. Scale in mm. D. The machaeridian Lepido-
coleus sp. cf. L. jamesi (Hall and Whitfield), University of Cincinnati collec-
tions, Corryville Formation, Boone Co., Kentucky. Scale in mm.

142 A Sea without Fish

 10
ANNELIDS AND WORM-LIKE FOSSILS

B e c a u s e w o r m s are largely s o f t - b o d i e d , their fossil r e c o r d is rather l i m i t e d . ... we can easily imagine

N o n e t h e l e s s , n u m e r o u s fossils o c c u r i n t h e C i n c i n n a t i a n that c a n b e at- that the ocean beneath
tributed t o the P h y l u m A n n e l i d a o r related w o r m s . A n n e l i d s , t h e s e g m e n t e d which the Cincinnati

w o r m s , i n c l u d e p r e d o m i n a n t l y freshwater a n d terrestrial l e e c h e s a n d e a r t h - group was deposited,

at times swarmed with
w o r m s , and the p r e d o m i n a n t l y m a r i n e p o l y c h a e t e s . I n t h e m o d e r n o c e a n s
innumerable worms,
p o l y c h a e t e s are h i g h l y d i v e r s e a n d a b u n d a n t a n d p l a y m a n y i m p o r t a n t
which have, so far as
e c o l o g i c a l roles. T h r o u g h o u t the C i n c i n n a t i a n c o m m o n tooth-like m i c r o -
we at present know,
fossils c a l l e d s c o l e c o d o n t s i n d i c a t e that p o l y c h a e t e s w e r e also c o m p o n e n t s left no traces of them-
o f the O r d o v i c i a n m a r i n e e c o s y s t e m ( E r i k s s o n a n d B e r g m a n 2003). A l - selves excepting their
t h o u g h t h e y r e s e m b l e c o n o d o n t s , a n o t h e r c a t e g o r y o f t o o t h - l i k e fossils, i n jaws, tracks, and pos-
size and form s c o l e c o d o n t s are distinct in h a v i n g a jet b l a c k a p p e a r a n c e in sibly a few rude impres-

contrast t o t h e a m b e r c o l o r t y p i c a l o f c o n o d o n t s ( P l a t e 5). T h e d e f i n i t e sions of their bodies.

p o l y c h a e t e affinity of s c o l e c o d o n t s is e s t a b l i s h e d by rare c a s e s (not C i n c i n - E. O. Ulrich 1878, 88

natian) of s c o l e c o d o n t s f o u n d w i t h t h e b o d y fossil of a p o l y c h a e t e w o r m as
a s s e m b l a g e s of paired tooth-like e l e m e n t s f o r m i n g a jaw a p p a r a t u s . In m o d -
ern p o l y c h a e t e s a n entire a p p a r a t u s consists o f t h r e e pairs o f d i f f e r e n t indi-
vidual e l e m e n t s . B e c a u s e s c o l e c o d o n t s u s u a l l y o c c u r a s d i s s o c i a t e d e l e -
m e n t s , their t a x o n o m y has b e e n c o m p l i c a t e d b y a s s i g n m e n t o f s e p a r a t e
n a m e s for e a c h e l e m e n t . I n r e c e n t w o r k t h e r e c o g n i t i o n o f l i k e l y a s s o c i a -
tions o f e l e m e n t s has b e g u n t o a l l e v i a t e this p r o b l e m .
The C i n c i n n a t i a n w a s the s o u r c e o f t h e earliest r e p o r t o f s c o l e c o d o n t s
as w o r m jaws ( G r i n n e l l 1 8 7 7 ) . In a r e c e n t study, s c o l e c o d o n t s c o l l e c t e d
from d i s a g g r e g a t e d s h a l e s o r a c i d - i n s o l u b l e r e s i d u e s f r o m l i m e s t o n e s w e r e
found t o b e c o m m o n t h r o u g h o u t t h e C i n c i n n a t i a n s e c t i o n w i t h a m a x i -
m u m a b u n d a n c e o f 1545 e l e m e n t s p e r k i l o g r a m o f r o c k ( E r i k s s o n a n d
B e r g m a n 2003). E r i k s s o n a n d B e r g m a n e s t i m a t e d that forty t o f i f t y m u l t i -
element species b e l o n g i n g to twelve families o c c u r in the C i n c i n n a t i a n .
A l t h o u g h s o m e s c o l e c o d o n t s r e s e m b l e jaws o f c e r t a i n m o d e r n p o l y c h a e t e
f a m i l i e s , n o n e o f t h e C i n c i n n a t i a n taxa r e p r e s e n t l i v i n g g r o u p s . B e c a u s e
m o d e r n j a w - b e a r i n g p o l y c h a e t e s are p r e d a t o r y , it is likely that C i n c i n n a -
tian s c o l e c o d o n t - b e a r i n g f o r m s w e r e also p r e d a t o r s , p r e s u m a b l y o n soft-
b o d i e d prey. Eriksson a n d B e r g m a n r e c o g n i z e d five s t r a t i g r a p h i c a s s o c i a -
tions o f s c o l e c o d o n t s w i t h i n t h e C i n c i n n a t i a n ; a l s o , m a n y C i n c i n n a t i a n
f a m i l i e s and g e n e r a o c c u r i n t h e U p p e r O r d o v i c i a n o f t h e B a l t i c r e g i o n o f
Europe, indicating an intercontinental distribution. Further refinement of
s c o l e c o d o n t t a x o n o m y w i l l u n d o u b t e d l y fulfill a g r e a t p o t e n t i a l for t h e u s e
of scolecodonts in biostratigraphic zonation and correlation.
In an e x t r e m e l y rare i n s t a n c e of soft-bodied p r e s e r v a t i o n , U l r i c h (1878)
described a c t u a l b o d y f o s s i l s o f w o r m s from a h o r i z o n i n the E d e n i a n . T h e

143
 s p e c i m e n s are very s m a l l , a n d have a distinctly s e g m e n t e d a p p e a r a n c e (Fig-
ure 10.1 B; R o b i s o n 1987, figure 12.41F). Ulrich n a m e d this w o r m Protoscolex
and d e s c r i b e d four s p e c i e s , all o c c u r r i n g i n t h e E c o n o m y b e d s o f t h e E d e -
n i a n (now t h e K o p e f o r m a t i o n ) . M i l l e r and Faber (1892b) d e s c r i b e d a fifth
s p e c i e s also from the E d e n i a n , from " n e a r the l o w water m a r k " o f the O h i o
River, e q u i v a l e n t to the F u l t o n M e m b e r of the K o p e F o r m a t i o n . A s p e c i m e n
i n the c o l l e c t i o n s o f t h e C i n c i n n a t i M u s e u m C e n t e r i s l a b e l e d from the
400-foot e l e v a t i o n , w h i c h w o u l d p l a c e it in the M a v s v i l l i a n C o r r y v i l l e For-
m a t i o n . A l t h o u g h t h e s e fossils h a v e not b e e n r e s t u d i e d , their identification
as w o r m s has not b e e n c h a l l e n g e d . In the Treatise on Invertebrate Paleontol-
ogy. Protoscolex is listed with w o r m s for w h i c h the phylum is u n c e r t a i n (How-
ell 1962). H o w e v e r . Robison (1987) illustrated a s p e c i m e n from the U p p e r
O r d o v i c i a n of K e n t u c k y as a fossil a n n e l i d .

Cornulites and The c o n i c a l e n c r u s t i n g fossil Gornulites is a c o m m o n C i n c i n n a t i a n m a c r o -

O t h e r Small, fossil of u n c e r t a i n z o o l o g i c a l position but with possible w o r m affinities. C o r -

Conical Fossils nulites o c c u r s t h r o u g h o u t t h e C i n c i n n a t i a n and several s p e c i e s h a v e b e e n

d e s c r i b e d . In t h e Treatise on Invertebrate Paleontology, Fisher (1962) described
Cornulites as s m a l l t u b e s of c a l c i u m c a r b o n a t e , with a c i r c u l a r cross-section
of 2 to 20 mm d i a m e t e r and a l e n g t h of 5 to 80 m m . S m a l l e r Gornulites are
s m o o t h , b u t larger t u b e s h a v e c i r c u m f e r e n t i a l rings and l o n g i t u d i n a l stria-
tions. B e c a u s e t h e p o r o u s or v e s i c u l a r internal w a l l s t r u c t u r e differs from the
m i c r o s t r u c t u r e of t u b e - f o r m i n g a n n e l i d s , Fisher did not assign Gornulites to
a definite p h y l u m . A l t h o u g h t h e n a t u r e o f t h e a n i m a l r e m a i n s u n c e r t a i n ,
Gornulites is s i m i l a r to m o d e r n p o l y c h a e t e s (like serpulids or spirorbids) that
f o r m e n c r u s t i n g t u b e s a n d live as filter feeders. In the C i n c i n n a t i a n , Gornu-
lites u s u a l l y o c c u r s as clusters of t u b e s e n c r u s t i n g a w i d e variety of shelly
substrata s u c h a s b r a c h i o p o d s , b r y o z o a n s , m o l l u s c s , trilobites, and e c h i n o -
d e r m s ( F i g u r e K M C ; see F i g u r e 12.9B), but c a n also o c c u r a s solitary, unat-
t a c h e d t u b e s ( M o r r i s a n d R o l l i n s 1 9 7 1 ; M o r r i s a n d Felton 1993, 2003). T h e
o r i e n t a t i o n of Gornulites w i t h the a p e r t u r e toward the m a r g i n s of b r a c h i o -
p o d s a n d p e l e c y p o d s s u g g e s t s a possible c o m m e n s a l relationship b y w h i c h
the t u b e - d w e l l i n g f i l t e r f e e d e r took a d v a n t a g e o f the f e e d i n g c u r r e n t s o f the
host ( M o r r i s a n d Felton 1993, 2003). M o r e o v e r , c o r n u l i t i d s often e n c r u s t e d
c r i n o i d c o l u m n s a n d shells o f t h e g a s t r o p o d C y c l o n e m a that f r e q u e n t l y at-
t a c h e d to t h e c a l y x of l i v i n g c r i n o i d s (see F i g u r e 12.9B). In this m a i m e r ,
c o r n u l i t i d s g a i n e d a p o t e n t i a l l y a d v a n t a g e o u s f e e d i n g position a b o v e the
s e d i m e n t - w a t e r interlace ( M o r r i s a n d Felton 2003).
T e n t a c u l i t o i d s are s m a l l , c a l c a r e o u s c o n i c a l fossils w i t h c i r c u m f e r e n -
tial rings. T e n t a c u l i t o i d s are s i m i l a r in s i z e a n d form to t h e p r o b a b l e a n -
n e l i d t u b e s c a l l e d Gornulites, r a n g i n g from 1 to 40 mm in l e n g t h (rarely
u p t o 8 0 m m ) . U n l i k e m o s t Gornulites, t e n t a c u l i t o i d s are n o n - e n c r u s t i n g
a n d h a v e a t h i n k l a y e r e d o r l a m e l l a r shell s t r u c t u r e a n d internal i m p e r f o -
rated septa ( B e r g s t r o m 1996b). There is e v i d e n c e of soft part a n a t o m y from
x - r a d i o g r a p h s of a D e v o n i a n t e n t a c u l i t o i d from G e r m a n y that tentacles a n d
a s i p h o n w e r e p r e s e n t ( B l i n d a n d S t u e r m e r 1 9 7 7 ) . This s u g g e s t s a possible
affiliation of t e n t a c u l i t o i d s w i t h c e p h a l o p o d s . In t h e Treatise on Inverte-

144 A Sea without Fish

brate Paleontology, Fisher (1962) p r o p o s e d that tentaculitoids and s o m e
o t h e r s m a l l c o n i c a l shells b e i n c l u d e d i n a n e x t i n c t class C r i c o c o n a r i d a
(meaning "small, ringed cones") b e l o n g i n g to the M o l l u s c a . M o r e recently,
tentaculitoids have b e e n treated as a class u n t o t h e m s e l v e s , T e n t a c u l i -
toidea, and not i n c l u d e d w i t h a n y k n o w n g r o u p ( B e r g s t r o m 1996b). C l u s -
ters of t e n t a c u l i t o i d s are s o m e t i m e s f o u n d on a b e d d i n g p l a n e in p a r a l l e l
a l i g n m e n t probably c a u s e d b y water m o v e m e n t ( F i g u r e 10.1A). T w o s p e c i e s
are k n o w n from the type-Cincinnatian, Tentaculites sterlingensis a n d T.
richmondensis, both from the Richmondian.
A n o t h e r g r o u p of s m a l l , c a l c a r e o u s , c o n i c a l fossils that c a n be f o u n d in
the C i n c i n n a t i a n section are the hyolithids. Hyolithids are m u c h smaller
than either C o r n u l i t e s o r tentaculitids; t h o s e o c c u r r i n g i n t h e C i n c i n n a t i a n
are a b o u t 2 to 3 mm long. U n l i k e Cornulites or t e n t a c u l i t i d s , hyolithids h a v e
a s m o o t h shell with a r o u g h l y t r i a n g u l a r cross-section a n d an o p e r c u l u m or
c a p c l o s i n g the a p e r t u r e . In the Treatise on Invertebrate Paleontology, Fisher
(1962) p l a c e d the hyolithids in the C a l y p t o m a t i d a , r e g a r d e d as an e x t i n c t
class of the P h y l u m M o l l u s c a . A l t h o u g h there is u n c e r t a i n t y as to t h e c o r r e c t
t a x o n o m i c classification of hyolithids, m o s t workers c o n t i n u e to favor affinity
with the m o l l u s c s ( M a l i n k y e t al. 2004). S o m e h y o l i t h i d s m a y h a v e b e e n
similar t o the p r e s e n t - d a y p l a n k t o n i c p t e r o p o d m o l l u s c s , b u t o t h e r s w e r e
probably b e n t h o n i c a n d c a p a b l e o f m o v e m e n t a l o n g t h e sea f l o o r (Fisher
1962). Well-preserved s p e c i m e n s retain the o p e r c u l u m a n d a pair of c u r v e d
spines e m e r g i n g from the a p e r t u r e that c o u l d h a v e b e e n u s e d i n l o c o m o t i o n .
C i n c i n n a t i a n h y o l i t h i d s are k n o w n from t h e M a y s v i l l i a n a n d R i c h m o n d i a n
and i n c l u d e t w o g e n e r a , Hyolithes a n d Coleolus (see D a l v e 1948).

M a c h a e r i d i a n s are a v e r y s m a l l a n d p r o b l e m a t i c g r o u p o f fossils k n o w n Mysterious

from r o c k s o f O r d o v i c i a n t h r o u g h P e r m i a n a g e , a n d are r e p r e s e n t e d i n t h e Machaeridians
Cincinnatian. T h e s e p e c u l i a r fossils are c o m p o s e d o f a series o f o v e r l a p -
p i n g , c a l c a r e o u s s e g m e n t s that a r e e a c h f o r m e d b y a p a i r o f plates ( c a l l e d
sclerites). The p a i r e d plates f o r m a h e a r t - s h a p e d t u b e in c r o s s - s e c t i o n .
M a c h a e r i d i a n s o c c u r e i t h e r a s a r t i c u l a t e d series o f m a n y s e g m e n t s , taper-
i n g at o n e e n d , or as d i s a r t i c u l a t e d sclerites. It is p o s s i b l e that t h e series of
sclerites is not a c o m p l e t e o r g a n i s m , b u t rather is a d i s s o c i a t e d body part.
In a d e t a i l e d study of C i n c i n n a t i a n s p e c i m e n s , John K. P o p e ( 1 9 7 5 ) a r g u e d
that Lepidocoleus is a s h e a t h of plates that c o v e r e d s p i n e s t h a t w e r e at-
t a c h e d to the t h e c a of the c a r p o i d e c h i n o d e r m Enoploura (see c h a p t e r 1 2 ,
F i g u r e 1 2 . 1 7 B ) . Pope's a r g u m e n t w a s b a s e d largely o n t h e f i n d i n g o f a b u n -
dant d i s a r t i c u l a t e d c a r p o i d plates a n d a r t i c u l a t e d series o f m a c h a e r i d i a n
sclerites a t t w o C i n c i n n a t i a n l o c a l i t i e s . M a c h a e r i d i a n s h a d b e e n inter-
preted p r e v i o u s l y a s b e l o n g i n g t o t h e c h i t o n s , b a r n a c l e s , a n n e l i d s , trilo-
bites, or e c h i n o d e r m s . In 2 0 0 4 H i n t s et al. treated m a c h a e r i d i a n s as plates
o f a n u n k n o w n , bilaterally s y m m e t r i c a l w o r m - l i k e o r g a n i s m , b u t i g n o r e d
Pope's c a r p o i d e c h i n o d e r m interpretation. Lepidocoleus jamesi (flail and
W h i t f i e l d ) o c c u r s t h r o u g h o u t t h e C i n c i n n a t i a n ( D a l v e 1948). M a c h a e r i d -
ians are u s u a l l y r e g a r d e d as rare fossils, b u t t h e o c c u r r e n c e s r e p o r t e d by
Pope show that t h e y c a n b e very a b u n d a n t i n s o m e b e d s .

Annelids and Worm-Like Fossils 145

746 A Sea without Fish
 11
ARTHROPODS: TRILOBITES AND
OTHER LEGGED CREATURES

I n t e r m s o f s h e e r a b u n d a n c e , s p e c i e s diversity, a n d e x p l o i t a t i o n o f h a b i t a t s ,
a r t h r o p o d s rank a s t h e m o s t s u c c e s s f u l o f all l i v i n g a n i m a l s . M o r e t h a n
750,000 s p e c i e s (mostly insects) i n h a b i t a cast r a n g e of e n v i r o n m e n t s on
l a n d , i n the sea, a n d i n fresh water. L i v i n g a r t h r o p o d s i n c l u d e t h e i n s e c t s ,
crustaceans, horseshoe crabs, arachnids, centipedes, and millipedes. Dur-
i n g the O r d o v i c i a n , a r t h r o p o d s h a d n o t yet i n v a d e d t h e l a n d , b u t trilobites
w e r e a b u n d a n t a n d d i v e r s e i n the sea, a l o n g w i t h t h e e u r y p t e r i d s , ostra-
c o d e s , a n d a few o t h e r m i n o r g r o u p s .
D e s p i t e their b e w i l d e r i n g variety o f f o r m , all a r t h r o p o d s s h a r e c e r t a i n
basic features. L i k e their c l o s e relatives, t h e a n n e l i d w o r m s , a r t h r o p o d s
have a s e g m e n t e d body. Unlike the a n n e l i d s , t h e b o d y a n d its a p p e n d a g e s
are e n c a s e d in an e x o s k e l e t o n c o m p o s e d of t h e p r o t e i n c h i t i n . The e x o s k e l -
eton is m u c h like a suit of a r m o r in h a v i n g rigid c o m p o n e n t s a r t i c u l a t e d
Figure 11.1. The Ordovi-
b y f l e x i b l e joints. ( T h e n a m e a r t h r o p o d m e a n s " j o i n t e d legs.") N o t o n l y
cian trilobite Triarthrus.
d o e s the e x o s k e l e t o n shield t h e internal o r g a n s f r o m p r e d a t i o n a n d s o m e Left, dorsal view, right,
e n v i r o n m e n t a l h a z a r d s , b u t it also p r o v i d e s rigid p o i n t s for m u s c l e a t t a c h - ventral view. Drawings by
m e n t . C o n s e q u e n t l y , a r t h r o p o d s are c a p a b l e o f rapid l o c o m o t i o n b y w a l k - Kevina Vulinec.
ing, s w i m m i n g , o r f l y i n g . T h e n a t u r e o f t h e e x o s k e l e t o n has t w o i m p o r t a n t
i m p l i c a t i o n s for t h e fossilization p o t e n t i a l o f a r t h r o p o d s . First, b e c a u s e t h e
c h i t i n o u s e x o s k e l e t o n d e c o m p o s e s after d e a t h , m a i n a r t h r o p o d s are p o o r
c a n d i d a t e s for fossil p r e s e r v a t i o n . H o w e v e r , a r t h r o p o d s that h a v e t h i c k e r
e x o s k e l e t o n s o r i n c o r p o r a t i o n o f c a l c i u m c a r b o n a t e into t h e i r s k e l e t o n s
(such a s s o m e c r u s t a c e a n s a n d trilobites) w i l l h a v e e n h a n c e d p o t e n t i a l for
p r e s e r v a t i o n . S e c o n d , all a r t h r o p o d s grow b y p e r i o d i c a l l y s h e d d i n g t h e
e x o s k e l e t o n a n d f o r m i n g a new skin that a c c o m m o d a t e s g r o w t h . E a c h in-
d i v i d u a l a r t h r o p o d c a n c o n t r i b u t e n u m e r o u s s h e d e x o s k e l e t o n s (molts) a s
potential fossils d u r i n g its l i f e t i m e . M o l t i n g may thus e x p l a i n in part t h e
a b u n d a n c e o f s o m e a r t h r o p o d fossils.

A l t h o u g h trilobites a c h i e v e d their m a x i m u m diversity d u r i n g t h e L a t e C a m - Trilobites

brian. T h e y r e m a i n e d diverse and a b u n d a n t d u r i n g the O r d o v i c i a n . A m o n g
the plethora of C i n c i n n a t i a n fossils, trilobites are a r g u a b l e the best k n o w n for
their preservation and a b u n d a n c e . Trilobites are u n i q u e a m o n g the a r t h r o -
p o d s in h a v i n g a characteristic l e n g t h w i s e s u b d i v i s i o n of the dorsal exoskel-
eton into three l o b e s , an axial l o b e flanked by t w o pleural l o b e s ( F i g u r e 11.1).
There is a distinct h e a d shield or c e p h a l o n , a flexibly s e g m e n t e d thorax, a n d
a tail shield or p y g i d i u m . The c e p h a l o n u s u a l l y has a c e n t r a l , s w o l l e n r e g i o n
c a l l e d the g l a b e l l a . A l t h o u g h it r e s e m b l e s a n o s e or f o r e h e a d , the g l a b e l l a

147
 a c t u a l l y p r o t e c t e d the trilobite's s t o m a c h . T r a n s v e r s e glabellar lobes and fur-
rows i n d i c a t e fused s e g m e n t a t i o n of the h e a d r e g i o n . A p r o m i n e n t pair of
c o m p o u n d eyes u s u a l l y flanks the g l a b e l l a . A s i n u o u s facial suture crosses the
cephalon a l o n g s i d e the e y e s , s e p a r a t i n g the lateral free c h e e k s from the fixed
cheeks. The facial s u t u r e p r o v i d e d a l i n e o f b r e a k a g e across t h e c e p h a l o n
d u r i n g m o l t i n g . For this r e a s o n , isolated free c h e c k s are c o m m o n l y f o u n d as
well as the c r a n i d i u m . a single unit c o m p r i s i n g the g l a b e l l a and fixed c h e e k s .
A pair of g e n a l spines is often d e v e l o p e d at the posterior c o r n e r s of the c e p h a -
l o n , as seen in Flexicalymene meeki, Isotelus maximus, and Cryptolithus tes-
sellatus from the C i n c i n n a t i a n .
T h e s e g m e n t s o f t h e t h o r a x w e r e c o n n e c t e d b y a t h i n i n t e g u m e n t that
a l l o w e d t h e trilobite to flex its body a n d in many cases to a c h i e v e c o m p l e t e
e n r o l l m e n t l i k e a m o d e r n p i l l b u g (an i s o p o d c r u s t a c e a n ) . After d e a t h , d e -
c a y o f the a r t i c u l a t i n g i n t e g u m e n t often r e l e a s e d i n d i v i d u a l t h o r a c i c seg-
m e n t s that r e s e m b l e b r a c k e t s ({) w h e n p r e s e r v e d . The p y g i d i u m is c o m -
n i o n l v p r e s e r v e d as a s i n g l e unit b e c a u s e its s e g m e n t s w e r e fused. As a
c o n s e q u e n c e o f m o l t i n g a n d p o s t - m o r t e m d e c a y , t r i l o b i t e f r a g m e n t s are
a b u n d a n t , b u t c o m p l e t e , a r t i c u l a t e d s p e c i m e n s are u n c o m m o n . There i s
c o n s i d e r a b l e d e b a t e a b o u t w h e t h e r c o m p l e t e s p e c i m e n s represent trilobites
buried intact, b e c a u s e s o m e may have molted without the exoskeleton
b r e a k i n g apart. U s u a l l y , h o w e v e r , a r t i c u l a t e d s p e c i m e n s , p a r t i c u l a r l y e n -
rolled o n e s , r e p r e s e n t trilobites b u r i e d alive or very s o o n after d e a t h .
O n t h e v e n t r a l side o f t h e c e p h a l o n , a plate c a l l e d the l a b r u m o r hy-
p o s t o m c w a s p o s i t i o n e d b e n e a t h the m o u t h a n d c o n n e c t e d t o t h e anterior
m a r g i n o f t h e c e p h a l o n ( F i g u r e s 1 1 . 1 , 1 1 . 2 , 11.4B). T h e l a b r u m i s often
f o u n d as an isolated fossil, b u t it rarely will be f o u n d in p l a c e . U n l i k e that
of a c r a b or lobster, t h e ventral e x o s k e l e t o n of trilobites was a t h i n , c h i t i n o u s
m e m b r a n e to w h i c h the appendages were attached. Trilobite appendages
were also weakly constructed and thus were preserved only under excep-
tional c o n d i t i o n s ( F i g u r e 1 1 . 4 D ) . A pair of jointed a n t e n n a e w a s a t t a c h e d
to the ventral side of the c e p h a l o n , f o l l o w e d by a series of p a i r e d , jointed
a p p e n d a g e s u n d e r l y i n g e a c h s e g m e n t o f the c e p h a l o n , thorax and pygid-
i u m . A l t h o u g h a p p e n d a g e s are k n o w n from v e r y few s p e c i e s o f trilobites,
t h e a p p e n d a g e s are s i m i l a r i n h a v i n g t w o b r a n c h e s : t h e w a l k i n g l e g o r
e n d o p o d i t e a n d a b r a n c h c a l l e d t h e e x o p o d i t e e x t e n d i n g from t h e basal
s e g m e n t o f t h e e n d o p o d i t e ( F i g u r e s 1 1 . 1 , 11.2). T h e e x o p o d i t e c a r r i e d n u -
m e r o u s filaments g i v i n g it a c o m b - l i k e a p p e a r a n c e .
A l t h o u g h at least sixteen g e n e r a of trilobites are k n o w n from the C i n c i n -
n a t i a n , o n l y a few are c o m m o n . Flexicalymene and Isotelus are the m o s t
c o m m o n and are distributed t h r o u g h o u t the C i n c i n n a t i a n Series. The w i d e -
spread distribution of t h e s e t w o s i g n a t u r e trilobites, in shales a n d l i m e s t o n e s
r e p r e s e n t i n g t h e full r a n g e o f C i n c i n n a t i a n d e p o s i t i o n a l e n v i r o n m e n t s ,
clearly s u g g e s t s that b o t h h a d very g e n e r a l i z e d habitat p r e f e r e n c e s . In c o n -
trast, m o s t o t h e r C i n c i n n a t i a n trilobites h a v e m u c h m o r e restricted strati-
g r a p h i c distributions, s u g g e s t i n g m o r e l i m i t e d e n v i r o n m e n t a l tolerances.
Flexicalymene is o n e of the world's b e s t - k n o w n trilobites, in large m e a -
sure d u e to its a b u n d a n c e in C i n c i n n a t i a n strata ( F i g u r e 11.3). A l t h o u g h a
m o d e r n systematic review has not b e e n d o n e , as m a n y as three species may

148 A Sea without Fish

 Figure 1 1 . 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-
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.

be present: F. meeki (Foerste), by far t h e m o s t a b u n d a n t a n d w i d e l y distrib-

uted species, F. granulosa (Foerste), a small form restricted to t h e l o w e r C i n -
c i n n a t i a n K o p e F o r m a t i o n , a n d F. retrorsa (Foerste), f o u n d o n l y in t h e u p p e r
Cincinnatian Waynesville F o r m a t i o n . In the first v o l u m e of the Geological
Survey of Ohio (1873), F. B. M e e k i n c l u d e d t h e C i n c i n n a t i a n c a l y m e n i d
trilobites with Calymene senaria, originally described from N e w York by
C o n r a d in 1841. Foerste (1910) p r o p o s e d t h e n a m e Calymene meeki for the

Arthropods 149
 s p e c i e s so well d e s c r i b e d by M e e k from the C i n c i n n a t i a n rocks of O h i o , but
p r o v i d e d o n l y a five-line description and a single illustration of an e n r o l l e d
s p e c i m e n . In the s a m e p a p e r Foerste n a m e d C. meeki-retrorsa, a form of
C a l y m e n e meeki from the Waynesville, w h i c h differs chiefly in t h e n a r r o w e r
posterior w i d t h o f the c e p h a l o n , r e s u l t i n g i n m o r e o b t u s e genal a n g l e s . T h e
anterior border of the c e p h a l o n is m o r e strongly reflexed, b r i n g i n g it closer
to the anterior m a r g i n of the g l a b e l l a . T h e British worker Shirley (1936)
p l a c e d t h e C i n c i n n a t i a n s p e c i e s in Flexicalymene. Ross (1967) provided a
m o r e c o m p l e t e d e s c r i p t i o n of F. meeki a n d also r e v i e w e d the status of Foer-
ste's C. meeki-retrorsa. Ross c o n s i d e r e d F. retrorsa to be a valid species al-
t h o u g h it has little to d i s t i n g u i s h it from meeki except the size, s h a p e , and

750 A Sea without Fish

inclination of the anterior c r a n i d i a l border. Ross was u n a b l e to verify t h e Figure 11.3. A, B. Flexicalymene meeki
other differences asserted by Foerste. A m o r p h o m e t r i c analysis c o n d u c t e d by
D a n i t a Brandt (1980) in her u n p u b l i s h e d master's thesis led h e r to c o n c l u d e (Foerste), University of
Cincinnati collections,
that only a single species, F. meeki, is valid, a n d that b o t h F. granulosa and
Maysvillian, Corryville
F. retrorsa s h o u l d be s y n o n y m i z e d w i t h F. meeki as intraspecific variants.
Formation, Hamilton Co.,
M o r e recent work b y B r e n d a H u n d a (pers. c o m m . ) supports t h e r e c o g n i t i o n
Ohio, enrolled specimen,
of F. meeki, F. retrorsa, and F. granulosa as valid s p e c i e s . cephalic width 28 mm.
Flexicalymene is c o m m o n l y f o u n d as isolated p a r t i a l e x o s k e l e t o n s (cc- C. Flexicalymene ret-
p h a l a , c r a n i d i a , free c h e e k s , t h o r a c i c s e g m e n t s , p y g i d i a ) i n C i n c i n n a t i a n rorsa (Foerste), CMC IP,
l i m e s t o n e s ; c o m p l e t e s p e c i m e n s are less c o m m o n a n d a r e u s u a l l y f o u n d i n Ferree Collection, Rich-
mondian, Arnheim For-
shales a s either e n r o l l e d o r e x t e n d e d i n d i v i d u a l s . O n rare o c c a s i o n s , t h e s e
mation, Highland Co.,
trilobites c a n be f o u n d in g r e a t n u m b e r s in yellowish s h a l e s k n o w n as but-
Ohio, x 1.4. D. Flexicalymene granulosa
ter shales. O n e o f the m o s t prolific trilobite d i s c o v e r i e s e v e r m a d e i n t h e
(Foerste), Mark Peter col-
C i n c i n n a t i a n was in such a shale within the lower R i c h m o n d i a n W a y n e s - lection, Edenian, Kope
ville 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 a n 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 Formation, Brown Co.,
Avenue and W e s t w o o d Northern Boulevard in northwest C i n c i n n a t i in the Ohio, x 2.5.
1950s. Literally t h o u s a n d s o f F l e x i c a l y m e n e w e r e c o l l e c t e d f r o m t w o shale-
b e d s 2.5 a n d 3 feet t h i c k , a n d as t h e e x p o s u r e w e a t h e r e d , trilobites b e c a m e
p e r c h e d on pedestals of cla\ for eas\ p i c k i n g ( C a s t e r , pers, c o m m . ; S c h w e i n -
furth 1958). T a p h o n o m i c s t u d i e s o f o c c u r r e n c e s o f a b u n d a n t , c o m p l e t e
trilobites i n C i n c i n n a t i a n s h a l e s i n d i c a t e that t h e s e a r e t h e result o f m a s s
m o r t a l i t i e s o f l i v i n g trilobite p o p u l a t i o n s s m o t h e r e d d u r i n g s t o r m s o r m a s s
m o v e m e n t s of hue-grained sediments (Brandt 1980, 1985; S c h u m a c h e r
a n d S h r a k e 1997; H u g h e s a n d C o o p e r 1999).

Life Habits of Flexicalymene

O u r u n d e r s t a n d i n g o f t h e life habits o f trilobites has b e e n h a m p e r e d b y t h e

fact that trilobites are e x t i n c t a n d h a v e n o c l o s e l i v i n g relatives, a l t h o u g h
h o r s e s h o e crabs a n d s o m e c r u s t a c e a n s are often r e g a r d e d a s p o s s i b l e m o d -
els (Plate 3 G ) . T h e f o r m o f t h e a p p e n d a g e s i s c l o s e l y related t o life habits
in l i v i n g a r t h r o p o d s , but in trilobites t h e a p p e n d a g e s are so rarely p r e s e r v e d
that little i n f o r m a t i o n c a n be g a i n e d from t h e m . A r e c e n t r e v i e w by Fortey
and O w e n s (1999) d e m o n s t r a t e d that o t h e r p r e s e r v e d m o r p h o l o g i c a l fea-
tures o f trilobites c a n b e u s e d t o d e t e r m i n e their f e e d i n g habits. Fortey a n d
O w e n s regard t h e calymenid trilobites like F l e x i c a l y m e n e t o h a v e b e e n
predators or s c a v e n g e r s b e c a u s e the h y p o s t o m e is rigidly a t t a c h e d to t h e
underturned edge of the c e p h a l o n (doublure). T h e hypostome may have
acted as a g r i n d i n g or m a n i p u l a t i n g s u r f a c e for s m a l l prey i t e m s h e l d b e -
t w e e n t h e basal s e g m e n t s o f t h e a p p e n d a g e s .
F u r t h e r e v i d e n c e for the predator) behavior of Flexicalymene c o m e s
from characteristic b u r r o w s (trace fossils n a m e d Rusophycus) f o r m e d by this
trilobite. Rusophycus trace fossils are well k n o w n in strata of late P r e c a m -
brian t h r o u g h D e v o n i a n a g e s , and m o s t are t h o u g h t t o h a v e b e e n p r o d u c e d
by trilobites d i g g i n g into the s e d i m e n t u s i n g the paired a p p e n d a g e s ( H a n t z -
schcl 1975). A l t h o u g h the d i g g i n g activity c o u l d reflect different possible-
behaviors i n c l u d i n g sheltering, resting, e g g l a y i n g , o r f e e d i n g , r e c e n t discov-
eries suggest that trilobites w e r e actively h u n t i n g prey b u r i e d w i t h i n the sedi-

Arthropods 151
 ment. A l t h o u g h by no m e a n s c o m m o n , several r e m a r k a b l e cases of Rusophy-
cus i n t e r s e c t i n g w o r m b u r r o w s a r e k n o w n from the C a m b r i a n (Jensen 1990)
a n d C i n c i n n a t i a n ( B r a n d t et al. 1995). In t h e C i n c i n n a t i a n , Flexicalymene
is u n e q u i v o c a l l y identified as the p r o d u c e r of o n e t y p e of Rusophycus (R.
puclicum) on the basis of a few e x c e e d i n g l y rare s p e c i m e n s that have the
characteristic bilobate burrow preserved in p l a c e b e n e a t h the c o m p l e t e cara-

152 A Sea without Fish

p a c e o f the trilobite (see F i g u r e 14.2B; O s g o o d 1970). A l t h o u g h n o s p e c i - Figure 1 1 . 4 . I s o t e l u s
m e n s of Rusophycus p u d i c u m from the C i n c i n n a t i a n h a v e b e e n f o u n d inter- maximus Locke. A. En-

s e c t i n g w o r m b u r r o w s , the d i g g i n g activity i s c o n s i s t e n t w i t h p r e d a t i o n o n rolled specimen, CMC IP

2250, Riehmondian, Arn-
small, infaunal o r g a n i s m s (Fortey a n d O w e n s 1999).
heim Formation, High-
D e s p i t e the e x c e l l e n t state of preservation f o u n d in C i n c i n n a t i a n Flexi-
land Co., Ohio, x 1.3.
calymene, r e m n a n t s o f the a p p e n d a g e s h a v e n e v e r b e e n f o u n d . S t u r m e r a n d
B. Large hypostome,
B e r g s t r o m (1973) carried out x-radiographic studies that r e v e a l e d p r e s e r v e d CMC IP 33067, Maysvil-
a p p e n d a g e s in s o m e trilobites, b u t s i m i l a r studies by B r a n d t (1980) a n d by lian, Clermont Co., Ohio,
H u g h e s and C o o p e r (1999) d e t e c t e d n o e v i d e n c e o f a p p e n d a g e s i n Flexica- x 7. C. CMC IP 51,
lymene. The study by H u g h e s and C o o p e r revealed pyritized material c o n - Robert Nestor collection,
Maysvillian, Corryville
centrated w i t h i n the b o d y cavity of Flexicalymene that m a y h a v e o r i g i n a t e d
Formation, Clermont Co.,
as d e c a y i n g soft parts. An a p p r o x i m a t e idea of the n a t u r e of the a p p e n d a g e s
Ohio, x 1.8. D. Ap-
in C i n c i n n a t i a n Flexicalymene s p e c i e s c a n be g a i n e d f r o m t h e restoration of
pendages on ventral side
the closely related F. senaria ( f i g u r e 1 1 . 2 A ; R a y m o n d 1920). of a complete specimen,
Isotelus i s t h e o t h e r h i g h l y c h a r a c t e r i s t i c a n d w i d e l y d i s t r i b u t e d t r i l o - USNM 33458, Riehmon-
bite of t h e 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 tri- dian, Oxford, Butler Co.,
lobite are f o u n d i n every C i n c i n n a t i a n f o r m a t i o n , i n b o t h l i m e s t o n e s a n d Ohio, x 0.75. This excep-

shales. C o m p l e t e s p e c i m e n s are q u i t e rare, b u t i n c e r t a i n s h a l e h o r i z o n s , tional specimen was orig-

inally illustrated by Mick-
p a r t i c u l a r l y i n the W a y n e s v i l l e F o r m a t i o n , n u m e r o u s c o m p l e t e s p e c i m e n s
leborough (1883). Photo
have b e e n f o u n d ( S c h u m a c h e r a n d S h r a k e 1 9 9 7 ) . S p e c i m e n s o f Isotelus
courtesy of Loren
from the C i n c i n n a t i a n arc a m o n g t h e l a r g e s t - k n o w n trilobites. A s p e c i m e n
Babcock.
of Isotelus on e x h i b i t at t h e S m i t h s o n i a n I n s t i t u t i o n c o l l e c t e d in 1919 dur-
i n g c o n s t r u c t i o n o f the H u f f m a n D a m n e a r D a y t o n m e a s u r e s 3 7 c m l o n g
(14.5 in) b y 2 6 c m w i d e (10.25 in). T h e c o m p l e t e s p e c i m e n a t C i n c i n n a t i
M u s e u m C e n t e r m e a s u r e s 37.5 c m i n l e n g t h (Plate 7 ) . O n t h e basis o f
partial s p e c i m e n s , Isotelus p r o b a b l y r e a c h e d l e n g t h s o f 8 0 - 9 0 c m ( B a b -
c o c k , pers. c o m m . ) . R e c e n t l y , a s p e c i m e n o f Isotelus w a s f o u n d i n U p p e r
O r d o v i c i a n strata i n n o r t h e r n M a n i t o b a that h o l d s t h e c u r r e n t w o r l d r e c o r d
as the largest trilobite, at a l e n g t h of o v e r 70 cm ( R u d k i n et al. 2003).
C u r r e n t l y , t w o s p e c i e s o f Isotelus a r e r e c o g n i z e d i n t h e C i n c i n n a t i a n :
I. maximus and I. gigas. I. maximus has w e l l - d e v e l o p e d g e n a l s p i n e s . I. gi-
gas has g e n a l s p i n e s that are e i t h e r shorter t h a n t h o s e of I. maximus or
l a c k i n g . W i t h o u t a m o d e r n , critical a n a l y s i s o f t h e r a n g e o f v a r i a t i o n w i t h i n
these s p e c i e s , it is not c l e a r h o w to s e p a r a t e I. maximus f r o m I. gigas on t h e
basis of g e n a l s p i n e l e n g t h . C i n c i n n a t i a n Isotelus h a v i n g a very b r o a d , flat-
tened c a r a p a c e w e r e g i v e n the n a m e I. brachycephalus by F o e r s t e (1919),
but this s p e c i e s w a s r e g a r d e d as a v a r i a n t of I. maximus by B a b c o c k ( 1 9 9 6 ) ,
and h e n c e a j u n i o r s y n o n y m of I. maximus. (If t w o d i f f e r e n t n a m e s h a v e
b e e n g i v e n t o the s a m e s p e c i e s b y d i f f e r e n t w o r k e r s , o n e n a m e c a n b e
s u b o r d i n a t e d as a j u n i o r anonym if it w a s d e s c r i b e d after t h e o r i g i n a l
n a m e a n d if it is d e t e r m i n e d to be e q u i v a l e n t to t h e o r i g i n a l l y d e s c r i b e d
taxon.) B e c a u s e I. maximus a n d his I. brachycephalus o c c u r t o g e t h e r in t h e
R i e h m o n d i a n strata o f t h e C i n c i n n a t i a n , Foerste s u g g e s t e d t h e possibility
that the b r o a d e r I. B r a c h y c e p h a l u s m i g h t r e p r e s e n t f e m a l e s a n d t h e nar-
rower I . m a x i m u s the m a l e s o f a s i n g l e s p e c i e s . This r e m a i n s a n i n t r i g u i n g
possibility that has n e v e r b e e n fully e x p l o r e d .

Anhropods 153
 Life Habits o f Isotelus

Isotelus w a s o n e of t h e l a r g e s t - k n o w n a n i m a l s in the C i n c i n n a t i a n sea, ri-

v a l e d o n l y by t h e less c o m m o n euryptcrid Megulograptus and endocerid
c e p h a l o p o d s . By v i r t u e o f its l a r g e s i z e a l o n e , Isotelus m i g h t b e suspected
to h a v e b e e n a predator, b u t a d d i t i o n a l e v i d e n c e also p o i n t s clearly toward
this i n t e r p r e t a t i o n of its e c o l o g i c a l role. L i k e Flexicalymene, Isotelus has a
h y p o s t o m e rigidly a t t a c h e d t o the c e p h a l i c d o u b l u r e ( F i g u r e s 1 1 . 2 B , 11.4B);
i n a d d i t i o n t h e a n t e r i o r c e p h a l i c m a r g i n i s s t r e n g t h e n e d ( F o r t e y and O w -

154 A Sea without Fish

e n s 1999). Fortey a n d O w e n s m e n t i o n e d several o t h e r u n i q u e features o f Figure 11.5. A. Deco-
the Isotelus h y p o s t o m e that s u g g e s t its f u n c t i o n as a rigid p l a t f o r m like an roproetus parviusculus

anvil for the m a n i p u l a t i o n of b u l k y f o o d : its forked s h a p e , a n d d e v e l o p m e n t (Hall), CMC IP 46429,

Edenian (figured in Davis
of anterior w i n g s p r o v i d e a larger s u r f a c e a r e a , a n d the fine raised ridges
[1992, plate 2, figure 23]
on the i n n e r s u r f a c e s of t h e fork c o u l d m a k e it easier to h o l d t h e prey rigidly
as Proetus parviuscu-
u s i n g t h e a p p e n d a g e s . This h y p o s t o m e i s t h e m o s t h e a v i l y c a l c i f i e d part o f
lus), x 7.7. B. Triar-
the Isotelus exoskeleton and is o f t e n found as a n isolated c o m p o n e n t thrus eatoni (Hall), Steve
( F i g u r e 11.4B). Brown collection, J. Rush
In the C i n c i n n a t i a n , large Rusophycus b u r r o w s h a v e b e e n a t t r i b u t e d collector, Edenian, Kope
to Isotelus on the basis of their s i z e (R. carleyi, see O s g o o d 1970). S p e c i - Formation, Hamilton Co.,

m e n s often show not o n l y f u r r o w s c r e a t e d b y t h e a p p e n d a g e s , b u t a l s o Ohio, x 4.6. C. Cryp-

tolithus tessellatus
impressions o f the c e p h a l i c a n d p y g i d i a l m a r g i n s a n d p l e u r a e . A r e m a r k -
Green, University of Cin-
able s p e c i m e n s h o w s a h o r i z o n t a l w o r m burrow a p p a r e n t l y t r u n c a t e d i n
cinnati collections, Ede-
the a p p r o x i m a t e l o c a t i o n o f t r i l o b i t e s m o u t h (see F i g u r e 14.2A; Brandt e t
nian, Kope Formation,
al. 1 9 9 5 ) a trace fossil r e c o r d i n g the v e r y act o f p r e d a t i o n . T h e trilobite Hamilton Co., Ohio, x
e v i d e n t l y d u g a n d drew itself d o w n into a s e m i - c o h e s i v e m u d s u b s t r a t u m 3.7.
so as to impress the m a r g i n s of its c a r a p a c e l i k e a c o o k i e cutter. T h e t r a c e
s h o w s i m p r e s s i o n s o f t h e basal s e g m e n t s (coxae) o f the a p p e n d a g e s that
probably s e i z e d the prey a l o n g the v e n t r a l m i d l i n e a n d w o r k e d it toward
t h e m o u t h . A s i n g l e e x c e p t i o n a l s p e c i m e n p r e s e r v i n g the a p p e n d a g e s o f
Isotelus was f o u n d in t h e C i n c i n n a t i a n n e a r O x f o r d , O h i o , a n d w a s first
r e p o r t e d b y M i c k l e b o r o u g h (1883) ( F i g u r e 11.41D). O n l y t h e w a l k i n g l e g s
are p o o r l y p r e s e r v e d , but the large s i z e of the c o x a e is e v i d e n t .

O t h e r Trilobites

Cryptolithus, the l a c e - c o l l a r e d trilobite, is a n o t h e r c o m m o n C i n c i n n a t i a n

trilobite in the Edenian a n d M a v s v i l l i a n f o r m a t i o n s ; a s i n g l e s p e c i e s , C.
tessellatus, is present ( F i g u r e 1 1 . 5 C ) . Its c o m m o n n a m e refers to the b r o a d ,
perforated c e p h a l o n that bears g e n a l spines. The t h o r a x a n d p y g i d i u m form
a s m a l l , short unit that is rarely f o u n d a t t a c h e d to the c e p h a l o n . E x c e p t i o n -
ally w e l l - p r e s e r v e d s p e c i m e n s from t h e O r d o v i c i a n o f N e w York r e v e a l e d
the a p p e n d a g e s of Cryptolithus. R e c o n s t r u c t i o n of t h e a p p e n d a g e s s h o w s
that e a c h w a l k i n g leg was a c c o m p a n i e d by a b r a n c h (exopodite) that c a r r i e d
long, comb-like filaments (Figure 11.2C). The e x o p o d i t e s p r o b a b l y w a v e d
in u n i s o n to stir up the s e d i m e n t and create a respiratory a n d f e e d i n g cur-
rent. Trace fossils attributable to Cryptolithus (Rusophycus cryptolithi) s u g -
gest that this trilobite e x c a v a t e d a pit that c r e a t e d a filter-feeding c h a m b e r
b e n e a t h the broad c e p h a l o n ( O s g o o d 1970; F o r t e y and O w e n s 1999). Dif-
ferent workers a g r e e that Cryptolithus u s e d the a p p e n d a g e s to e x t r a c t food
particles from the s e d i m e n t stirred up in d i g g i n g the pit, b u t t h e r e are vary-
i n g interpretations of the f u n c t i o n of the p e r f o r a t i o n s of the c e p h a l i c fringe.
The tunnel-like perforations c o u l d h a v e a c t e d as a sieve to s e p a r a t e fine
particles from a f e e d i n g c u r r e n t p a s s i n g from the exterior of t h e c e p h a l o n
to the interior ( C i s n e 1970; S c i l a c h e r 1970). A c c o r d i n g to Fortey a n d O w e n s
(1999), it is m o r e r e a s o n a b l e to s u p p o s e that a f e e d i n g c u r r e n t c r e a t e d by the
e x o p o d i t e s (gills) stirred up food p a r t i c l e s b e n e a t h t h e trilobite, t h e n exited
t h r o u g h the perforations from t h e interior t o t h e exterior. C a m p b e l l ( 1 9 7 5 ) ,

Arthropods 155
Figure 11.6. A. Acidaspis cincinnatiensis Meek, Steve Brown collection, J. Rush collector, x 2.6.
B, D. Odontopleurid, gen. and sp. Undetermined. B. MUGM 29056, Richmondian, Oxford, Butler Co.,
Ohio, x 4.0. D, Steve Brown collection, J. Rush collector, x 2.6. C. Primaspis crosotus (Locke), University
of Cincinnati collections, Kope Formation, Hamilton Co., Ohio, x 6. E. Primaspis crosotus (Locke) on
bryozoan Peronopora sp., MUGM 4001-A, x 2.8, from Shrake (1989, plate 3C). Note Catellocaula val-
lata bioclaustrations in upper right part of bryozoan. F. Primaspis crosotus (Locke) on bryozoan, MUGM
4010, ventral side of trilobite showing hypostome in place, x 5.2, from Shrake (1989, plate 4B).

156 A Sea without Fish

Figure 11.7. Cincinnatian phacopid trilobites. A. Platycoryphe christyi (Hall), Mark Peter collection, Rieh-
mondian, Waynesville Formation, Montgomery Co., Ohio, x 2.6. B. Tricopelta breviceps (Hall), MUGM
29057, Riehmondian, Waynesville Formation, Franklin Co., Indiana, x 5.5. C. Ceraurus milleranus Miller
and Gurley, CMC IP 5199, enrolled, no unit or locality (figured in Davis [1992, plate 3, figure 28]), x 2.5.
D. Ceraurinus icarus (Billings), Steve Brown collection, J. Rush collector, Riehmondian, x 2.

Arthropods 157
Figure 11.8. A, C from Caster and Kjellesvig-Waering (1964, plate 45, figure 1, plate 46, figure 4) and re-
printed by permission of the Paleontological Research Institution. A. Eurypterid Megalograptus ohioen-
sis Caster and Kjellesvig-Waering, Richmondian, Elkhorn Formation, Adams Co., Ohio; ventral side of post-
abdomen and last (sixth) pair of legs (darker segments), dorsal imprint of prosoma (grey segments turned
to right near top), holotype, CMC IP 24119A, x 0.14. B. Professor Kenneth E. Caster with type specimens
of Megalograptus ohioensis. C. First walking leg, paratype, CMC IP 24117A, x 0.9. D. Aglaspid Neo-
strabops martini Caster and Macke, holotype, CMC IP 25569, Maysvillian, Corryville Formation, Clermont
Co., Ohio, x 1.4. From Caster and Macke (1952, plate 109, figure 2), and reprinted by permission of the
Society for Sedimentary Geology.

158 A Sea without Fish

GALLERY
Plate 1. Ordovician continents and oceans. The reconstruction shown is for the Middle Ordovician, about 4 5 8 million years a g o ,
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 w a s south of the equator. Map courtesy of C. R. Scotese, PALEOMAP Project (www.scotese.com).
Plate 2. Late Ordovician continents and oceans. This reconstruction is
for the youngest t w o stages of the Ordovician, the Ashgillian and Hirnan-
tian. This timeslice includes the Cincinnatian interval. Dots indicate sites
where reefs and related deposits are known for these stages, and colors
indicate the dominant groups contributing to these deposits. Tan = land
areas, light blue = epicontinental seas, and darker blue = d e e p oceans.
Map from Kiessling et al. (2003), and courtesy of Wolfgang Kiessling.

Plate 3. Living invertebrates related to type-Cincinnatian fossil groups. All

photos by D. L. Meyer except as noted. A. Sponge, Pericharax sp., Lizard Island,
Great Barrier Reef, Australia. Green fluorescent dye w a s released at base of
sponge, taken into sponge's filtration system, and ejected as stream from oscu-
lum at top, as a demonstration of the sponge's filter-feeding capability.
B. Coralline sponge, Acanthochaetetes wellsi Hartman and Goreau, submarine
cave at 18 m depth, Palau Islands. Diameter 8 cm. Sponge tissue is a thin veneer
over a calcareous skeleton. Coralline sponges are similar to Ordovician stromato-
poroids. C. Scleractinian coral, Tubastrea coccinea Lesson, showing tentacle-
bearing polyps, Curacao, Netherlands Antilles. Photo by William K. Sacco. D.
Bryozoan, Heteropora pacifica Borg, San Juan Islands, Washington, depth 24 m.
Colony form very similar to some Ordovician bryozoans. E. Bryozoan, unidenti-
fied cheilostome, Lizard Island, Great Barrier Reef. Australia, depth about 18 m.
Note extended tentacles along funnel-shaped section at center. Funnel-shaped
section will act as a "chimney" to direct water away from the colony after
it has been filtered by the tentacles. F. Brachiopod, Thecidellina, Curacao,
Netherlands Antilles. These brachiopods are cemented by the pedicle valve to
the undersides of coral colonies. Note 90 degree g a p e of valves and extended
tentacles of the lophophore. Width about 4 mm. Photo by William K. Sacco.
G. Horseshoe crab, Family Limulidae, Subfamily Tachypleinae, Mersing, Malaysia.
Plate 4. Nautilus, the only living cephalopod with an external shell. Photographs by Richard
Arnold Davis. A., B. Nautilus macromphalus, specimen trapped at Lifou and photographed
in aquarium at Noumea, New Caledonia. A. Note the hyponome, the circular opening below
the head. B. Note countershading pattern in which stripes are confined to upper side of
shell. C. Nautilus pompilius, cross-section of shell, showing chambers and siphuncular tube.
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. C o n o -
dont, one element of an
apparatus, Phragmodus
undatus Branson and
Mehl, CMC IP 50705,
Kope Formation, Campbell
Co., Kentucky, x 227.

Plate 6. Allolichas halli

(Foerste), Dan Cooper
collection, Richmondian,
Waynesville Formation,
Franklin Co., Indiana.
This exceptionally rare,
complete specimen
shows a darker coloration
around the margin of the
carapace that may be a
remnant of original col-
oration. This species w a s
assigned to Amphilichas
until 2002 w h e n Holloway
and Thomas placed it in
Allolichas. Scale in mm.
Plate 7. Isotelus maximus Locke,
CMC IP 50168, Richmondian,
Adams Co., Ohio. This exceptional
complete specimen is 37.5 cm
in length and w a s collected
and prepared by Thomas T.
Johnson. Isotelus is the State
Invertebrate Fossil of Ohio.
Plate 8. A. Flexicalymene retrorsa on internal mold
of nautiloid, CMC IP 50844, Arnheim Formation,
Thomas Weaver Collection. Trilobite is located
on phragmocone (chambered) section of the
nautiloid mold, with head toward body chamber.
Scale in mm. B. Parallel alignment of orthoconic
nautiloids, C M C IP, Cincinnatian. Scale in mm. Note
that not all specimens on the slab are aligned.
Plate 9. Living echinoderms. 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 - 1 0 mm.
D. Echinoid, Strongylocentrotus franciscanus, San Juan Islands, Washing-
ton, diameter about 15 cm. E. Asteroid (sea star), Fromia nodosa Clark,
Seychelles, Indian Ocean, arm length about 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 10. Cincinnatian disparid crinoids. A. locrinus subcrassus (Meek and
Worthen), C M C IP 46012, Waynesville Formation, Franklin Co., Indiana;
note large crown size and dense packing of crinoidal material. B. "Logjam"
of Ectenocrinus simplex (Hall), C M C IP, Fairview Formation, Hamilton Co.,
Ohio; note parallel alignment of crinoids.
Plate 11. Cincinnatian cladid crinoids. A. Cupulocrinus polydactylus (Shumard), MUGM 28344,
Oxford area, Butler Co., Ohio, six crowns, XI. B. Plicodendrocrinus casei(Meek), M U G M 28353,
Liberty Formation, southwestern Ohio, t w o crowns, one overlaps the cup of the other, X1.3.
Plate 12. Late Ordovician paleogeography of the eastern United States.
Light blue = shallowest water over Lexington Platform,
medium blue = shallow subtidal depths, dark blue = deeper subtidal depths,
buff = Queenston Delta, brown = Taconic highlands, the eventual loca-
tion of the Appalachian Mountains. Courtesy of Steven M. Holland.

Plate 1 3 . Dioramas of the Cincinnatian sea. A. Cincinnati Museum of

Natural History, by Paul Marchand, about 1959, under direction of Ken-
neth E. Caster. B. Museum of Comparative Zoology, Harvard University.
Plate 14. The Cincinnatian, by John A g n e w , 2007.
Figure 11.9. Eurypterid Megalograptus ohioensis Caster and Kjellesvig-Waering. C, D. From Caster and
Kjellesvig-Waering (1964, plate 49, figure 2, plate 48, figure 1). A. Reconstruction of dorsal surface of
adult female (from Caster and Kjellesvig-Waering [1964, plate 43]), x 0.14. B. Reconstruction of ventral
surface of adult female (from Caster and Kjellesvig-Waering [1964, plate 44]), x 0.14. C. Distal end of
second walking leg, note bulbous expansion (sensory?) at end of longest spine, paratype, CMC IP 24115, x
0.7. D. Third paired appendage, showing large coxa and part of first four joints, paratype, CMC IP 24103A.
x 0.4. A-D reprinted by permission of the Paleontological Research Institution.

Arthropods 159
 h o w e v e r , p o i n t e d o u t that very little f l o w c o u l d h a v e passed t h r o u g h the
m i n u t e p o r e s a n d t h u s it is m o r e likely that t h e pores h a d a sensory f u n c t i o n
to orient the a n i m a l into the c u r r e n t . It this is true, the pores may h a v e b e e n
the sites of s e n s o r y hairs that w e r e not p r e s e r v e d .
Triarthrus is restricted to the l o w e r m o s t K o p e f o r m a t i o n b u t is signifi-
c a n t for several r e a s o n s ( f i g u r e 11.5B). Triarthrus is t h e last of the o l e n i d
trilobites t h a t w e r e p r o m i n e n t d u r i n g t h e C a m b r i a n . P y r i t i z e d s p e c i m e n s
f r o m t h e U p p e r O r d o v i c i a n U t i c a S h a l e o f N e w York a r e a m o n g t h e m o s t
w e l l - p r e s e r v e d trilobites, f r o m w h i c h d e t a i l e d r e c o n s t r u c t i o n s o f the ap-
p e n d a g e s a n d i n t e r n a l soft a n a t o m y h a v e b e e n m a d e ( C i s n e 1970; W l i i t -
t i n g t o n a n d A l m o n d 1987). In t h e C i n c i n n a t i a n , t w o s p e c i e s , T. eatoni a n d
T. spinosus, are r e c o g n i z e d ( B a b c o c k 1996a) but p r e s e r v e d a p p e n d a g e s
h a v e n o t b e e n f o u n d . T h e s t r u c t u r e o f t h e a p p e n d a g e s i n Triarthrus sug-
gests t h a t it w a s a p a r t i c l e f e e d e r , s o r t i n g food p a r t i c l e s w i t h the t h o r a c i c
a p p e n d a g e s , w h i c h t h e n p a s s e d t h e m t o w a r d the m o u t h a l o n g the ventral
axis ( F o r t e y a n d O w e n s 1999). E n l a r g e d , s p i n e - b e a r i n g basal l i m b seg-
m e n t s ( g n a t h o b a s e s ) b e n e a t h t h e c e p h a l o n a c t e d like jaws t o p r o c e s s t h e
f o o d a n d transfer it to the m o u t h . T h e restriction of Triarthrus to the dark
s h a l e s o f t h e l o w e r K o p e f o r m a t i o n , a n d its d o m i n a n c e i n s o m e t h i n b e d s ,
b o t h s u g g e s t that this trilobite w a s u n i q u e l y a d a p t e d t o d e e p e r water e n v i -
r o n m e n t s low i n o x y g e n , a s w e r e o t h e r o l e n i d trilobites (Fortey and O w e n s
1999).
A m o n g C i n c i n n a t i a n trilobites, t w o g e n e r a b e l o n g i n g t o the O r d e r O d -
o n t o p l e u r i d a are u n i q u e in b e i n g f e s t o o n e d with spines and tubercles; these
are Acidaspis and Primaspis ( F i g u r e 11.6). Acidaspis is d i s t i n g u i s h e d by hav-
i n g a l o n g spine o r i g i n a t i n g on the occipital l o b e of the c e p h a l o n and e x t e n d -
i n g o v e r t h e axial l o b e of the t h o r a x , as well as l o n g g e n a l spines and a fringe
of short spines a l o n g the m a r g i n of the free c h e c k s ( F i g u r e 11.6A). A c o m -
plete d e s c r i p t i o n of A. cincinnatiensis M e e k is given by W h i t t i n g t o n (1956).
Primaspis lacks t h e o c c i p i t a l spine a n d has t h e c e p h a l i c spines l y i n g in a
c u r v e ( F i g u r e s 1 1 . 6 C , E , F). B o t h g e n e r a h a v e spines d e v e l o p e d from the
p l e u r a l l o b e s o f t h e thorax a n d the p y g i d i u m . Ross (1979) provided e x c e l l e n t
illustrations of t h e s e very s m a l l trilobites that are u s u a l l y less than 1 cm in
l e n g t h a n d rarely f o u n d c o m p l e t e . Acidaspis o c c u r s from the basal K o p e
Figure 11.10. Cincinnatian
ostracodes. A. Cer- F o r m a t i o n t h r o u g h the top o f the M a v s v i l l i a n M l . A u b u r n F o r m a t i o n , and

atopsis chambersi r e a p p e a r s i n t h e R i c h m o n d i a n W a y n e s v i l l e F o r m a t i o n . W h i t t i n g t o n (1956)

(Miller), CMC IP 40171, s u g g e s t e d that a s i n g l e s p e c i e s , A. cincinnatiensis, is present. Primaspis cro-
Kope Formation, Cincin- sotus ( L o c k e ) o c c u r s in t h e K o p e F o r m a t i o n w h e r e it was found in associa-
nati, Ohio. From War- tion w i t h a cluster of Flexicalymene ( H u g h e s a n d C o o p e r 1999).
shauer (1972, plate 5), x
T h e e x t r e m e s p i n o s i t y o f o d o n t o p l e u r i d s has p r o m p t e d m u c h debate-
30. B. Ctenobolbina
a b o u t t h e i r m o d e o f life. T h e f r i n g e o f v e r t i c a l c e p h a l i c s p i n e s i n Acidaspis
alata Ulrich, CMC IP
e n a b l e d t h e c e p h a l o n t o b e rested a g a i n s t t h e s u b s t r a t u m w i t h the thorax
33085, Kope Formation,
a n d p y g i d i u m o u t s t r e t c h e d a n d e l e v a t e d slightly, in a p r o b a b l e f e e d i n g posi-
Cincinnati, Ohio, x
33. C. Quadrijugator t i o n ( W h i t t i n g t o n 1956). I n this p o s i t i o n t h e h y p o s t o m e a n d m o u t h w e r e

regularis (Emmons), situated c l o s e t o t h e s u b s t r a t u m . C l a r k s o n (1969) p o i n t e d out that w h e r e a s

CMC IP 28245, Waynes- Acidaspis h a d a s i n g l e , fixed life p o s i t i o n , Primaspis c o u l d a s s u m e t h e s a m e
ville Formation, Butler f e e d i n g p o s i t i o n a n d also a r e s t i n g a t t i t u d e by t i l t i n g the c e p h a l o n back-
Co., Ohio, x 90. w a r d s s o that t h e e n t i r e b o d y w a s s u p p o r t e d a g a i n s t the s u b s t r a t u m . O d o n -

760 A Sea without Fish

t o p l e u r i d s arc c o n s i d e r e d t o h a v e b e e n p o t e n t i a l p r e d a t o r s b y F o r t e y a n d
O w e n s 11999), b u t their prey m u s t h a v e b e e n very s m a l l .
Plate 6 a n d F i g u r e 11.7 illustrate e x c e p t i o n a l l y w e l l - p r e s e r v e d s p e c i -
m e n s of s o m e of the rarest of C i n c i n n a t i a n trilobite s p e c i e s .

O n e o f the most i n t r i g u i n g fossils k n o w n from the C i n c i n n a t i a n i s the e u r y p - Eurypterids

terid Megalograptus ( F i g u r e s 11.8, 11.9). E u r y p t e r i d s are c h e l i c e r a t e a r t h r o -
p o d s , d i s t i n g u i s h e d b y h a v i n g t h e f i r s t pair o f a p p e n d a g e s ( c h e l i c e r a e )
e q u i p p e d w i t h small p i n c e r s . M o d e r n c h e l i c e r a t e s are t h e h o r s e s h o e crabs
(Plate 3 G ) , and a r a c h n i d s (scorpions, spiders, m i t e s , a n d ticks). The b o d y of
the eurypterid is u n i q u e , w i t h a distinct h e a d ( p r o s o m a ) b e a r i n g c o m p o u n d
eyes followed b y a n e l o n g a t e d , s e g m e n t e d s e c t i o n c a l l e d the o p i s t h o s o m a ,
divided into a w i d e r p r e a b d o m e n a n d a narrower, tail-like p o s t a b d o m e n (Fig-
ures 11.9A, B). Six pairs of a p p e n d a g e s w e r e a t t a c h e d to the u n d e r s i d e of t h e
head and served f u n c t i o n s of f e e d i n g a n d l o c o m o t i o n . B e c a u s e t h e exoskel-
eton was not c a l c i f i e d , preservation of the c h i t i n o u s r e m a i n s of e u r y p t e r i d s
was unlikely, a n d therefore they are u s u a l l y very rare fossils.
E u r y p t e r i d s first a p p e a r e d in the Early O r d o v i c i a n a n d a t t a i n e d t h e i r
m a x i m u m diversity in the S i l u r i a n , b u t Megalograptus is s i g n i f i c a n t for
b e i n g o n e o f the oldest a n d most u n u s u a l . W h e n f r a g m e n t s o f t h e a n i m a l
w e r e first d e s c r i b e d in 1874 by S. A. M i l l e r , they w e r e t h o u g h t to be parts
o f a graptolite, h e n c e the n a m e . L a t e r w o r k e r s c o r r e c t e d t h e e r r o r o n t h e

Arthropods 161
basis o f a d d i t i o n a l , a l b e i t f r a g m e n t a r y d i s c o v e r i e s . A t r u l y p h e n o m e n a l
d i s c o v e r y m a d e i n 1938 o f e x c e p t i o n a l l y w e l l - p r e s e r v e d and nearly c o m -
plete s p e c i m e n s from a single bed in the uppermost C i n c i n n a t i a n Elkhorn
F o r m a t i o n o f t h e R i c h m o n d G r o u p i n A d a m s C o u n t y , O h i o , led t o a better
u n d e r s t a n d i n g o f the a n i m a l ( F i g u r e 11.8B). This m a t e r i a l , w h i c h i n c l u d e d
m a l e a n d f e m a l e s p e c i m e n s , b e c a m e t h e basis for a n e w s p e c i e s , M. ohio-
ensis, d e s c r i b e d by K e n n e t h F. C a s t e r a n d F r i k K j e l l e s v i g - W a e r i n g in 1964.
One additional eurypterid species, Eocarcinosoma batrachophthalmus, was
d e s c r i b e d f r o m this b e d o n the basis o f a n isolated p r o s o m a .
Megalograptus ohioensis w a s o n e of the largest creatures in the C i n c i n -
natian sea floor c o m m u n i t y , r e a c h i n g a l e n g t h of over 50 c m . The first pair
o f a p p e n d a g e s , t h e c h e l i c e r a e , i s s m a l l a n d l o c a t e d b e n e a t h the h e a d . T h e
n e x t t h r e e pairs o f a p p e n d a g e s b e a r w e l l - d e v e l o p e d spines ( F i g u r e 11.9). The
third a p p e n d a g e s are m o s t striking for their l e n g t h and l o n g spines directed
toward t h e m i d l i n e ( F i g u r e s 1 1 . 9 C , D ) . E x a c t l y how the eurypterid used
t h e s e spiny a p p e n d a g e s is u n c e r t a i n . C a s t e r a n d K j e l l e s v i g - W a e r i n g consid-
ered Megalograptus to h a v e b e e n a predator, a n d thus the a p p e n d a g e s likely
h a d s o m e f u n c t i o n i n g r a s p i n g prey. T h e basket-like structure o f the l o n g
spines o f t h e third a p p e n d a g e s s u g g e s t s that the a n i m a l m i g h t have raked
t h e m t h r o u g h the s e d i m e n t in order to extract prey in a s i e v i n g fashion. Only
a few o t h e r e u r y p t e r i d s h a v e similar l o n g spiny a p p e n d a g e s . Tubular castings
f i l l e d w i t h f r a g m e n t s o f e u r y p t e r i d i n t e g u m e n t associated w i t h the e u r y p -
terid m a t e r i a l c o u l d represent feces o f t h e a n i m a l , i n d i c a t i n g c a n n i b a l i s t i c
b e h a v i o r like that f o u n d in l i v i n g e u r y p t e r i d relatives a m o n g the scorpions
and spiders. The fourth pair of a p p e n d a g e s lacks spines, and the fifth pair
has e x p a n d e d , flattened segments giving them a paddle-like appearance
( F i g u r e s 11.9A, B). T h e s e w e r e m o s t l i k e l y e m p l o y e d i n s w i m m i n g .
O n e o f t h e m o s t p e c u l i a r features o f Megalograptus i s t h e d e v e l o p m e n t
at t h e e n d of the p o s t a b d o m e n of a pair of e x p a n d e d , h o o k - l i k e c e r e a l
b l a d e s flanking a s p i n e - l i k e telson ( F i g u r e s 11.9A, B). C a s t e r a n d Kjellesvig-
W a e r i n g t h o u g h t that the c e r e a l b l a d e s c o u l d m o v e laterally in a scissor-like
m o t i o n , possibly serving to grasp either in defense or copulation. The
p a i r e d , i n c u r v e d p o s t e r i o r s p i n e s o f e a r w i g s a r e s i m i l a r , b u t n o t h i n g like
this structure appears in any other eurypterid.
In a d d i t i o n to t h e A d a m s C o u n t y o c c u r r e n c e of Megalograptus ohioen-
sis in t h e Elkhorn b e d s . C a s t e r and K j e l l e s v i g - W a e r i n g d e s c r i b e d two other
s p e c i e s from the C i n c i n n a t i a n . M. shideleri is k n o w n from f r a g m e n t s o c c u r -
r i n g in the S a l u d a F o r m a t i o n of the R i c h m o n d C r o u p and M. williamsae
from the W a y n e s v i l l e F o r m a t i o n o f the R i c h m o n d G r o u p . F r a g m e n t s o f M .
shideleri s u g g e s t that it m a y h a v e r e a c h e d t w o meters in l e n g t h . The t y p e
s p e c i e s , M . welchi, w a s o r i g i n a l l y d e s c r i b e d from the Liberty F o r m a t i o n ,
t h e r e b y i n d i c a t i n g that t h e g e n u s r a n g e s t h r o u g h virtually the entire u p p e r
C i n c i n n a t i a n . Megalograptus w a s not restricted to the C i n c i n n a t i A r c h re-
g i o n , as M. alveolatus is k n o w n from the Upper O r d o v i c i a n of V i r g i n i a .
In t h e L i b e r t y o c c u r r e n c e , d e s c r i b e d by Foerste (1912), Megalograptus
w a s f o u n d in a p o c k e t t o g e t h e r w i t h c r i n o i d s . In t h e Elkhorn, Megalograp-
tus w a s a s s o c i a t e d w i t h a diverse m a r i n e fauna i n c l u d i n g trilobites, b r a c h i o -
p o d s , b r y o z o a n s , a n d m o l l u s c s . D e s p i t e the c o m m o n n o t i o n that e u r y p -

A Sea without Fish

terids lived in s o m e w h a t restricted or a t y p i c a l m a r i n e e n v i r o n m e n t s , t h e
C i n c i n n a t i a n o c c u r r e n c e s a r g u e s t r o n g l y for a s s o c i a t i o n w i t h t h e n o r m a l
m a r i n e biota. Their c h i t i n o u s i n t e g u m e n t m a y a c c o u n t largely f o r t h e i r
rarity i n the C i n c i n n a t i a n . T h e e x t r a o r d i n a r y q u a l i t y a n d q u a n t i t y o f e u -
rypterid p r e s e r v a t i o n a t the A d a m s C o u n t y site m a y h a v e resulted f r o m
s m o t h e r i n g o f the m a r i n e f a u n a b y v o l c a n i c a s h f a l l , b e c a u s e the 1 5 c m -
thick s h a l e w i t h i n w h i c h t h e fossils w e r e c o n c e n t r a t e d w a s f o u n d t o c o n t a i n
b e n t o n i t i c clays ( C a s t e r and K j e l l e s v i g - W a e r i n g 1964).

Neostrabops

In 1952 C a s t e r and M a c k e described w h a t they t e r m e d a m a v e r i c k m e r o s t o m e ,

Neostnibops martini, from a single s p e c i m e n found in the M a y s v i l l i a n C o r r y -
ville f o r m a t i o n i n 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). This fossil c o u l d
be taken to be a trilobite, but lacks the characteristic l e n g t h w i s e division into
three distinct lobes. It also resembles the a f o r e m e n t i o n e d eurypterids in hav-
ing n u m e r o u s narrow s e g m e n t s , a l t h o u g h it lacks a n y d e m a r c a t i o n of p r e - a n d
p o s t a b d o m e n . Neostrabops d o e s resemble other arthropods k n o w n as aglaspids
from the C a m b r i a n . Aglaspids are regarded as early offshoots of the evolution-
ary lineage of m o d e r n horseshoe crabs, the w e l l - k n o w n Limulus.

C r u s t a c e a n s are o n e o f the most a b u n d a n t and diverse g r o u p s o f l i v i n g a r t h r o - Ostracodes

p o d s , yet t h e y are represented in the C i n c i n n a t i a n by o n l y o n e g r o u p , the
o s t r a c o d e s (major l i v i n g c r u s t a c e a n s s u c h a s s h r i m p s , lobsters, a n d crabs
evolved m u c h later than the O r d o v i c i a n , a n d thus are not f o u n d in the C i n -
c i n n a t i a n ) . O s t r a c o d e s are g e n e r a l l y very s m a l l , less t h a n 1 mm in l e n g t h , but
c a n e x c e e d 1 cm in forms like the O r d o v i c i a n Eoleperditia. T h e y are distin-
g u i s h e d by a calcitic, bivalved shell or c a r a p a c e that c a n be s m o o t h or have
various s u r f a c e features such a s k n o b s , ridges, o r spines ( f i g u r e 11.10). W h e n
t h e h i n g e d valves o f the c a r a p a c e o p e n , the a p p e n d a g e s c a n b e e x t e n d e d for
f e e d i n g and l o c o m o t i o n . M o d e r n o s t r a c o d e s have a w i d e r a n g e o f f e e d i n g
habits, but m a n y live as b e n t h i c s u s p e n s i o n feeders a n d deposit feeders. It is
very difficult t o d e t e r m i n e the s p e c i f i c habits o f C i n c i n n a t i a n s p e c i e s , b e -
c a u s e the a p p e n d a g e s and o t h e r internal a n a t o m y are not preserved
O s t r a c o d e s are diverse and a b u n d a n t t h r o u g h o u t t h e C i n c i n n a t i a n
S e r i e s , a l t h o u g h this is g e n e r a l l y u n a p p r e c i a t e d b e c a u s e few c o l l e c t o r s or
researchers e n c o u n t e r t h e s e m i c r o f o s s i l s . Two m a j o r s t u d i e s o n C i n c i n -
natian o s t r a c o d e s p r o v i d e a m o d e r n analysis of their diversity a n d classifica-
tion, but there is no s i n g l e c o m p r e h e n s i v e s t u d y that gives an o v e r v i e w of
total C i n c i n n a t i a n o s t r a c o d e diversity. W a r s h a u e r a n d B e r d a n (1982) re-
ported f i f t y - t h r e e s p e c i e s a n d t h i r t y - n i n e g e n e r a o f o s t r a c o d e s b e l o n g i n g t o
t w o m a j o r orders, the P a l a e o c o p i d a a n d t h e P o d o c o p i d a , f r o m t h e M i d d l e
O r d o v i c i a n L e x i n g t o n L i m e s t o n e a n d the U p p e r O r d o v i c i a n (basal C i n -
c i n n a t i a n ) C l a y s Kerry f o r m a t i o n o f K e n t u c k y . S p e c i e s i n t h e s e g r o u p s are
all very s m a l l , a n d w e r e e x t r a c t e d f r o m d i s a g g r e g a t e d s h a l e s or by a c i d dis-
solution o f t h e L e x i n g t o n L i m e s t o n e i n which t h e f o s s i l s are silicified.
f o u r t e e n g e n e r a o c c u r i n t h e C l a y s Ferry f o r m a t i o n , b u t d i s t r i b u t i o n

Arthropods 163
 w i t h i n t h e rest o f t h e C i n c i n n a t i a n was not s t u d i e d . B e r d a n (1984) reported
on ostracodes of the order Leperditicopida from the M i d d l e and Upper
O r d o v i c i a n o f K e n t u c k y a n d v i c i n i t y . T h e s e o s t r a c o d e s are n o t e w o r t h y for
their size (sometimes > 1 cm long) and high a b u n d a n c e in single beds of
f i n e - g r a i n e d l i m e s t o n e . T h e o c c u r r e n c e o f l e p e r d i t i c o p i d s i s restricted t o
f i n e - g r a i n e d l i m e s t o n e f a d e s d e p o s i t e d i n e x t r e m e l y s h a l l o w subtidal t o
intertidal e n v i r o n m e n t s p a r t i c u l a r l y w e l l k n o w n f r o m t h e M i d d l e O r d o v i -
c i a n H i g h B r i d g e G r o u p o f K e n t u c k y ( C r e s s m a n a n d N o g e r 1976). T h i s
u n i q u e f a c i e s i s a b s e n t f r o m t h e d e e p e r w a t e r f a d e s o f t h e l o w e r and m i d d l e
C i n c i n n a t i a n but recurs in the R i e h m o n d i a n Sunset M e m b e r of the Arn-
h e i m F o r m a t i o n , i n w h i c h four s p e c i e s are f o u n d .

164 A Sea without Fish

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.
Comactinia sp., Carib-
bean. Scanning electron
micrograph, x 79.

Figure 1 2 . 2 . Arm of
modern crinoid (dark)
with pinnules (light
branches) bearing fine
tube feet in feeding pos-
ture. Pinnule length
about 1 cm. Aquarium
photo, comasterid cri-
noid, Curacao, Nether-
lands Antilles

166 A Sea without Fish

 12
ECHINODERMS: A WORLD
UNTO THEMSELVES

E c h i n o d e r m s are a m o n g the rarest and m o s t sought-after fossils in t h e C i n - I . . here salute the

c i n n a t i a n rocks. N o t o n l y are they c o m p l e x in form a n d s t r u c t u r e , but they noble echinoderms
also possess a c e r t a i n b e a u t y a n d mystery that n e v e r fail to attract interest. as a noble group es-

A n y o n e w h o has visited the seashore is f a m i l i a r w i t h l i v i n g e c h i n o d e r m s s u c h pecially designed to

puzzle the zoologist.
as sea stars or starfish (asteroids), sea u r c h i n s , a n d s a n d d o l l a r s ( b o t h e c h i -
noids) (Plate 9), O t h e r l i v i n g e c h i n o d e r m s found in d e e p e r m a r i n e waters are L. H. H y m a n 1955, vi
the sea lilies and feather stars (crinoids), brittle stars (ophiuroids), and sea
c u c u m b e r s ( h o l o t h u r o i d s ) (Plate 9). T h e r e are a b o u t 6650 l i v i n g s p e c i e s of
e c h i n o d e r m s , a n d over 3500 g e n e r a a n d 13,000 d e s c r i b e d fossil s p e c i e s .
The O r d o v i c i a n Period m a r k e d a very significant t i m e in the e v o l u t i o n
o f e c h i n o d e r m s , b e c a u s e m a n y different m a j o r g r o u p s ( u s u a l l y r e g a r d e d a s
classes) o f e c h i n o d e r m s c o e x i s t e d . L i k e o t h e r i n v e r t e b r a t e p h y l a , t h e oldest
fossil e c h i n o d e r m s are found in Early C a m b r i a n r o c k s o v e r 500 m i l l i o n
years o l d , but it was not until O r d o v i c i a n t i m e that e c h i n o d e r m s b e g a n to
leave a significant fossil r e c o r d . Early in t h e O r d o v i c i a n , e c h i n o d e r m s di-
versified a l o n g w i t h m a n y o t h e r m a r i n e i n v e r t e b r a t e s , a n d b y L a t e O r d o v i -
c i a n t i m e a b e w i l d e r i n g variety of e c h i n o d e r m s h a d a p p e a r e d in shallow
m a r i n e e n v i r o n m e n t s w o r l d w i d e . A l t h o u g h the f i v e classes o f m o d e r n e c h i -
n o d e r m s ( c r i n o i d s , asteroids, o p h i u r o i d s , e c h i n o i d s , a n d h o l o t h u r o i d s ) ex-
isted t h e n , m a n y o t h e r classes w e r e also p r e s e n t . In s o m e strata of Late
Ordovician | M o h a w k i a n ) age, as m a n y as fourteen classes of e c h i n o d e r m s
are f o u n d t o g e t h e r ( S p r i n k l e a n d C u e n s b u r g 1997). B e c a u s e s o m e o f t h e s e
classes soon b e c a m e e x t i n c t , e c h i n o d e r m d i v e r s i t y h a d d e c l i n e d b y C i n c i n -
natian t i m e , but still e x c e e d e d p r e s e n t levels, w i t h s e v e n classes f o u n d in
the C i n c i n n a t i A r c h r e g i o n .
M o r e t h a n o n e o b s e r v e r has n o t e d ( t o n g u e i n c h e e k ) that i f e v e r a n y
a n i m a l g r o u p c o u l d h a v e o r i g i n a t e d a s " a l i e n b e i n g s " that l a n d e d o n E a r t h
from outer s p a c e , i t w o u l d b e the e c h i n o d e r m s s o b i z a r r e a r e their b o d y
f o r m s , p a r t i c u l a r l y a m o n g t h e w i d e variety w e s e e a m o n g O r d o v i c i a n fos-
sils. T h e features that identify all t h e s e s t r a n g e fossils as e c h i n o d e r m s are
not the traits w e u s u a l l y c o n s i d e r c h a r a c t e r i s t i c o f e c h i n o d e r m s , t h e "spiny
skin" that gives the g r o u p its n a m e , a n d the f i v e - f o l d ( p e n t a m e r a l ) s y m -
metry o f the body. T h e s i n g l e m o s t c h a r a c t e r i s t i c trait o f e c h i n o d e r m s i s
the n a t u r e of their skeleton. All e c h i n o d e r m s h a v e a m i n e r a l i z e d s k e l e t o n
c o m p o s e d o f c a l c i u m c a r b o n a t e a s the m i n e r a l c a l c i t e . U n l i k e o t h e r inver-
tebrates that have calcitic shells, the e c h i n o d e r m skeleton is f o r m e d w i t h i n
the m i d d l e cell layer of the body ( m e s o d e r m ) , a n d has a t h i n o u t e r layer of
c e l l s c o v e r i n g it ( e c t o d e r m ) , t h u s m a k i n g it an i n t e r n a l s k e l e t o n . B e c a u s e
the outer cell layer is so t h i n , we often t h i n k of e c h i n o d e r m s k e l e t o n s s u c h

767
 a s sea u r c h i n shells a s e x t e r n a l , b u t i n o t h e r c a s e s , s u c h a s t h e a r m s o f m a n y
sea stars, t h e skeletal c o m p o n e n t s , c a l l e d o s s i c l e s , are clearly i n t e r n a l , b e -
n e a t h a l e a t h e r y " s k i n . " In a d d i t i o n , b e c a u s e it is truly m e s o d e r m a l , the
e c h i n o d e r m s k e l e t o n has a u n i q u e m i c r o s t r u c t u r e not f o u n d i n a n y o t h e r
a n i m a l g r o u p . T h e c a l c i t e i s f o r m e d a r o u n d m e s o d e r m a l c e l l s into a n in-
tricate t h r e e - d i m e n s i o n a l l a t t i c e w o r k c a l l e d the s t e r e o m ( F i g u r e 12.1). Skel-
e t a l p l a t e s , s p i n e s , o r ossicles t h u s h a v e a h i g h l y p o r o u s s t r u c t u r e i n w h i c h
o v e r 50 p e r c e n t of t h e v o l u m e c a n be t a k e n up by p o r e s . In life t h e nurtur-
i n g c e l l s o c c u p y t h e s e p o r e s , b u t after d e a t h , the c e l l u l a r m a t e r i a l d e c a y s ,
l e a v i n g t h e p o r o u s s k e l e t o n . B u r i e d i n s e d i m e n t , t h e s e p o r e s are u s u a l l y
i n f i l l e d w i t h s e c o n d a r y c a l c i t e , a n d the entire skeletal plate displays t h e
t y p i c a l r h o m b i c c l e a v a g e o f c a l c i t e . O f t e n t h e m i c r o s t r u c t u r e i s still visible
i n t h i n o r p o l i s h e d s e c t i o n s . T h e s e f e a t u r e s h a v e e n a b l e d m a n y fossil e c h i -
n o d e r m s t o b e c o r r e c t l y i d e n t i f i e d , e v e n t h o u g h their b o d y form m a y b e
considerably different from any of the five familiar living groups.
Echinoderms are a l s o p e c u l i a r in l a c k i n g s t r u c t u r e s like a d i s t i n c t
h e a d , e y e s , or i n t e r n a l s y s t e m s s u c h as a b l o o d c i r c u l a t o r y system or respira-
tory s y s t e m . I n s t e a d , t h e y are u n i q u e i n h a v i n g a n i n t e r n a l system o f
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 a t e r y fluid that c i r c u l a t e s
d i s s o l v e d o x y g e n a n d d i s s o l v e d w a s t e s a n d p r e s s u r i z e s t h e vessels t h e m -
selves. T h e vessels t e r m i n a t e i n c h a r a c t e r i s t i c s t r u c t u r e s c a l l e d t u b e f e e t ,
w h i c h s e r v e i m p o r t a n t f u n c t i o n s for all e c h i n o d e r m s ( F i g u r e 12.2). D i s -
s o l v e d o x y g e n i s e x c h a n g e d across the t h i n m e m b r a n e o f the t u b e feet a n d
d i s s o l v e d w a s t e s are e x p e l l e d . In g r o u p s like c r i n o i d s a n d o p h i u r o i d s , tube-
feet are l i k e m i n u t e bristles that c a p t u r e f o o d p a r t i c l e s s u s p e n d e d i n t h e
water. S e a stars a n d sea u r c h i n s h a v e t u b e feet w i t h s u c t i o n disks b y w h i c h
t h e y c l i n g t o r o c k s , a n d w h i c h aid i n p u l l i n g c l a m s h e l l s apart, i n the c a s e
of sea stars, a n d in " w a l k i n g " a c r o s s t h e sea floor. Tube feet are p r a c t i c a l l y
n e v e r p r e s e r v e d in fossils, b u t traces of t h e c a n a l s are r e v e a l e d in t h e skel-
e t o n , p r o v i d i n g y e t a n o t h e r i n d i c a t i o n o f e c h i n o d e r m affinity. R e c e n t l y
p y r i t i z e d t u b e feet w e r e d i s c o v e r e d i n a C i n c i n n a t i a n o p h i u r o i d , p r o v i d i n g
o n e o f t h e f e w c a s e s i n t h e e n t i r e fossil r e c o r d a n d rare e v i d e n c e for pre-
s e r v e d soft tissues a m o n g C i n c i n n a t i a n fossils ( C l a s s 2006).
The five-fold or pentameral body plan seen in living e c h i n o d e r m s
a p p e a r s i n m a n y o f t h e C i n c i n n a t i a n fossil e c h i n o d e r m s ( F i g u r e s 12.5,
12.10-12.16), b u t this is by no m e a n s a u n i v e r s a l f e a t u r e . The enigmatic
" c a r p o i d s " l a c k a n y t r a c e o f p e n t a m e r a l form ( F i g u r e 12.17). T h e l a r v a e o f
l i v i n g e c h i n o d e r m s are a c t u a l l y b i l a t e r a l l y s y m m e t r i c a l , a n d p e n t a m e r y
a p p e a r s o n l y in a d u l t s t a g e s . The five-part s t r u c t u r e is v i r t u a l l y u n i q u e to
e c h i n o d e r m s i n t h e A n i m a l K i n g d o m , a n d z o o l o g i s t s h a v e d e b a t e d its sig-
n i f i c a n c e a n d o r i g i n s . S p r i n k l e (1973) f o u n d that p e n t a m e r a l s y m m e t r y f i r s t
a p p e a r e d in the food-gathering system of Early and M i d d l e C a m b r i a n
e o c r i n o i d s , a n d later d e v e l o p e d i n t h e c a l y x plates a n d respiratory struc-
t u r e s . T h i s s u g g e s t s that p e n t a m e r a l s y m m e t r y p r o v i d e d a n a d v a n t a g e for
t h e sessile, f i l t e r - f e e d i n g h a b i t s o f t h e s e early e c h i n o d e r m s . Initially, s o m e
e o c r i n o i d s a c t u a l l y h a v e a three-fold radial a r r a n g e m e n t o f t h e f o o d - g a t h -
e r i n g s t r u c t u r e s that later b e c a m e five-fold b y the b r a n c h i n g o f o n l y two
f o o d g r o o v e s o r a m b u l a c r a . B r a n c h i n g b e y o n d the f i v e - f o l d pattern m a y

168 A Sea without Fish

h a v e b e e n l i m i t e d b y the a v a i l a b l e s p a c e a r o u n d t h e m o u t h , a n d t h u s t h e
p e n t a m e r a l pattern m a y h a v e b e e n the most efficient s o l u t i o n . O n c e p e n -
t a m e r a l s t r u c t u r e b e c a m e g e n e t i c a l l y p r o g r a m m e d in Echinoderms, it per-
sisted e v e n i n g r o u p s that g a v e u p t h e a n c e s t r a l m o d e o f life t o b e c o m e
m o b i l e sea stars or sea u r c h i n s .
D e s p i t e their m a n y " a l i e n " features, E c h i n o d e r m s a r e s i g n i f i c a n t a s o n e
of the invertebrate phyla m o s t closely related to o u r o w n , the c h o r d a t e s . For
a l o n g t i m e zoologists s t u d y i n g the e m b r y o n i c d e v e l o p m e n t of echinoderms
have r e c o g n i z e d close similarities in the early d e v e l o p m e n t of echinoderms
and chordates. B o t h g r o u p s have s y m m e t r i c a l cell division in the fertilized
e g g , i n d e t e r m i n a t e d e v e l o p m e n t ( e m b r y o n i c cells are not p r e p r o g r a m m e d
to form a specific adult tissue), a n d the internal b o d y cavity, t h e c o e l o m ,
forms in the s a m e way in the e m b r y o . R e c e n t studies of m o l e c u l a r c o m p o s i -
tion o f a n i m a l phyla d e m o n s t r a t e that e c h i n o d e r m s are m u c h m o r e closely
allied to h e m i c h o r d a t e s and c h o r d a t e s than to any o t h e r g r o u p (Raff 1996).
C o m p l e t e fossil echinoderms are i n d e e d rare fossils in C i n c i n n a t i a n
strata, but the a b u n d a n c e of their isolated skeletal c o m p o n e n t s suggests that
t h e y w e r e very c o m m o n m e m b e r s o f sea floor c o m m u n i t i e s d u r i n g the O r -
d o v i c i a n . The reason for their rarity as c o m p l e t e fossils is f o u n d in the n a t u r e
of the e c h i n o d e r m skeleton, c o m p o s e d of m y r i a d tiny plates or ossicles, all
held together in life by fibers of c o n n e c t i v e tissue that p e n e t r a t e pores of the
s t e r e o m i c skeleton. U p o n d e a t h , these f i b e r s rapidly d e c a y ; the skeletal c o m -
p o n e n t s separate, a n d are u s u a l l y dispersed b y w a t e r m o v e m e n t ( F i g u r e
12.3A). A m o n g t h e m o s t c o m m o n fossils f o u n d a r o u n d C i n c i n n a t i are t h e
disk-like or ring-like s e g m e n t s of the c r i n o i d stem ( c o l u m n a l s ; F i g u r e s 12.3A,
12.5). T h e s e are often isolated like tiny LifeSavers or still c o n n e c t e d in vary-
ing l e n g t h s like strings of b e a d s . Entire l i m e s t o n e layers often consist of
n o t h i n g but dissociated crinoid c o l u m n a l s , o r i g i n a l l y c r i n o i d a l sands, s o m e -
times sorted b y c u r r e n t s into z o n e s o f c o m m o n size, a n d f o r m e d into rippled
surfaces. B e c a u s e echinoderms are so s u s c e p t i b l e to b r e a k u p of t h e skeleton
after d e a t h , the o c c u r r e n c e of c o m p l e t e , a r t i c u l a t e d skeletons u s u a l l y re-
quires rapid burial in s e d i m e n t . Thus, c o m p l e t e c r i n o i d s are f o u n d w i t h i n
shales or on the u p p e r s u r f a c e of l i m e s t o n e s c o v e r e d by shales, or m o r e rarely
at the base of c a l c a r e o u s siltstones. T h e very n a t u r e of this c o m p l e t e preserva-
tion provides a v a l u a b l e c l u e that the e n c l o s i n g s e d i m e n t w a s in fact d e p o s -
ited rapidly, probably t i m i n g storms, either as stirred up b o t t o m s e d i m e n t s or
underwater mudslides.

T h e seven classes o f e c h i n o d e r m s f o u n d i n C i n c i n n a t i a n strata are t h e Echinoderms of

Crinoidea, Rhomhifera, Kdrioasteroidca, Asteroidea, Ophiuroidea, C y c l o - t h e Cincinnatian
c y s t o i d e a , a n d S t y l o p h o r a . o f t h e s e , the c r i n o i d s a r e t h e m o s t a b u n d a n t i n
the f i e l d a n d the m o s t d i v e r s e i n n u m b e r o f s p e c i e s .

Crinoids

S t e m m e d crinoids (sea lilies) are a t t a c h e d to the sea floor by m e a n s of a hold-

fast or root, and elevated a b o v e the b o t t o m by a stem c o m p o s e d of m a n y c o -

Echinoderms 169
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 disarticu-
lated skeletal compo-
nents. Upper Ordovician,
Corryville Formation, Cin-
cinnati, Ohio. University
of Cincinnati collections.
B. Two articulated
crowns, detached from
stalk, oriented parallel
but in opposite direction,
preserved on base of
bed. Upper Ordovician,
Corryville Formation,
Clermont Co., Ohio. CMC
IP 44362. Scale in mm. B
in upper right denotes
basal plate; R denotes
radial plate.

l u m n a l s ( F i g u r e s 12.3-12.5). T h e b o d y is e n c l o s e d w i t h i n a plated c a l y x c o m -
p o s e d of t h e l o w e r c u p a n d a roof-like tegmen. T h e c u p has a regular
a r r a n g e m e n t of plates: a circlet of five radials, a n d either o n e circlet ( m o n o -
There were also s o m e
cyclic) of basals or t w o circlets (dicyclic) of basals a n d infrabasals o c c u r r i n g
beautiful forms of Cri-
a b o v e t h e stem ( F i g u r e s 12.3B, 12.10A). T h e a r m s arise from the radials of the
noidea, or stone-lilies . .
c u p a n d usually b r a n c h a n d m a y support finer b r a n c h e s , the p i n n u l e s , g i v i n g
C h a r l e s Lyell 1845, 50 the a r m a feathery a p p e a r a n c e ( F i g u r e 12.4). T h e a r m s and p i n n u l e s carry the
food g r o o v e s (ambulacra) w h i c h c o n v e y tiny food particles to the m o u t h situ-
ated on the u p p e r surface of t h e c a l y x . I .iving crinoids are passive suspension
feeders, d e p e n d e n t on currents to supply food particles to the a w a i t i n g feed-
i n g apparatus. H i e a n i m a l c o n s t r u c t s a filtration fan of the ar m s and pinnules

770 A Sea without Fish

 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 with
living crinoids, current
flow was from left to
right. Drawing by John
Agnew.

Echinoderms 171
Figure 12.5. Cincinnatian
crinoid columnals and
holdfasts. A, H. Cincin-
naticrinus varibrachialus
Warn and Strimple; A.
Articular surface x 9.6;
H. Lateral view of mature
section x 5.9. B, I. Ect-
enocrinus simplex
(Hall). B. Articular surface
y.9.6; I. Lateral view*
12.6. C, J. locrinus sub-
crassus (Meek and
Worthen). C. Articular
surface x 5.2; J. iafera/
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
curtus Ulrich, articular
surface x 5.2. F. Anom-
alocrinus incurvus (Meek
and Worthen), lateral view
of compressed section
with fracture separating
meres, x 1. A - K , from
Meyer et al. (2002, figure
2) and reprinted by permis-
sion of The Paleontological
Society. I, M. Lichenocri-
nus crateriformis Hall,
FMNH8810. L. Two speci- arrayed p e r p e n d i c u l a r to c u r r e n t Flow, thus a c t i n g as a " f l o w - t h r o u g h " filter
mens attached to brachio- (Plates 9A, B). T i n y t u b e feet l i n i n g the p i n n u l a r food grooves snare food
pod, x 1.6. M. Enlarge- particles s u c h as o r g a n i c detritus a n d p l a n k t o n and stuff t h e m into the food
ment of larger specimen in g r o o v e for transport to the m o u t h . The close similarity of O r d o v i c i a n crinoids
I, y.3.7, Waynesville For- to m o d e r n forms suggests that a n c i e n t crinoids used similar f e e d i n g postures
mation, Warren Co., Ohio.
a n d m o d e s o f c a p t u r i n g food, and w e c a n i m a g i n e that they looked quite like
From Faber (1929, plate 32,
s t e m m e d c r i n o i d s that today exist o n l y in the d e e p sea (Plate qA).
figures 5, 6) and reprinted
C r i n o i d c o l u m n a l s a r e a m o n g t h e m o s t c o m m o n l y e n c o u n t e r e d fossils
by permission of the
American Midland Nat- i n t h e C i n c i n n a t i a n . T h e y are Found e i t h e r a s s i n g l e c o l u m n a l s o r a r t i c u -

uralist. N. Lichenocrinus lated s e c t i o n s o F t h e s t e m ( f i g u r e s 12.3, 12.5). A l t h o u g h s u p e r f i c i a l l y very

milleri Faber, CMC IP
10047, Elkhorn Formation,
cross-section showing
plate with central node,
crater plates, and base of
column, y.3.7, from Faber
(1929, plate 33, figure 9).
s i m i l a r i n a p p e a r a n c e , c o l u m n a l s c a n u s u a l l y b e identified t o g e n u s . C r i -
n o i d s p e c i e s are u s u a l l y d e s c r i b e d from s p e c i m e n s o f the c a l y x o r c r o w n
floor that are less c o m m o n .
T w e n t y - o n e g e n e r a o f c r i n o i d s a r e k n o w n From the C i n c i n n a t i region.
S o m e g e n e r a are m o n o s p e c i f i c ( h a v i n g o n l y o n e species) and others have
m o r e t h a n o n e s p e c i e s , b r i n g i n g to t w e n t y - e i g h t the total n u m b e r oF species
in the local area. C i n c i n n a t i a n c r i n o i d s represent all three oF the subclasses

172 A Sea without Fish

Figure 12.6. Cincinnatian disparid crinoids. A, B. Cincinnaticrinus varibrachialus Warn andStrimple.
A, B. Holotype, CMC IP 3871, Kope Formation, Cincinnati, Ohio, x 1.0 and x 3.1. C. C. pentagonus
(Ulrich), YPM 24801, Fairview Formation, Cincinnati, Ohio, x 1.3. D. Dystactocrinus constrictus (Hall),
showing regeneration of arms, USNM 93223, Fairview Formation, Hamilton Co., Ohio, x 2.0. E. Ecteno-
crinus geniculatus (Ulrich), CMC IP 36313, Kope Formation, Cincinnati, Ohio, x 7.4. F, G.
E. simplex (Hall) F. CMC IP, x 7.5. G. CMC IP 42679, x 0.3. A - C , E, G from Warn and Strimple (1977,
plate 3, figures 1, 2, plate 6, figure 1, plate 13, figure 3, plate!4, figure 8) and reprinted by permission of
the Paleontological Research Institution.

Echinoderms 173
Figure 12.7. Cincinnatian disparid crinoids. A. locrinus subcrassus (Meek and Worthen), MUGM 28048,
horizon and locality unknown, x 1.4. B-D. Anomalocrinus. B. A. sp, CMC IP 170, Bellevue-Corryville
Formation, Cold Spring, Kentucky, x 1.8. C. Arms of A. incurvus Meek and Worthen, CMC IP, uncata-
logued, Cincinnati, OH, x 1.4. D. A. sp, CMC IP 7341, Cincinnati, Ohio.

174 A Sea without Fish

 Figure 12.8. Cincinnatian
monobathrid camerate
crinoids. A. Glyptocri-
nus decadactylus Hall,
MUGM 28046, Fairview
Formation (?), locality
unknown, x 1.5. B.
Pycnocrinus dyeri
(Meek), USNM 40762,
Corryville Formation,
Morrow, Warren Co.,
Ohio, x 1.2.

Echinoderms 175
Figure 12.9. Cincinnatian monobathrid camerate crinoids. A. Glyptocrinus fornshelli Miller, MUGM
28121, Liberty Formation, Butler Co., Ohio, x 7.7. B. Pycnocrinus dyeri (Meek) with attached gastropod,
Cyclonema sp., MUGM uncatalogued, Arnheim Formation, Dent, Hamilton Co., Ohio, x 2.0. C. P. dyeri,
Wayne State University Collection, Arnheim Formation, Dent, Hamilton Co., Ohio, lens cap 55 mm diam-
eter. Note remarkable preservation of articulated crowns, some splayed oralside down.

Figure 12.10. A. Cladid crinoid, Merocrinus curtus Ulrich, No. 366, Bruce and Charlotte Gibson Collec-
tion, Kope Formation, Hamilton Co., Ohio, x 4.2. B. Xenocrinus baeri (Meek), MUGM 28347, Liberty
Formation, southwestern Ohio, x 7.2. C. Diplobathrid camerate crinoid, Gaurocrinus nealli (Hall) USNM
S9I, Waynesville Formation, east of Lebanon, Warren Co., Ohio, x 1.3. This specimen was illustrated by
Meek (1873, plate 2, figure 3a).

176 A Sea without Fish

Echinoderms 177
Figure 12.11. Cincinnatian of c r i n o i d s , the disparids, the c l a d i d s , a n d tlic c a m c r a t c s . Disparids have a
rhombiferans. A, B. Le- s m a l l , m o n o c y c l i c c u p - o r h o w l - s h a p e d c a l y x w i t h b r a n c h i n g a r m s that d o
padocystis moorei
n o t b e a r p i n n u l e s . An e l o n g a t e , plated t u b e , the a n a l sac. is often present
(Meek), CMC IP 24680,
b e t w e e n the a r m s a n d has the a n a l o p e n i n g a t its e n d . T h e m o s t c o m m o n
Elkhorn Formation, Pre-
Cincinnatian c r i n o i d s , Cincinnaticrinus, Ectenocrimts, and locrinus are dis-
ble Co., Ohio, x 4.3.
C. Cheirocystis fulton- parids ( F i g u r e s 12.6,12.7; Plate 10). Cincinnaticrinus a n d probably Ectenocri-
ensis Sumrall and Schu- mis h a d a u n i q u e , button-like holdfast c o m p o s e d of m a i n tiny plates, often
macher, holotype, CMC f o u n d a t t a c h e d to b r a c h i o p o d shells a n d o t h e r hard substrata. Before it was
IP 50402, Kope Forma- r e c o g n i z e d that this holdfast b e l o n g s to these types of crinoids, if was given
tion, Bracken Co., Ken- t h e n a m e Lichenocrinus w i t h n u m e r o u s s p e c i e s ( F i g u r e s 1 2 . 5 L - N ; Faber
tucky, x 3.5. D. Recon-
1929; W a r n a n d S t r i m p l e 1977). Q u i t e often in p a l e o n t o l o g y isolated parts of
struction of late
o n e o r g a n i s m are d e s c r i b e d as distinct s p e c i e s before sufficiently well pre-
Riehmondian sea floor of
served fossils are f o u n d that reveal the entire a n i m a l .
southeastern Indiana and
southwestern Ohio, Cincinnaticrinus and Ectenocrinus are frequently found together in
showing Lepadocystis t h e K o p e F o r m a t i o n , w h e r e their d i s a r t i c u l a t e d c o l u n m a l s c a n form entire
moorei (A) attached to l i m e s t o n e b e d s . S o m e t i m e s t h e a r t i c u l a t e d stalks a r e p a c k e d tightlv to-
bryozoans (B), brachio- g e t h e r like l o g j a m s w h e r e t h e c o l l e c t o r s h o u l d l o o k c l o s e l y for t h e s m a l l ,
pods, Zygospira mod- d e l i c a t e c r o w n s ( Plate 10B). T h e s e " l o g j a m s " w e r e p r o b a h l v f o r m e d d u r i n g
esta (C), and an edrioast-
a n c i e n t s t o r m s that d i s r u p t e d t h e sea floor.
eroid, Carneyella sp. (D).
locrinus is a n o t h e r c o m m o n disparid c r i n o i d in t h e C i n c i n n a t i a n , but
A, B, D from Kesling and
Mintz (1961, plate 6, fig- is larger a n d m o r e robust in s t r u c t u r e than (lincinnaticrinus and Ectenocri-
ures 8, 10, plate 7) and nus ( F i g u r e 12.7A; Plate 10A). locrinus has a low, c o n i c a l c u p with dec]) in-
reprinted by permission d e n t a t i o n s m a r k i n g the j u n c t i o n of e a c h basal plate with the two radials
of the Museum of Pale- a b o v e . T h e c o l u n m a l s are p e n t a g o n a l or star-shaped ( F i g u r e s 12.5C, J). A d u l t
ontology, University of locrinus a t t a c h e d to b r y o z o a n s by c o i l i n g t h e stalk tightly a r o u n d the
Michigan.
b r a n c h e s . Slabs c o v e r e d w i t h locrinus c r o w n s h a v e b e e n f o u n d in the u p p e r
C o r r y v i l l e F o r m a t i o n . In o n e e a s e , t h e c r o w n s are c o n c e n t r a t e d at the base
of t h e fine-grained l i m e s t o n e b e d a n d are a l i g n e d parallel to a narrow gutter
e r o d e d into the sea floor ( F i g u r e s 12.^B; M e y e r el al. 19(81). C u r v e d l e n g t h s of
the locrinus stalks c o v e r the u p p e r surface of the bed in a r a n d o m fashion.
These preservational features s u g g e s t that the c r o w n s were s n a p p e d off the
stalks by a violent d i s t u r b a n c e (possibly storm-related) and swept d o w n s l o p e ,
to he f o l l o w e d by their stalks that settled on top of t h e m .
Anomalocrinus w a s t h e g i a n t a m o n g t h e c r i n o i d s o f t h e C i n c i n n a t i a n
( F i g u r e s 1 2 . 7 B - D ) . Its s t e m r e a c h e d a l e n g t h o f a b o u t o n e m e t e r , with c o -
l u n m a l s o v e r a c e n t i m e t e r in d i a m e t e r ( F i g u r e 12.5F). The c r o w n was at
least 10 cm h i g h a n d had up to 500 b r a n c h e s . The s t e m w a s a t t a c h e d by
m e a n s o f a s t u m p - l i k e holdfast c e m e n t e d d i r e c t l y t o hard substrata. T h e
holdfasts are f o u n d a s w o r n , crater-like l u m p s e n c r u s t i n g l i m e s t o n e n o d -
u l e s , h a r d g r o u n d s , b r y o z o a n s , o r shells.
C l a d i d s h a v e a c o n i c a l , d i e v c l i c c u p . Mcrocrinus is restricted to t h e
l o w e r m o s t K o p e F o r m a t i o n w h e r e its s t e m s are easily r e c o g n i z e d b y their
t h i n , wafer-like c o l u n m a l s ( F i g u r e s 12.5E, 12.10A). Cupulocrinusand Plico-
dendrocrinus are c l a d i d s f o u n d in t h e R i e h m o n d i a n f o r m a t i o n s (Plate 11).
C a m e r a t e c r i n o i d s h a v e a rigid c o n i c a l calyx that i n c l u d e s m a i n ' fixed
b r a c h i a l s , that is, a r m plates a b o v e the radials that are incorporated into the
c a l y x . T h e r e is a plated t c g i n e n c o v e r i n g the m o u t h . The p i n n u l e - b e a r i n g
free a r m s h a v e a feathery a p p e a r a n c e . ('dyptocrinus and Pycnocrinus, 1110110-

178 A Sea without Fish

Echinoderms 179
 Figure 12.12. Edrioaster-
oid Isorophus cincinna-
tiensis (Roemer), recon-
structed as in life, with
food grooves open for
feeding. Drawing by John
Agnew.

Figure 12.13. Cincinnatian

edrioasteroids A,
B. Isorophus cincinna-
tiensis (Roemer).
A. Corryville Formation,
Hamilton Co., Ohio, Uni-
versity of Cincinnati col-
bathrid c a m e r a t e s ( m o n o c y c l i c calyx), are very similar a n d are the m o s t c o m -
lection, x 2.2. B. Large
individual crowding m o n C i n c i n n a t i a n c a m e r a t e s ( F i g u r e 12.8). B o t h h a v e a distinct g e o d e s i c
against smaller individual, pattern of ridges on t h e c a l y x plates. Glyptocrinus decadactylus o c c u r s in the
Corryville Formation, l o w e r C i n c i n n a t i a n K o p e a n d F a i r v i e w F o r m a t i o n s and has t w o s e c u n d i -
Boone Co., Kentucky, braehs (calyx plates f o l l o w i n g the first b r a n c h i n g of a rav) and twenty free
University of Cincinnati
a r m s . Pycnocrinus o c c u r s in t h e C o r r y v i l l e , A r n h e i m , and W a y n e s v i l l e For-
collection, x 3. C-E.
m a t i o n s a n d has several s e e u n d i b r a c h s and ten free a r m s that b r a n c h o n c e
Carneyella pilea
n e a r t h e base, p r o d u c i n g a total of t w e n t y a r m s . B o t h Glyptocrinus and Pyc-
(Hall). C. CMC IP
34537, x 3.2. D, E. At- nocrinus c o i l e d t h e s t e m a r o u n d o b j e c t s s u c h as b r y o z o a n s for a t t a c h m e n t

tached to crinoid column, ( F i g u r e 12.4). O c c a s i o n a l l y , m u d s stirred u p d u r i n g storms s m o t h e r e d dense

CMC IP 26324, x 2.1. a g g r e g a t i o n s of these c r i n o i d s . A s p e c t a c u l a r Pycnocrinus a g g r e g a t i o n of this
F. Streptaster vorticel- type was found in the A r n h e i m Formation at D e n t , Hamilton C o u n t y ,
latus (Hall), CMC IP a r o u n d i 9 6 0 ( F i g u r e 1 2 . 9 C ) , a n d recently a d e n s e a g g r e g a t i o n of h u n d r e d s
24700, x3.2. G. Cys-
of Glyptocrinus, p r e s e r v e d c o m p l e t e w i t h a r m s , p i n n u l e s , and attached stalks
taster stellatus (Hall),
w a s r e c o v e r e d f r o m t h e F a i r v i e w F o r m a t i o n n e a r M a y s v i l l e , K e n t u c k y (in
CMC IP 40481, x 6.2.
preparation at t h e C i n c i n n a t i M u s e u m C e n t e r ) . Glyptocrinus fornshelli is
Photos C - G from Bell
(1976a, plate 16, figure very d i s t i n c t f r o m G. decadactylus in h a v i n g finer ridges on the calyx plates
10, plate 17, figures 1, 3, a n d u n i q u e t h i n , p e n t a g o n a l c o l u m n a l s ( F i g u r e 12.9A).
plate 10, figure14, plate Xenocrinus is a n o t h e r m o n o b a t h r i d c a m e r a t e crinoid f o u n d in the L i b -
13, figure 7) and re- erty F o r m a t i o n of t h e R i c h m o n d i a n S t a g e ( F i g u r e 12.10B). T h i s crinoid is
printed by permission of
n o t e w o r t h y for h a v i n g four-sided c o l u m n a l s and also h a v i n g the ability to coil
the New York State Mu-
a r o u n d o t h e r objects ( D o n o v a n et al. 1995). D i p l o b a t h r i d camerates have an
seum, Albany, N.Y.,
additional circlet of plates, the infrabasals, at the base of the calyx and are quite
12230.
rare in the C i n c i n n a t i a n . Gaurocrinus nealli is s h o w n in F i g u r e 12.10C.

180 A Sea without Fish

Echinoderms 181
Figure 12.14. Cincinnatian Associations of Crinoids with O t h e r Species
asteroids. A, B. Pro-
Interesting i n f o r m a t i o n a b o u t the e c o l o g y o f C i n c i n n a t i a n crinoids c a n
mopalaeaster specio-
sus (Meek). A. Oral b e g a i n e d b y o b s e r v i n g c l o s e associations b e t w e e n crinoids a n d other o r g a n -
side. B. Aboral side, isms. A s m e n t i o n e d a b o v e , s o m e C i n c i n n a t i a n crinoids used other o r g a n i s m s
holotype, Maysvillian, o n t h e sea f l o o r , u s u a l l y b r y o z o a n s , for a t t a c h m e n t . T h i s habit a l l o w e d the
Cincinnati, Ohio, MCI c r i n o i d s to g a i n e l e v a t i o n a b o v e t h e sea floor that e x p o s e d t h e m to u n r e -
108059, x 2. C, D. Pro-
stricted c u r r e n t flow, an a d v a n t a g e in passive suspension f e e d i n g a n d a m e a n s
mopalaeaster magnifi-
o f a v o i d i n g c o m p e t i t o r s . C r i n o i d s w e r e thus m e m b e r s o f s o m e o f the earliest-
e s (Miller) C. Oral
k n o w n " t i e r e d " m a r i n e b o t t o m c o m m u n i t i e s , i n w h i c h p a r t i c u l a r species,
side. D. Aboral side,
holotype, USNM 40883, e s p e c i a l l y s u s p e n s i o n feeders, o c c u p y preferred levels or tiers a b o v e the sub-

Richmondian, Waynes- s t r a t u m . C r i n o i d s h a v e b e e n " u p p e r story" o c c u p a n t s o f tiered c o m m u n i t i e s

ville, Warren Co., Ohio, x from O r d o v i c i a n t i m e up t h r o u g h the present (Ausich and Bottjer 1982).
0.8. Photos courtesy of Man\- C i n c i n n a t i a n s p e c i e s i n t u r n u s e d c r i n o i d s a s a t t a c h m e n t sites.
Jon W. Branstrator. C r i n o i d s t e m s are f r e q u e n t l y e n c r u s t e d w i t h b r v o z o a n s , c o r a l s , c o r n u l i t i d
w o r m t u b e s , b r a c h i o p o d s , e d r i o a s t e r o i d e c h i n o d e r m s ( F i g u r e s 12.13D, F ) ,
a n d e v e n o t h e r c r i n o i d holdfasts. B e c a u s e t h e s e e n c r u s t e r s often e n c i r c l e
t h e stalk, it is likelv s o m e w e r e a t t a c h e d d u r i n g t h e life of t h e c r i n o i d ,
p r o v i d i n g t h e e n c r u s t e r s t h e a d v a n t a g e o f a h i g h e r t i e r i n g level.
Pits, b o r i n g s , a n d gall-like s w e l l i n g s in c r i n o i d stems indicate that other
o r g a n i s m s u s e d t h e s t e m s as d w e l l i n g sites or e v e n parasitized t h e crinoid
host. S t e m s of Cincinnaticrinus s o m e t i m e s h a v e gall-like s w e l l i n g s p e n e -
trated by a pit. C a l c i u m p h o s p h a t i c rings found either w i t h i n the pit or on
the surfaces o f u n p i t t e d c o l u m n a l s ( W a r n 1974) w c r c
probably f o r m e d b y the
o r g a n i s m m a k i n g t h e pits, a l t h o u g h its identification is disputed. W a r n (1974)
c o n c l u d e d that t h e O r d o v i c i a n d e f o r m i t i e s a n d rings w e r e c a u s e d b y m y z o s -
t o m i d a n n e l i d w o r m s that also form galls in living crinoids. W e l c h (1976)
p o i n t e d o u t that n o m o d e r n m y z o s t o m i d s form p h o s p h a t i c l i n i n g s w i t h i n
their pits. H e r e c o g n i z e d that t h e O r d o v i c i a n s t r u c t u r e s r e s e m b l e others
f o u n d i n y o u n g e r P a l e o z o i c c r i n o i d s t e m s a n d thus c o u l d b e assigned t o the
g e n u s Phosphannulus. B i s c h o f f (1989) c o n s i d e r e d Phosphannulus to be a
junior s y n o n y m of Byronia, w h i c h b e l o n g s to a g r o u p of p h o s p h a t i c and/or
o r g a n i c t u b e - s h a p e d fossils (byroniids) f o u n d i n C a m b r i a n t h r o u g h P e r m i a n
strata. A l t h o u g h s o m e w o r k e r s interpret b y r o n i i d s as t u b e s of tiny s u s p e n s i o n
f e e d i n g w o r m s , B i s c h o f f a r g u e d that the}' are s h e a t h s of the p o l y p o i d larval
stage o f s c y p h o z o a n jellyfish. ( W e discussed the association b e t w e e n crinoids
a n d g a s t r o p o d s [ F i g u r e 12.9I in c h a p t e r 9.)
T h a n k s to m o d e r n o c e a n o g r a p h y , m a r i n e biologists c a n e x a m i n e
m a n y l i v i n g d e e p sea a n i m a l s s u c h a s stalked c r i n o i d s that w e r e p r e v i o u s l y
i n a c c e s s i b l e e x c e p t b y d r e d g i n g . O n e o f t h e m o s t s u r p r i s i n g r e c e n t discov-
eries a b o u t l i v i n g c r i n o i d s w a s o c c a s i o n a l c o l u m n s l a c k i n g c r o w n s s t a n d i n g
in t h e m i d s t of d e e p sea c r i n o i d " g a r d e n s " ( C o n a n et al. 1981). A f u r t h e r
s u r p r i s e c a m e w h e n J a p a n e s e w o r k e r s d i s c o v e r e d that " h e a d l e s s " c o l u m n s
o f R e c e n t Metacrinus c a n live i n d e f i n i t e l y i n a q u a r i a a n d e v e n r e g e n e r a t e
t h e c r o w n i f r e g e n e r a t i o n o c c u r s a b o v e t h e basal circlet o f plates ( A m e m i y a
a n d O j i 1992). T h i s s u r v i v a l a n d r e g e n e r a t i o n i s p r e s u m a b l y e n a b l e d b y
d i r e c t a b s o r p t i o n o f d i s s o l v e d n u t r i e n t s . F o s s o f the c r o w n s a m o n g wild
p o p u l a t i o n s of c r i n o i d s is m o s t likelv the result of p r e d a t i o n .

\82 A Sea without Fish

 C i n c i n n a t i a n fossil c r i n o i d s p r o v i d e s o m e o f t h e earliest e v i d e n c e t h a t
predators inflicted s i m i l a r d a m a g e o n O r d o v i c i a n c r i n o i d s a n d t h a t r e g e n -
eration w a s a survival m e c h a n i s m . In a s t u d y of t h e i m p a c t of p r e d a t i o n on
P a l e o z o i c c r i n o i d s , B a u m i l l e r a n d C a l m (2004) illustrated a r e m a r k a b l e
s p e c i m e n of the disparid Dystactocrinus comtrictus ( H a l l ) f r o m t h e C i n -

Echinoderms 183
Figure 12.15. Cincinnatian asteroids. A, B. Lanthanaster intermedius (Schuchert). A. Aboral
side. B. Oral side, holotype, FMNH 9575, Maysvillian, Cincinnati, Ohio, x 3. C. Petraster speciosus
(Miller and Dyer), holotype, MCZ 108063, Maysvillian, Preble Co., Ohio, x 7. D. Promopalaeaster dyeri
(Meek), MUGM 29664, with arms wrapped around bivalve in presumed feeding posture, Waynesville For-
mation, Brookville, Franklin Co., Indiana. E. Salteraster grandis (Meek), USNM 40885, Richmondian,
Waynesville, Warrern Co., Ohio, x 3. Photos A, B courtesy of Jon W. Branstrator.

184 A Sea without Fish

Figure 12.16. A. Asteroid, Hudsonaster simplex (Miller and Dyer), USNM 40882, Richmondian, near
Waynesville, Warren Co., Ohio, x 3.1. B. Taeniaster spinosus (Billings), USNM, Middle or Upper Ordovi-
cian, southern Pennsylvania, x 1.4. C. Cyclocystoid, Zygocycloides magnus (Miller and Dyer). B, from
Boardman et al. (1987, figure 18.45B), courtesy of James Sprinkle, and reprinted by permission of Blackwell
Publishing; C, courtesy of Colin Sumrall.

Echinoderms 185
Figure 12.17. Cincinnatian c i n n a t i a n i n w h i c h all o f t h e a r m s a r e i n t h e p r o c e s s o f r e g e n e r a t i o n ( F i g -
stylophoran carpoids. u r e 12.6D). T h e y c o n c l u d e d that t h e r e g e n e r a t i o n f o l l o w e d t h e loss o f t h e
A. 1-6. Enoploura p o -
a r m s t o a n a t t a c k b y a n u n k n o w n predator. A u s i c h a n d B a u m i l l e r (1993)
p e / Caster. 1 -3. Holo-
r e p o r t e d r e g e n e r a t i o n of a c o l u m n a t t a c h e d to t h e holdfast Lichenocrinus
type, CMC IP 25993,
dubius ( k n o w n to be t h e h o l d f a s t of Cincinnaticrinus or o t h e r disparids).
Corryville Formation,
Clermont Co., Ohio. D o n o v a n a n d S c h m i d t (2001) illustrated p l u r i c o l u m n a l s (sections o f several
1. Ventral view. 2. Dor- c o l u m n a l s ) o f Cincinnaticrinus s h o w i n g r o u n d e d o v e r g r o w t h s o f o n e e n d .
sal view. 3. Lateral T h e y s u g g e s t e d t h a t t h e o v e r g r o w t h s f o r m e d after d e c a p i t a t i o n o f t h e
view, x 2.3. 4-6. Para- c r o w n s b y p r e d a t i o n , l e a v i n g a " h e a d l e s s " c o l u m n . I n light o f t h e n e w
type, CMC IP 25257, Cor- k n o w l e d g e of p r e d a t i o n d a m a g e in l i v i n g c r i n o i d s , it is m o s t likely that
ryville Formation, Hamil-
p r e d a t o r s a l s o c a u s e d loss o f c r o w n s a n d a r m s i n O r d o v i c i a n c r i n o i d s , al-
ton Co., Ohio. 4. Dorsal
t h o u g h the identity of the culprits remains uncertain.
view. 5. Lateral view.
6. Ventral view, x 2.3.
1-6 from Caster (1952, Rhombiferan "cystoids"
plate 1). B. Reconstruc-
tion of E. popei, from R h o m b i f e r a n s are stalked e c h i n o d e r m s that a p p e a r e d i n the Early O r d o v i -
Parsley (1991, text-figure cian and b e c a m e extinct by the Late D e v o n i a n . T h e term " c y s t o i d " (sac-
1). All reprinted by per- like) refers to t h e plated t h e c a that has four to five circlets of large plates
mission of the Paleonto- a r r a n g e d in a p e n t a m e r a l p a t t e r n . R h o m b i f e r a n s w e r e o r i g i n a l l y a s u b -
logical Research
g r o u p o f t h e cystoids b u t h a v e b e e n e l e v a t e d t o a separate class. T h e y lived
Institution.
as suspension feeders, but unlike crinoids, the feeding appendages of
r h o m b i f e r a n s are t h i n p l a t e d b r a c h i o l c s a r i s i n g f r o m a m b u l a c r a that are
e i t h e r i n c o r p o r a t e d into t h e t h e c a o r less c o m m o n l y stand e r e c t ( F i g u r e
12.11C). R h o m b i f e r a n s are n a m e d for a u n i q u e r h o m b i c s t r u c t u r e f o u n d
on t h e t h e c a l plates. A set of n a r r o w slits in a r h o m b i c o u t l i n e crosses
b o u n d a r i e s o f a d j a c e n t p l a t e s ; t h e s e slits led t o i n t e r c o n n e c t i n g i n t e r n a l
c a n a l s . T h e p u r p o s e o f t h e s e d i s t i n c t i v e c a n a l s w a s t o p r o v i d e w a t e r flow
t h r o u g h t h e interior o f t h e t h e c a for respiratory p u r p o s e s . T h e relatively
short b r a c h i o l e s o f r h o m b i f e r a n s m a y h a v e c a r r i e d f e w e r t u b e feet t h a n
t h o s e u s e d for r e s p i r a t i o n i n c r i n o i d s , o r m a y h a v e l a c k e d t h e m entirely.
T h e r e f o r e t h e h e a v i l y p l a t e d r h o m b i f e r a n s m a y h a v e n e e d e d a different
m e a n s t o s u p p l y o x y g e n a t e d w a t e r a n d t o r e m o v e dissolved c a r b o n d i o x i d e
f r o m t h e t h e c a l interior.
T w o s p e c i e s o f r h o m b i f e r a n s are f o u n d i n t h e C i n c i n n a t i a n . Cheiro-
cystis fultonensis is restricted to t h e l o w e r m o s t K o p e Formation (Fulton
S h a l e ) ( S u m r a l l a n d S c h u m a c h e r 2002). T h i s s p e c i e s has a n e l o n g a t e t h e c a
a n d a l o n g , t a p e r i n g , a t t a c h e d stalk ( F i g u r e 12.11C). Lepadocystis moorei
( M e e k ) is found only in the Elkhorn Formation (Richmondian). This
r h o m b i f e r a n h a s a r o u n d e d t h e c a a n d a shorter, t a p e r i n g u n a t t a c h e d stalk
( S u m r a l l a n d S p r i n k l e 1999; F i g u r e s 12.11A, B , D ) . S o m e c o m p l e t e s p e c i -
m e n s are still a t t a c h e d by a h o l d f a s t c e m e n t e d to v a r i o u s o b j e c t s .

E-drioasteroids

N e x t t o t h e c r i n o i d s , e d r i o a s t e r o i d s a r e t h e m o s t c o m m o n e c h i n o d e r m fos-
sils in C i n c i n n a t i a n strata. E d r i o a s t e r o i d s c o n s t i t u t e a class of e c h i n o d e r m s
that o r i g i n a t e d d u r i n g t h e C a m b r i a n a n d b e c a m e e x t i n c t i n the P e r m i a n .
At first g l a n c e , an e d r i o a s t e r o i d r e s e m b l e s a sea star w i t h a r i n g a r o u n d it

786 A Sea without Fish

Echinoderms 187
 (the n a m e m e a n s "seated-star"). T h e sea star r e s e m b l a n c e derives from the
five u s u a l l y c u r v i n g food g r o o v e s o r a m b u l a c r a ! tracts that c o n v e r g e o n the
c e n t r a l m o u t h ( F i g u r e 12.12). T h i n , o v e r l a p p i n g c a l c i t i c plates c a l l e d inter-
a m b u l a c r a l s take u p t h e s p a c e b e t w e e n t h e a m b u l a c r a . A m o r e rigid series
o f m a r g i n a l plates f o r m s t h e ring. U s u a l l y t h e a m b u l a c r a l a n d i n t e r a m b u -
lacral plates a p p e a r to h a v e c o l l a p s e d inside t h e m a r g i n a l r i n g ; thus it is
a s s u m e d that t h e m u l t i p l a t e d t h e c a was flexible in life. Rarely, u n c o l l a p s e d
o r " i n f l a t e d " s p e c i m e n s are f o u n d that reveal h o w the a n i m a l probably a p -
p e a r e d i n life. M o s t C i n c i n n a t i a n e d r i o a s t e r o i d s h a d a l o w d o m e s h a p e , b u t
s o m e w e r e m o r e c y l i n d r i c a l ( S u m r a l l 1994). A total of six g e n e r a a n d e l e v e n
s p e c i e s are k n o w n f r o m C i n c i n n a t i a n strata ( F i g u r e 12.13).
B e c a u s e e d r i o a s t e r o i d s are e x t i n c t , their m o d e o f life m u s t b e inferred
b y a n a l o g v t o l i v i n g e c h i n o d e r m s . F d r i o a s t e r o i d s are a l w a y s f o u n d a t t a c h e d
to a hard surface s u c h as a b r a c h i o p o d shell, b r y o z o a n , or hardground. T h e
lower surface was plated in s o m e species, but in C i n c i n n a t i a n species ap-
p a r e n t l y o n l y a soft tissue m e m b r a n e s e r v e d to a d h e r e to the s u b s t r a t u m ,
possibly i n t h e w a y sea a n e m o n e s a t t a c h b y their p e d a l disk. S o m e s p e c i e s
a c t u a l l y c e m e n t e d t o t h e s u b s t r a t u m like a b a r n a c l e , but those a d h e r i n g b y
a basal m e m b r a n e m a y have b e e n capable of limited m o v e m e n t . B e c a u s e
t h e m o u t h a n d a m b u l a c r a l tracts are d i r e c t e d u p w a r d s i n e d r i o a s t e r o i d s ,
t h e y a p p a r e n t l y l i v e d a s passive filter f e e d e r s . T h e a m b u l a c r a l g r o o v e s are
l i n e d w i t h tinv, z i p p e r - l i k e e o v e r p l a t e s c a p a b l e o f o p e n i n g a n d c l o s i n g (Fig-
u r e 12.12). O p e n i n g t h e e o v e r p l a t e s e x p o s e d t h e food g r o o v e l i n e d w i t h
t u b e feet and/or c i l i a that s e r v e d t o c o l l e c t s u s p e n d e d food p a r t i c l e s a n d
c o n v e y t h e m t o t h e m o u t h i n a m a n n e r s i m i l a r t o that o f l i v i n g c r i n o i d s .
Fecal waste was expelled t h r o u g h a valve-like anal o p e n i n g on the thecal
s u r f a c e . T w o o t h e r o p e n i n g s l o c a t e d n e a r t h e m o u t h are a s s u m e d t o b e a
g o n o p o r e (for r e l e a s e o f g a m e t e s ) a n d a h y d r o p o r e . The hydropore allowed
for w a t e r i n t a k e t o t h e w a t e r v a s c u l a r s y s t e m a n d for c o n t r o l o f t h e a m o u n t
o f t h e c a l i n f l a t i o n . C y l i n d r i c a l f o r m s s u c h a s t h e C i n c i n n a t i a n Streptaster
w e r e c a p a b l e o f t e l e s c o p i n g t h e t h e c a l plates d u r i n g inflation s o a s t o
a c h i e v e a n e l e v a t e d f e e d i n g p o s i t i o n a n d d u r i n g d e f l a t i o n a s s u m i n g a low-
profile, p r o t e c t i v e p o s i t i o n ( S u m r a l l 1994).
A l t h o u g h e d r i o a s t e r o i d s a r e u s u a l l y c o n s i d e r e d t o b e rare fossils, t h e y
c a n occasionally be found by the h u n d r e d s or thousands in C i n c i n n a t i a n
strata w h e n t h e a p p r o p r i a t e p r e s e r v a t i o n a l c o n d i t i o n s are p r e s e n t ( M e y e r
1990). T h i n l i m e s t o n e s c o v e r e d w i t h s t r o p h o m e n i d b r a c h i o p o d s (shell
p a v e m e n t s ) o r h a r d g r o u n d s ( c a l c a r e o u s s e d i m e n t s l i t h i h e d o n t h e sea f l o o r )
are t h e ideal substrata for e d r i o a s t e r o i d s ( W i l s o n 1985). D e s p i t e the a b u n -
d a n c e of these types of beds in the C i n c i n n a t i a n , edrioasteroid-bearing
p a v e m e n t s are rare. P r e s e r v a t i o n of e d r i o a s t e r o i d s in their life p o s i t i o n on
p a v e m e n t s r e q u i r e d a rapid, c a t a s t r o p h i c b u r i a l w i t h f i n e m u d . W i t h o u t
s u c h a rapid s m o t h e r i n g , t h e d e l i c a t e , m u l t i p l a t e d e d r i o a s t c r o i d t h e c a dis-
a r t i c u l a t e d s o o n after d e a t h , l e a v i n g little o r n o trace.
The d i s c o v e r y of p a v e m e n t s b e a r i n g a b u n d a n t edrioasteroids in the C i n -
c i n n a t i a n led to p i o n e e r i n g s t u d i e s of t h e life history of edrioasteroids and
their p o p u l a t i o n p a l e o e c o l o g v . Because pavements contained specimens
r a n g i n g in size from a few m i l l i m e t e r s in d i a m e t e r to the largest individuals

188 A Sea without Fish

over thirty m i l l i m e t e r s in d i a m e t e r , Bell (1976) was able to d e t e r m i n e how
several s p e c i e s c h a n g e d i n m o r p h o l o g y d u r i n g g r o w t h . T h i s i n f o r m a t i o n w a s
used by M e y e r (1990) to study t h e p o p u l a t i o n p a l c o e c o l o g y of t h r e e different
species f o u n d o n a s i n g l e p a v e m e n t . S m a l l i n d i v i d u a l s o f the c o m m o n spe-
cies Isorophus cincinnatiensis ( F i g u r e s 12.13A, B) clustered n e a r t h e m a r g i n s
of articulated (likely living) shells of t h e host b r a c h i o p o d Rafinesquina. T h e
small edrioasteroids m a y h a v e lived in a c o m m e n s a l r e l a t i o n s h i p w i t h the
living b r a c h i o p o d , t a k i n g a d v a n t a g e o f t h e f e e d i n g c u r r e n t s g e n e r a t e d b y t h e
b r a c h i o p o d and protection a l o n g the o v e r h a n g i n g m a r g i n o f the host shell.
B e c a u s e a single large Isorophus o c c u p i e s a l m o s t an entire b r a c h i o p o d shell,
it is likely that either m o r t a l i t y or r e l o c a t i o n to avoid o v e r c r o w d i n g d e p l e t e d
the juvenile clusters. In t i m e , t h e edrioasteroids m a y h a v e o u t l i v e d t h e bra-
c h i o p o d host, b e c a u s e t h e larger i n d i v i d u a l s are f o u n d o n d i s a r t i c u l a t e d ,
abraded ( h e n c e d e a d ) host shells. P a v e m e n t s f o u n d at different stratigraphic
h o r i z o n s i n t h e C i n c i n n a t i a n h a v e different edrioasteroid s p e c i e s p o p u l a -
tions ( S u m r a l l et al. 2001).

Asteroids

S e a stars are e x c e p t i o n a l l y rare fossils in t h e C i n c i n n a t i a n ; o f t e n , s p e c i m e n s

are hard to r e c o g n i z e b e c a u s e they are f r a g m e n t a r y or distorted. N e v e r t h e -
less, six g e n e r a a n d ten s p e c i e s are valid f r o m the C i n c i n n a t i a n ( F i g u r e s
12.14-12.16). S e a stars are t h e m o s t r e c o g n i z a b l e e c h i n o d e n n s , h a v i n g the
stereotypical pentaradial s y m m e t r y . they differ from brittle stars (ophiuroids)
in h a v i n g a r m s that are not sharply differentiated f r o m t h e c e n t r a l disk.
C i n c i n n a t i a n sea stars, like m o d e r n f o r m s (Plate 9 E ) , h a d a w i d e r a n g e
o f b o d y f o r m s that c a n b e related t o a diversity o f f e e d i n g habits. T h e best-
k n o w n C i n c i n n a t i a n sea star, Promopalaeaster, is f o u n d t h r o u g h o u t t h e
stratigraphic section. An extraordinary specimen of a Promopalaeaster
w r a p p e d a r o u n d a b i v a l v e m o l l u s c p r o v i d e s a rare e x a m p l e of a sea star
c a u g h t i n a n a n c i e n t act o f p r e d a t i o n ( F i g u r e 12.15D; B l a k e a n d G u e n s b u r g
1994). This c a s e d e m o n s t r a t e s that s o m e sea stars o f t h e O r d o v i c i a n h a d t h e
ability t o prey o n p e l e c y p o d s b y p r y i n g t h e i r v a l v e s a p a r t just a s d o m o d e r n
sea stars. T h e s t o m a c h is e v e r t e d t h r o u g h t h e m o u t h a n d t h e p r e v is d i -
g e s t e d w i t h i n its o w n s h e l l . B r a n s t r a t o r (1975) c o n c l u d e d that Mesopa-
laeaster w a s a l s o a s t o m a c h - e v e r t i n g sea star. Hudsonaster is a s m a l l sea star
with large, b l o c k y plates, at least s u p e r f i c i a l l y s i m i l a r to c e r t a i n p r e s e n t - d a y
m e m b e r s o f t h e G o n i a s t e r i d a e o r O p h i d i a s t e r i d a e , w h i c h f e e d o n passive,
c o l o n i a l o r g a n i s m s , s m a l l prey, a n d d e t r i t u s ( F i g u r e 12.16A; J a n g o u x 1982).
Another Cincinnatian asteroid, Petraster speciosus (Richmondian), has
short, broad a r m s a n d a flattened profile ( F i g u r e 12.15C). It is v e r y s i m i l a r
t o m o d e r n sea stars that live b y i n g e s t i n g s e d i m e n t a n d d i g e s t i n g o r g a n i c
materials ( B l a k e a n d G u e n s b u r g 1988). In c o n t r a s t , Salteraster h a d v e r y
l o n g , n a r r o w a r m s , w ith s m a l l ossicles ( F i g u r e 12.15E). S p e c i m e n s are often
p r e s e r v e d in a c o n t o r t e d m a n n e r s u g g e s t i n g great flexibility a n d ability to
handle small or large prey. Both Salteraster a n d Lanthanaster ( F i g u r e s
12.15A, B) are t h o u g h t to h a v e b u r r o w e d into fine s e d i m e n t in s e a r c h of

Echinoderms
 prey, a n d b o t h m a y h a v e p r o d u c e d the rare star-shaped b u r r o w s i n the
C i n c i n n a t i a n c a l l e d Asteriacites (see F i g u r e 14.3D; Branstrator 1975).

Ophiuroids
T h e o p h i u r o i d s (brittle stars o r s e r p e n t stars) a r e d i s t i n g u i s h e d from the
asteroids b y h a v i n g t h e f i v e a r m s sharply differentiated from t h e c e n t r a l
disk (Plate 9 C ) . T h e a r m s are q u i t e flexible b e c a u s e t h e y a r e c o m p o s e d o f
a series of vertebra-like ossicles c o n n e c t e d by m u s c l e s a n d l i g a m e n t s . O p h i -
uroids c a n m o v e q u i t e rapidly o n t h e sea floor b y l a s h i n g t h e a r m s and
s o m e c a n flex t h e a r m s v e r t i c a l l y a s w e l l . T h e a r m s are e q u i p p e d w i t h t u b e
feet that are u s e d to g a t h e r o r g a n i c p a r t i c l e s either d i r e c t l y from t h e sedi-
m e n t o r a s s u s p e n d e d p a r t i c l e s . O p h i u r o i d s a r e usually c o n s i d e r e d t o b e
d e p o s i t f e e d e r s , b u t s o m e are also c a p a b l e o f s u s p e n s i o n f e e d i n g a n d e v e n
predation. Taeniaster spinosus (Billings) occurs in the Cincinnatian
( I l o t c h k i s s 1970; F i g u r e 12.16B), a n d like asteroids, it is very rare. M o d e r n
o p h i u r o i d s c a n live in v e r y d e n s e a g g r e g a t i o n s on the sea floor. Rarely slabs
h a v e b e e n f o u n d in t h e C i n c i n n a t i a n b e a r i n g d e n s e a s s e m b l a g e s of Tae-
niaster, s u g g e s t i n g that a g g r e g a t i o n b e h a v i o r w a s a c h i e v e d in this very early
m e m b e r of the group. In the only other k n o w n C i n c i n n a t i a n ophiuroid,
Protasterina flexuosa, p y r i t i z e d t u b e feet h a v e b e e n reported in a s p e c i m e n
f r o m t h e K o p e F o r m a t i o n ( G l a s s 2006).

Cyclocystoids

T h e c y c l o c y s t o i d s are a m o n g the rarest a n d m o s t e n i g m a t i c o f C i n c i n n a -

tian e c h i n o d e r m s . T h e y c a n b e easily m i s t a k e n for a n edrioasteroid b e -
c a u s e t h e y h a d a c i r c l e t o f s q u a r i s h , m a r g i n a l plates, 7 - 2 0 m m i n d i a m e t e r ,
but lack t h e c h a r a c t e r i s t i c f i v e c u r v i n g a m b u l a c r a o f edrioasteroids ( F i g u r e
12.16C). A c o m p r e h e n s i v e study of c y c l o c y s t o i d s by S m i t h a n d Paul (1982)
p r o v i d e s t h e m o s t u p t o date u n d e r s t a n d i n g o f their c o m p l e x m o r p h o l o g y .
T h e skeleton of a c y c l o c y s t o i d c o n s i s t s of t h r e e parts: a c e n t r a l disk, a m a r -
g i n a l r i n g , a n d a p e r i p h e r a l skirt. U s u a l l y o n l y t h e m a r g i n a l r i n g is f o u n d ,
c o n s i s t i n g o f e i g h t e e n t o sixty s q u a r i s h o r b l o c k v ossicles. N u m e r o u s s m a l l
plates c o v e r o n e s u r f a c e o f t h e disk: g r o o v e d radial plates, u n g r o o v e d inter-
radial plates, a n d c o v e r plates c o v e r i n g t h e g r o o v e d radial plates. T h e r e is
a c e n t r a l o p e n i n g o n this s u r f a c e , p r e s u m a b l y the m o u t h . S m a l l p o l y g o n a l
plates c o v e r t h e o p p o s i t e s u r f a c e , w i t h a s m a l l c o n e at the c e n t e r , p r e s u m -
ably s u r r o u n d i n g t h e a n a l o p e n i n g . A l t h o u g h s o m e c y c l o c y s t o i d s h a v e f i v e -
fold s y m m e t r y o f t h e p l a t i n g , o t h e r s h a v e four-fold o r six-fold s y m m e t r y ,
b r e a k i n g the s t e r e o t y p i c a l e c h i n o d e r m p a t t e r n . The n a t u r e o f t h e p l a t i n g
s u g g e s t s t h a t i n life, t h e r e w a s v e r y little b o d y s p a c e b e t w e e n the t w o plated
s u r f a c e s . The m a r g i n a l ossicles h a v e a d i s t i n c t i v e f o r m a n d are perforated
b y tiny c a n a l s . E a c h m a r g i n a l has o n e t o seven c u p u l c s , w h i c h a r e s p o o n -
s h a p e d d e p r e s s i o n s . E a c h c u p u l e l e a d s to a radial d u c t l y i n g w i t h i n a cir-
c u m f e r e n t i a l g r o o v e . T h e p e r i p h e r a l skirt consists o f s m a l l , i m b r i c a t e plates
that c o v e r e d t h e r i n g o f c u p u l e s i n life.

190 A Sea without Fish

 B e c a u s e t h e r e a p p e a r s t o h a v e b e e n very little s p a c e b e t w e e n t h e
plated s u r f a c e s o f the l i v i n g c v c l o e y s t o i d , t h e a n i m a l s o m e w h a t r e s e m b l e d
a t a m b o u r i n e rather t h a n a d r u m . L a c k i n g internal s p a c e for o r g a n s , c y c l o -
cystoids w e r e restricted t o f e e d i n g o n m i n u t e o r g a n i c p a r t i c l e s . S m i t h a n d
Paul c o n c l u d e that the p a r t i c l e s w e r e c o l l e c t e d a t t h e c u p u l e s b y c i l i a r y
a c t i o n a n d c o n v e y e d via the d u c t s t o t h e radial g r o o v e s that c o n v e r g e d o n
t h e c e n t r a l m o u t h . A l t h o u g h t u b e feet are n o t p r e s e r v e d , S m i t h a n d P a u l
suggest that t u b e feet c o u l d h a v e e m e r g e d f r o m p o r e s b e t w e e n plates o n
the ventral s u r f a c e a n d p r o v i d e d a m e a n s o f l o c o m o t i o n . T h e e y c l o c v s t o i d
thus m o v e d over the substrate a n d g a t h e r e d o r g a n i c p a r t i c l e s u s i n g c i l i a t e d
c u p u l e s . O t h e r s p e c i a l i s t s , h o w e v e r , d o not a c c e p t t h e life o r i e n t a t i o n fa-
vored by S m i t h a n d Paul, a n d instead regard the o p p o s i t e side as l o w e r m o s t
( S p r i n k l e , pers. c o m m . ) C y c l o c y s t o i d s f i r s t a p p e a r e d i n t h e Farlv O r d o v i -
c i a n a n d last in the Late D e v o n i a n or Karlv C a r b o n i f e r o u s ( S m i t h a n d Paul
1982). A total of n i n e g e n e r a a n d f o r t y - o n e s p e c i e s h a v e b e e n d e s c r i b e d . In
the C i n c i n n a t i a n , three g e n e r a a n d f i v e s p e c i e s a r e k n o w n .

Stvlophorans

f o r m a n y p a l e o n t o l o g i s t s , s t v l o p h o r a n s surpass e v e n the c y c l o c y s t o i d s a s
s o m e o f t h e m o s t b i z a r r e fossil e c h i n o d e r m s . For a m i n o r i t y o f s p e c i a l i s t s ,
s t v l o p h o r a n s are c o n s i d e r e d not e v e n t o h e e c h i n o d e r m s , b u t rather a n c e s -
tors of t h e vertebrates b e l o n g i n g to a g r o u p c a l l e d c a l c i c h o r d a t e s . ( T h e
interested reader is referred to G e e 119961 for a b l o w - b y - b l o w a c c o u n t of
this debate.) This c o n t r o v e r s y , c o u p l e d w i t h their e x c e e d i n g rarity i n C i n -
c i n n a t i a n strata, m a k e the s t v l o p h o r a n s o n e o f the m o s t i n t r i g u i n g fossils
ever to be f o u n d in the C i n c i n n a t i r e g i o n . A s i n g l e g e n u s , Enoploura, is
f o u n d in the C i n c i n n a t i a n , w i t h t w o s p e c i e s , E. popei C a s t e r in the
M a v s v i l l i a n ( F i g u r e 12.17) a n c
' balanoides 1 M e e k ) in the Mavsv illian a n d
R i c h m o n d i a n ( C a s t e r 19S2; Parsley 1991).
Enoploura is t y p i c a l of the s t v l o p h o r a n s , an e x t i n c t e c h i n o d e r m class
that r a n g e s from t h e M i d d l e C a m b r i a n t h r o u g h t h e E a r l y P e n n s v l v a n i a n ,
with a b o u t e i g h t y g e n e r a in all. Enoploura has a flattened, p l a t e d t h e c a that
i s r o u g h l y r e c t a n g u l a r ; the p l a t i n g has n o radial o r p e n t a m e r a l s y m m e t r y
w h a t s o e v e r , hut rather is bilaterally s y m m e t r i c a l . A p a i r of s p i n e s is at-
tached at one end, and at the opposite end, a single, s e g m e n t e d a p p e n d a g e
is a t t a c h e d . This a p p e n d a g e has a very c o m p l e x s t r u c t u r e a n d its f u n c t i o n
has b e e n v i g o r o u s l y d e b a t e d . Il was o n c e t h o u g h t to be a stalk-like h o l d f a s t ,
but those f a v o r i n g the e c h i n o d e r m affinity of (he s t v l o p h o r a n s c o n s i d e r it
to be a feeding a p p e n d a g e , called the a u l a c o p h o r e . T h e basal p a r t o f t h e
a u l a c o p h o r e i s m a d e u p o f a series o f t h i n e l e m e n t s e a c h c o n s i s t i n g o f four
c o m p o n e n t s ; il was p r o b a b l y flexible. F o l l o w i n g this flexible r e g i o n is t h e
s o - c a l l e d styloid, b e a r i n g a pair of b l a d e l i k e flanges that p r e s u m a b l y d u g
into the s e d i m e n t . The t e r m i n a l p a r t o f t h e a u l a c o p h o r e has a t a p e r i n g
series o f sharply k e e l e d ossicles b e a r i n g a g r o o v e w i t h c o v e r i n g plates. T w o
possible f e e d i n g positions h a v e b e e n p o s t u l a t e d for t h e a u l a c o p h o r e : h e l d
up into the water or a r c h e d slightly a b o v e t h e s u b s t r a t u m . In a n y c a s e t h e
food consisted of very tine o r g a n i c p a r t i c l e s . Particles taken in a l o n g a food

Echinoderms 191
 groove entered an internal m o u t h , and wastes were emitted through an
a n a l o p e n i n g b e t w e e n t h e s p i n e s . P r o p o n e n t s o f t h e e a l c i c h o r d a t e inter-
p r e t a t i o n of s t v l o p h o r a n s regard t h e a p p e n d a g e to be a t r u e , w r i g g l i n g tail,
with the m o u t h located at the opposite end of the theca. Study of well-
p r e s e r v e d skeletal m i c r o s t r u c t u r e in C i n c i n n a t i a n V.noploura by C a r l s o n
a n d f i s h e r (1981) r e v e a l e d c l o s e s i m i l a r i t y t o t y p i c a l e c h i n o d e r m s t c r e o m ,
further supporting the classification of stvlophorans with e c h i n o d e r m s .
S o m e e n i g m a t i c fossils c a l l e d m a e h a e r i d i a n s m i g h t also b e related t o stv-
l o p h o r a n s (see c h a p t e r 10).

192 A Sea without Fish

194 A Sea without Fish
GRAPTOLITES AND CONODONTS:
OUR CLOSEST RELATIVES?

G r a p t o l i t e s are a m o n g t h e m o s t d i s t i n c t i v e fossils f o u n d i n C i n c i n n a t i a n Graptolites

strata a n d are also u n i q u e l y s i g n i f i c a n t . C r a p t o l i t e s are c o m m o n l y pre-
served in shales in a h i g h l y flattened c o n d i t i o n , a p p e a r i n g like b l a c k p e n c i l
m a r k i n g s w i t h a s a w - t o o t h e d m a r g i n (the n a m e g r a p t o l i t e i n fact m e a n s
" w r i t t e n stone"; F i g u r e 1 3 . l C ) . I n s o m e C i n c i n n a t i a n l i m e s t o n e s g r a p t o l i t e s
can be preserved in an u n c o m p a c t e d , three-dimensional condition. B e -
c a u s e their skeletal s t r u c t u r e ( p e r i d e r m ) i s o r g a n i c t h e s e " i n f l a t e d " g r a p t o -
lites c a n b e e t c h e d free o f t h e m a t r i x u s i n g a c i d t o r e v e a l e x c e p t i o n a l s t r u c -
tural details ( F i g u r e 13.1B). G r a p t o l i t e s r e p r e s e n t t h e skeletal s h e a t h of a
c o l o n i a l , soft-bodied m a r i n e invertebrate w h o s e soft parts are n o t p r e s e r v e d .
G r a p t o l i t e c o l o n i e s existed a s f r e e - f l o a t i n g p l a n k t o n (order G r a p t o l o i d e a )
o r a s b r a n c h i n g , b e n t h i c c o l o n i e s (order D e n d r o i d e a ) . T h e c o l o n i a l skele-
ton ( r h a b d o s o m e ) h o u s e d m a n y s o f t - b o d i e d z o o i d s e a c h w i t h i n a c u p - l i k e
t h e c a ( F i g u r e 13.1A). T h e w a l l s o f t h e t h e c a e are c o n s t r u c t e d i n a u n i q u e
w a y that i s o f g r e a t i m p o r t a n c e i n e s t a b l i s h i n g t h e n a t u r e o f t h e g r a p t o l i t e Figure 13.1. A. Grapto-
o r g a n i s m : narrow h a l f - r i n g s (fusellae) a l t e r n a t e t o f o r m a z i g z a g s u t u r e lite morphology. Drawing
a l o n g the t h e c a l t u b e . A n o u t e r c o r t i c a l layer o f c o l l a g e n fibrils r e i n f o r c e s by Kevina Vulinec. B,
the fusellar layer b y c r i s s - c r o s s i n g t h e s u r f a c e ; h e n c e t h e s e are c a l l e d c o r t i - C. Geniculograptus
cal b a n d a g e s . I n p l a n k t i c g r a p t o l o i d s , t h e c a e are a r r a n g e d a s b r a n c h e s o r typicalis (Hall). B. Sin-

stipes either in a s i n g l e l i n e a r series ( m o n o s e r i a l ) , a d o u b l e Series (biserial), gle rhabdosome, courtesy

of Daniel Goldman.
or e v e n triserial or q u a d r i s e r i a l . S t i p e s w e r e a t t a c h e d s i n g l y or in m u l t i p l e s
C. Cluster of partly paral-
by t h e t h r e a d - l i k e n e m a to v a n e - l i k e s t r u c t u r e s or p o s s i b l y to gas-filled
lel-oriented rhabdosomes,
bulbs that p r o v i d e d flotation. A l t h o u g h i t w a s o n c e t h o u g h t that g r a p t o l o i d s
MUGM 29469, Cincinna-
attached by m e a n s of the n e m a to other floating material like s e a w e e d , it
tian, Butler Co., Ohio,
i s n o w g e n e r a l l y a c c e p t e d that p l a n k t i c g r a p t o l o i d s u s e d t h e i r o w n m e a n s scale in mm.
of flotation, b u t t h e r e is d e b a t e as to w h e t h e r flotation w a s a c t i v e l y or p a s -
sively m a i n t a i n e d ( R i g b y a n d Fortey 1991).
T h e f l o a t i n g ability o f g r a p t o l o i d s c a u s e d t h e m t o b e c a r r i e d g r e a t
distances by o c e a n currents. C o n s e q u e n t l y a single graptolite species c a n
b e f o u n d o v e r a vast g e o g r a p h i c a r e a , e v e n o n s e p a r a t e c o n t i n e n t s . T h i s
w i d e d i s t r i b u t i o n , c o u p l e d w i t h t h e i r relatively rapid e v o l u t i o n a r y c h a n g e
over t i m e , m a k e graptolites s o m e o f t h e m o s t ideal i n d e x fossils for biostrati-
g r a p h i c z o n a t i o n a n d c o r r e l a t i o n o f strata. G r a p t o l i t e s f i r s t a p p e a r e d i n t h e
Middle C a m b r i a n and b e c a m e extinct as a group in the Pennsylvanian,
b u t for O r d o v i c i a n a n d S i l u r i a n strata in p a r t i c u l a r , g r a p t o l i t e s ( a l o n g w i t h
c o n o d o n t s ) p r o v i d e t h e e s s e n t i a l basis for relative a g e d a t i n g a n d c o r r e l a -
tion w o r l d w i d e . The U p p e r O r d o v i c i a n o f N o r t h A m e r i c a i s d i v i d e d into
four graptolite b i o z o n e s o n t h e basis o f o v e r l a p p i n g r a n g e s o f several s p e c i e s
( G o l d m a n a n d B e r g s t r o m 1997).

195
Figure 1 3 . 2 . Cincinnatian \ l t h o u g h several s p e c i e s of graptolites o c c u r in the C i n c i n n a t i a n of
conodonts. Those shown t h e C i n c i n n a t i A r c h r e g i o n , o n l y a few are c o m m o n ( B e r g s t r o m 1997). The
are among the most m o s t c o m m o n a n d c h a r a c t e r i s t i c C i n c i n n a t i a n graptolite i s Geniculogrcip-
common forms. All are
tus (identified as Climacograptus in o l d e r literature; f i g u r e s 13.1B, C ) . T h i s
from the lower Riehmon-
g r a p t o l i t e has a hiserial r h a b d o s o m e a n d is very c h a r a c t e r i s t i c of t h e K o p e
dian Stage near
f o r m a t i o n , w h e r e a g g r e g a t i o n s o f stipes often o c c u r i n p a r a l l e l a l i g n m e n t
Brookville, Franklin Co.,
Indiana. A, G, H. Plec- o n b e d d i n g s u r f a c e s ( f i g u r e 13.1C). T w o s p e c i e s r a n g e from t h e K o p e

todina tenuis (Branson t h r o u g h F a i r v i e w f o r m a t i o n s ( B e r g s t r o m 1996a). ' I w o hiserial graptolites

and Mehl). A. Pb ele- o c c u r in 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 qucidrimu-
ment. G. M element. cronatus a n d Arnheimograptus anacanthus (see B e r g s t r o m 1996a). S e v e r a l
H. Sc element. s p e c i e s of Mastigograptus, a d e l i c a t e , b u s h - l i k e d e n d r o i d graptolite, o c c u r
B. Oulodus oregonia i n the K o p e , A r n h e i m , a n d W a y n e s v i l l e f o r m a t i o n s .
(Branson, Mehl, and
The z o o l o g i c a l affinities o f g r a p t o l i t e s w e r e for m a n y years a m o n g t h e
Branson), Pb element.
most c h a l l e n g i n g p r o b l e m s i n p a l e o n t o l o g y . G r a p t o l i t e s w e r e classified
C, D. Amorphogna-
w i t h several d i f f e r e n t 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 y o z o -
thus ordovicicus Bran-
ans, and hemichordates, or were considered to be unrelated to any living
son and Mehl. C. Sc
element. D. Pa ele- g r o u p ( B u l i n a n 1970). R e s e a r c h b y t h e Polish p a l e o n t o l o g i s t R o m a n K o -
ment. E. Drepanoisto- z l o w s k i (1966) n o t e d several s i m i l a r i t i e s b e t w e e n graptolites a n d the l i v i n g
dus suberectus (Branson h e m i c h o r d a t e s c a l l e d p t e r o b r a n c h s that a r g u e stronglv for a c l o s e e v o l u -
and Mehl). F. Phrag- t i o n a r y r e l a t i o n s h i p . P t e r o b r a n c h s are a g r o u p o f s m a l l , m a r i n e , t u b e - d w e l l -
modus undatus Branson i n g i n v e r t e b r a t e s t h a t are c l a s s e d t o g e t h e r w i t h t h e a c o r n w o r m s i n t h e
and Mehl, S element.
P h y l u m 1 l e m i c h o r d a t a o n t h e basis o f e m b r y o l o g i c a l s i m i l a r i t i e s ; there are
I Rhodesognathus el-
o n l y t h r e e liv i n g g e n e r a o f p t e r o b r a n c h s ( B a r n e s 19S7). T h e c r e e p i n g t u b e
egans (Rhodes), Pb ele-
c o n s t r u c t e d by t h e l i v i n g p t e r o b r a n c h Rhabdopleura has the u n i q u e fu-
ment. J. Aphelogna-
sellar h a l f - r i n g a n d c o r t i c a l s t r u c t u r e f o u n d i n graptolites ( B u l i n a n 1970).
thus grandis Branson,
Mehl and Branson, Pb In a d d i t i o n , some e n c r u s t i n g t y p e s of g r a p t o l i t e s h a v e a b l a c k stolon pre-

element. Scanning elec- s e r v e d w i t h i n t h e t u b e s that is verv s i m i l a r to t h e s t r u c t u r e of the stolon in

tron micrographs cour- pterobranchs that interconnects zooids within the colony. The identifica-
tesy of Stig M. Berg- tion of the protein c o l l a g e n in the tubes of both graptolites and ptero-
strom, approximately x b r a n c h s i n d i c a t e s not o n l y their c l o s e r e l a t i o n s h i p , b u t also affinity to chor-
130. P elements are prin- d a t e s , i n w h i c h c o l l a g e n f o r m s t h e c o n n e c t i v e tissue ( A r m s t r o n g e t al.
cipal elements, a and b
19S4). T h e e x t i n c t g r a p t o l i t e s a r e t h u s g e n e r a l l y treated a s a n e x t i n c t class
denoting position in the
w i t h i n the 1 l e m i c h o r d a t a . A l t h o u g h t h e soft part s t r u c t u r e of t h e graptolite
apparatus, M elements
z o o i d s r e m a i n s u n k n o w n , h y p o t h e t i c a l r e c o n s t r u c t i o n s s u g g e s t that the
are medial elements, and
z o o i d s p r o b a b l y h a d p a i r e d t e n t a c l e - b e a r i n g a r m s that w e r e e x t e n d e d for
S elements are symmetry
series elements, c denot- purposes of suspension feeding and deposition of the outer cortical ban-

ing position in the d a g e s o f the t h e c a l w a l l .

apparatus.

Conodonts It is i r o n i c that p e r h a p s t h e m o s t s i g n i f i c a n t C i n c i n n a t i a n fossils, in a g e o -

l o g i c a l s e n s e , a r e a l s o a m o n g t h e least c o n s p i c u o u s t o m o s t o b s e r v e r s .
T h e s e are t h e c o n o d o n t s , t o o t h - l i k e m i c r o f o s s i l s ( < 1 m m ) , s o u n l i k e a n y
o t h e r fossil or l i v i n g o r g a n i s m s that they w e r e r e g a r d e d until r e c e n t l y as
r e p r e s e n t i n g a d i s t i n c t p h y l u m . C o n o d o n t s c a n b e f o u n d t h r o u g h o u t the
C i n c i n n a t i a n by dissolving blocks of limestone in acetic acid, although
they c a n also be found in disaggregated shales. B e c a u s e conodonts have a
c a l c i u m p h o s p h a t e c o m p o s i t i o n , t h e y a r e i n s o l u b l e i n a c e t i c acid. C o n o -
d o n t s are c o n s p i c u o u s in t h e a c i d - i n s o l u b l e r e s i d u e s c a n n e d u n d e r a dis-

196 A Sea without Fish

Graptolites and Conodonts 197
 secting m i c r o s c o p e b e c a u s e of their u n i q u e forms and a beautiful a m b e r
c o l o r (Plate 5B).
C o n o d o n t s h a v e a w i d e r a n g e o f s h a p e s , i n c l u d i n g single c o n e s , m u l t i -
p r o n g e d " t e e t h , " serrated b l a d e - l i k e s h a p e s , a n d s o - c a l l e d p l a t f o r m - t y p e
f o r m s ( F i g u r e 13.2). M o s t c o n o d o n t s h a v e a tooth-like a p p e a r a n c e , l e a d i n g to
t h e a s s u m p t i o n that t h e y w e r e u s e d a s t e e t h . H o w e v e r , c o n o d o n t s also s h o w
r e g e n e r a t i o n . This s u g g e s t s that at least s o m e c o n o d o n t s w e r e e m b e d d e d in
soft tissue b y w h i c h t h e y w e r e s e c r e t e d . I n d i v i d u a l c o n o d o n t s , c a l l e d ele-
m e n t s , w e r e g i v e n s p e c i e s n a m e s b y earlier workers. H o w e v e r , the discovery
of rarely p r e s e r v e d a s s e m b l a g e s of different e l e m e n t s in rock matrix or fused
t o g e t h e r led to t h e r e c o g n i t i o n that e l e m e n t s w e r e a r r a n g e d in bilaterally
s y m m e t r i c a l a s s e m b l a g e s o f pairs o f e l e m e n t s , e a c h o f w h i c h i s c a l l e d a n
a p p a r a t u s . A l t h o u g h few a p p a r a t u s e s are preserved intact, it has b e e n possi-
ble to d e t e r m i n e w h i c h e l e m e n t s f o u n d in a s a m p l e likely f o r m e d an appa-
ratus o n t h e basis o f statistical analysis o f the ratios o f c o m m o n l y associated
e l e m e n t s . M o d e r n t a x o n o m i s t s o f c o n o d o n t s a t t e m p t t o i n c l u d e t h e six o r
s e v e n different e l e m e n t s of a g i v e n a p p a r a t u s u n d e r a single species n a m e .
B e c a u s e the tooth-like elements apparently were the only mineralized
parts o f t h e o r g a n i s m , t h e i d e n t i t y o f t h e c o n o d o n t - b e a r i n g o r g a n i s m a n d
t h e n a t u r e o f its soft p a r t a n a t o m y h a v e b e e n a m o n g t h e great m y s t e r i e s o f
p a l e o n t o l o g y . A m a j o r b r e a k t h r o u g h c a m e in 1983 w h e n a fossil of an e e l -
like, s o f t - b o d i e d o r g a n i s m f r o m t h e L o w e r C a r b o n i f e r o u s o f S c o t l a n d w a s
s h o w n t o h a v e a c o n o d o n t a p p a r a t u s a t o n e e n d a n d fin-like s t r u c t u r e s a t
t h e o p p o s i t e ( B r i g g s e t al. 11)83). T h i s " c o n o d o n t a n i m a l " i s o n l y 4 c m l o n g
a n d s h o w s little of its i n t e r n a l a n a t o m y , save for a m i d l i n e s u g g e s t i n g bilat-
eral s y m m e t r y . A l t h o u g h i t b e a r s s i m i l a r i t i e s t o b o t h c h a e t o g n a t h s a n d
c h o r d a t e s , t h e a u t h o r s c o n c l u d e d that t h e c o n o d o n t a n i m a l b e l o n g s t o a
distinct p h y l u m . M o r e recent phylogenetic studies based on more extensive
data p l a c e t h e c o n o d o n t s w i t h i n t h e c h o r d a t e s , c l o s e t o t h e l i v i n g l a m p r e y s
( D o n o g h u e e t al. 2000). B e c a u s e c o n o d o n t a n i m a l s h a d m i n e r a l i z e d c a l -
c i u m p h o s p h a t e e l e m e n t s , a m o n g o t h e r c h a r a c t e r i s t i c s , t h e y are c o n s i d e r e d
to be vertebrates, even t h o u g h they lacked internal skeletons. T h e c o n o -
dont a n i m a l c o u l d be considered to have b e e n the fish of the C i n c i n n a t i a n
sea. H o w e v e r , g i v e n t h e i r s m a l l s i z e a n d g r e a t d i f f e r e n c e s from a n y t h i n g
like p r e s e n t - d a y b o n y fish, t h e title of this b o o k , A Sea without Fish, re-
m a i n s v a l i d . T h e f o r e g o i n g i n t r o d u c t i o n i s l a r g e l y b a s e d o n C l a r k (1987).
F o r m o r e i n f o r m a t i o n a b o u t t h e m o r p h o l o g y a n d affinities o f c o n o d o n t s
t h e r e a d e r s h o u l d c o n s u l t A l d r i d g e e t al. (1993), A l d r i d g e a n d P u r n e l l
(1996), a n d D o n o g h u e e t al. (2000).
C o n o d o n t s of p a r t i c u l a r t y p e s are f o u n d in m a r i n e rocks over a very
w i d e g e o g r a p h i c r a n g e , e v e n i n t e r c o n t i n e n t a l i n extent. T h i s w i d e distribu-
tion s u g g e s t e d that t h e c o n o d o n t o r g a n i s m was p e l a g i c l o n g before the f i n -
b e a r i n g c o n o d o n t a n i m a l w a s d i s c o v e r e d ( C l a r k 1987). C o n o d o n t s apparently
lived a s s w i m m i n g m e m b e r s o f t h e m a r i n e n e k t o n r a n g i n g from close t o the
sea f l o o r t o v a r i o u s positions i n the w a t e r c o l u m n . T h e tooth-like apparatus
m i g h t h a v e b e e n u s e d to s e i z e or strain food items from the water.
I n a d d i t i o n t o their w i d e g e o g r a p h i c r a n g e , g e n e r a o f c o n o d o n t s are
r e s t r i c t e d i n t h e i r v e r t i c a l (stratigraphic) r a n g e s t h r o u g h o u t their M i d d l e

798 A Sea without Fish

 Figure 13.3. Reconstruc-
tion of the Ordovician
jawless fish, As t rasp is
desiderata Walcott,
from the Harding Sand-
stone of Colorado.
Length about 13 cm.
From Cowen (2005, 86,
figure 7.4). Reprinted
with permission of Black-
well Publishing.

C a m b r i a n t h r o u g h Triassic g e o l o g i c r e c o r d . T h e s e attributes, t o g e t h e r w i t h
a b u n d a n c e i n m a r i n e strata a s m i c r o t o s s i l s , m a k e c o n o d o n t s e x c e p t i o n a l l y
useful for b i o s t r a t i g r a p h i c s u b d i v i s i o n o f t h e P a l e o z o i c s e d i m e n t a r y r o c k
record. For the entire U p p e r O r d o v i c i a n (above t h e b a s e o f t h e L e x i n g t o n
L i m e s t o n e ) in t h e C i n c i n n a t i r e g i o n . S w e e t (1979) r e c o r d e d thirtv-live
conodont species representing twenty genera (Figure 13.2). Most genera
o c c u r also i n E u r o p e , A s i a , A u s t r a l i a , o r A f r i c a , b u t s p e c i e s a r e m o r e g e o -
g r a p h i c a l l y restricted. Most C i n c i n n a t i a n c o n o d o n t s represent the N o r t h
A m e r i c a n M i d c o n t i n c n t P r o v i n c e , b u t s o m e are c o m p o n e n t s o f t h e N o r t h
A t l a n t i c P r o v i n c e that is also r e p r e s e n t e d in E u r o p e . The t y p e - C i n c i n n a -
tian s e c t i o n s p a n s all or parts of six c o n o d o n t - d e f i n e d b i o s t r a t i g r a p h y
z o n e s ( W e b b y , C o o p e r , B e r g s t r o m , a n d Paris 2004). Although many cono-
d o n t taxa h a v e l o n g s t r a t i g r a p h i c r a n g e s w i t h i n t h e C i n c i n n a t i a n , varia-
tions in relative a b u n d a n c e are p r o b a b l y related to d i f f e r i n g d e p t h prefer-
e n c e s a m o n g s p e c i e s (Sweet 1979, 1996).

C o n o d o n t s m a y have b e e n the o n l y representatives of the vertebrates in the Ordovician "fish"

C i n c i n n a t i a n sea, but fossil e v i d e n c e from m a n y localities e l s e w h e r e s h o w s
that s o m e of the earliest tish did i n d e e d exist d u r i n g the O r d o v i c i a n Period.
T h e O r d o v i c i a n H a r d i n g S a n d s t o n e o f C o l o r a d o c o n t a i n s a b u n d a n t , tiny
plates c o m p o s e d of an e n a m e l - c o a t e d d e n t i n e , n a m e d Astnispis desiderata by
W a l c o t t in 1892. In 1997 S a n s o n ) and others illustrated an e x t r a o r d i n a r y fossil
from the H a r d i n g w i t h a c o m p r e s s e d , n e a r l y c o m p l e t e , fish-like b o d y t h a t
proved the l o n g - h e l d a s s u m p t i o n that the tiny plates f o r m e d a h e a d shield
(Figure 13.3). Lyes and a series of o p e n i n g s p r o b a b l y related to respiration are

Qraptolites and Conodonts 199

 p r e s e r v e d . A s i m i l a r fossil fish f r o m O r d o v i c i a n rocks in Bolivia shows that
t h e s e early fish h a d b l u n t , r o u n d e d h e a d s , an e l o n g a t e d fish-like s h a p e w i t h
a tail fin, b u t l a c k e d b o n y jaws a n d separate fins. T h e s e jawless fish are c a l l e d
a g n a t h a n s , b u t o t h e r O r d o v i c i a n fossils r e p r e s e n t t h e earliest jawed fish or
g n a t h o s t o m e s ( S a n s o m et al. 2001). T h u s , a variety of early fish had already
e v o l v e d b y C i n c i n n a t i a n t i m e , b u t t h e y are u n k n o w n from t h e C i n c i n n a t i
r e g i o n . H a d t h e s e early fish b e e n present in t h e C i n c i n n a t i a n sea, it w o u l d
s e e m r e a s o n a b l e t o e x p e c t their m i n e r a l i z e d plates t o b e p r e s e r v e d i n the
l i m e s t o n e s or shales. A l t h o u g h their a b s e n c e m a y indicate that then preferred
-

an e n v i r o n m e n t n o t represented in t h e t y p e - C i n c i n n a t i a n , the potential for

their e v e n t u a l d i s c o v e r ) s h o u l d not b e o v e r l o o k e d .

200 A Sea without Fish

202 A Sea without Fish
TYPE-CINCINNATI ANTRACE FOSSILS:
TRACKS, TRAILS, AND BURROWS

The C i n c i n n a t i a n is r e n o w n e d for its a b u n d a n c e of w e l l - p r e s e r v e d shells

and skeletons o f Orclo\ ician m a r i n e invertebrates, a n d b e c a u s e t h e s e fossils
represent the r e m a i n s o f l o n g - d e a d o r g a n i s m s , a t first g l a n c e o n e w o u l d n o t
e x p e c t t h e m t o yield m u c h i n f o r m a t i o n a b o u t t h e a c t i v i t y a n d b e h a v i o r o f
these a n i m a l s d u r i n g life. O f c o u r s e , w e c a n d e d u c e a g r e a t d e a l a b o u t t h e Figure 14.1. A. Repich-
life habits o f O r d o v i c i a n a n i m a l s d i r e c t l y from the m o r p h o l o g y o f shells nia of the trilobite Isote-
a n d skeletons ( b o d y fossils) by c o m p a r i s o n s to their l i v i n g relatives, b u t a lus, Asaphoidichnus
vast r a n g e o f e v i d e n c e a b o u t a n c i e n t b e h a v i o r also c o m e s f r o m a c o m - trifidum Miller, CMC IP
pletely different s o u r c e , n a m e l y the trace fossils that are b o t h a b u n d a n t a n d 37569, Edenian, Kope
Formation, Cincinnati,
diverse in C i n c i n n a t i a n strata.
Ohio, x 7. B. Repichnia
Trace fossils are e v i d e n c e o f t h e activities o f a n c i e n t o r g a n i s m s pre-
of the trilobite Cryptoli-
served in s e d i m e n t a r y rocks in t h e form of tracks, trails, b u r r o w s , b o r i n g s ,
thus, similar to Cruzi-
a n d o t h e r fossils s u c h as coprolites (fossil feces). M o s t t r a c e fossils w e r e ana, CMC IP 37622, hori-
f o r m e d a s o r g a n i s m s d i s r u p t e d l o o s e s e d i m e n t s b e f o r e l i t h i f i c a t i o n , al- zon and locality
t h o u g h b o r i n g s are the results of d r i l l i n g , r a s p i n g , or e t c h i n g into h a r d unknown, x 7. C. Trilo-
substrata such as shells or a l r c a d v - l i t h i f i e d h a r d g r o u n d s . The study of trace bite trail, intermediate

fossils (also k n o w n as i c b n o f o s s i l s or Lebensspuren) has b e c o m e a s p e c i a l - between Rusophycus

and Cruziana, Maysvil-
ized held o! g e o l o g y k n o w n as i e h n o l o g v . C i n c i n n a t i a n trace fossils with
lian, Corryville Formation,
their exquisite preservation h a v e l o n g f a s c i n a t e d s t u d e n t s o f t h e s e r o c k s .
Clermont Co., Ohio (from
A m o n g them were S. A. Miller, w h o described m a n y C i n c i n n a t i a n trace Osgood [1970, plate 66,
fossils b u t regarded t h e m t o b e the r e m a i n s o f m a r i n e p l a n t s ( s o - c a l l e d figure 3[), x 0.8.
f u c o i d s ) , and J. F. James, w h o w a s o n e of t h e earliest p r o p o n e n t s of t h e D. Paschichnia, ?Paleod-
c o r r e c t o r g a n i c i n t e r p r e t a t i o n o f trace fossils ( O s g o o d 1975a). R i c h a r d C . ictyon, CMC IP 17431,
O s g o o d , jr., w h o c o m p l e t e d his d o c t o r a l dissertation at t h e U n i v e r s i t y of Edenian, Kope Forma-
tion, Cincinnati, Ohio, x
C i n c i n n a t i in 1905 ( O s g o o d 1970), c o n d u c t e d a t h o r o u g h s y s t e m a t i c r e v i e w
3. E. Fodinichnia or
o f C i n c i n n a t i a n trace fossils a n d t h e r e b y c o n t r i b u t e d m u c h t o t h e d e v e l o p -
domichnia, the "turkey
m e n t of m o d e r n i e h n o l o g v .
track," Trichophycus
'Trace fossils greatly e n h a n c e the ability of the p a l e o n t o l o g i s t to r e c o n - venosum Miller, CMC IP
struct the life of the past i( )sgood 1975). In rare eases a trace fossil c a n be 37575, Campbell Co.,
preserved in close association with the b o d y fossil of the a c t u a l t r a c e i n a k e r Kentucky, x 0.4. From
( f i g u r e 14.2). U s u a l l y the identity of the t r a c e i n a k e r is u n k n o w n , often b e - Osgood (1970, plate 60,

c a u s e the traceinaker was soft-bodied and u n p r e s c r v a b l c . Different o r g a n - figure 7). C, E reprinted

by permission of the Pa-
isms and different prcscrvational processes c a n s o m e t i m e s p r o d u c e a similar
leontological Research
type of trace fossil. C o n s e q u e n t l y , trace fossils are not identified by a g e n u s
Institution.
and species n a m e of a b o d y fossil, but instead are g i v e n g e n u s and s p e c i e s
n a m e s of their o w n . The n e e d for this a p p r o a c h is further i n d i c a t e d b e c a u s e
a single o r g a n i s m c a n p r o d u c e several different t y p e s of traces, d e p e n d i n g
on its behavior. 'Thus, a one-for-onc c o r r e s p o n d e n c e b e t w e e n a t r a c e m a k i n g
biological species a n d a given t y p e of trace fossil c a n n o t be establi s hed. T h e

203
 L i n n a e a n b i n o m i a l system w a s d e v e l o p e d for trace fossils b e c a u s e they were
o n c e t h o u g h t t o b e b o d y fossils, and this p r o c e d u r e has persisted ( S i m p s o n
1975). Ideally, t h e i c h n o g e n u s m i g h t be d e f i n e d to represent a p a r t i c u l a r
b e h a v i o r a l pattern w h i l e t h e i c h n o s p e c i e s represents variations on this pat-
t e r n , a l t h o u g h this p r o c e d u r e has n o t b e e n u n i f o r m l y applied (Bromley
1990). T r a c e fossils p r o v i d e i n f o r m a t i o n a b o u t u n p r e s e r v a b l e soft part struc-
tures of a n i m a l s that are k n o w n as skeletal fossils. In the C i n c i n n a t i a n , g o o d
e x a m p l e s of this are t h e varieties of Rusophycus, the resting trace of trilobites,
w i t h i m p r e s s i o n s f o r m e d by t h e d i g g i n g activities of the legs ( F i g u r e 14.2A).
T r a c e fossils also a u g m e n t o u r record of diversity by p r e s e r v i n g activities of
entirely soft-bodied s p e c i e s that are o t h e r w i s e u n k n o w n in the record.
T h e g r e a t e s t s i g n i f i c a n c e o f t r a c e fossils is, h o w e v e r , the i n f o r m a t i o n
they yield about the behavior of long-dead animals. The G e r m a n paleon-
tologist R u d o l f R i c h t e r p i o n e e r e d t h e a n a l y s i s o f a n c i e n t b e h a v i o r from
t r a c e fossils b y s t u d y i n g t r a c e s m a d e b y shallow m a r i n e o r g a n i s m s i n R e -
c e n t s e d i m e n t s o f t h e N o r t h S e a . I n m a n y c a s e s m o d e r n tracks a n d trails
c o u l d be c o m p a r e d to fossilized traces. A remarkable book by W i l h e l m
S c h a f e r (1972) s u m m a r i z e s t h e w o r k o f R i c h t e r a n d m a n y s u b s e q u e n t G e r -
m a n s t u d e n t s o f t h e N o r t h S e a traces i n E n g l i s h . A d o l f S e i l a c h e r (1964)
classified t r a c e fossils into b e h a v i o r a l c a t e g o r i e s that facilitated the inter-
p r e t a t i o n o f a n c i e n t e n v i r o n m e n t s o n t h e basis o f trace fossil a s s e m b l a g e s .

Behavioral R e p i c h n i a are t r a c e s m a d e b y t h e d i r e c t e d l o c o m o t i o n o f b e n t h i c o r g a n -

Categories of i s m s . T h e s e are t h e m o s t c o m m o n t r a c e s i n t h e C i n c i n n a t i a n , w i t h s e v e n -
t e e n i c h n o s p e c i e s r e c o r d e d b y H o l l a n d (2005). T h e a p p e n d a g e s o f trilobites
Trace Fossils
o r o t h e r a r t h r o p o d s d i g g i n g into t h e s u b s t r a t u m d u r i n g c r a w l i n g f o r m e d
several repichnial t r a c e s . Asaphoidichnus trifidum (Miller) (Figure 14.1A)
a n d Allocotkhnus dyeri M i l l e r r e p r e s e n t different forms of the crawling
trace of t h e trilobite lsotelus (Osgood 1970). Trachomatichnus numerosum
M i l l e r is t h e c r a w l i n g t r a c e of t h e trilobite Cryptolithus, and Petaliclmus
multipartitum M i l l e r is t h e c r a w l i n g t r a c e of a m e d i u m - s i z e d , u n i d e n t i f i e d
trilobite ( O s g o o d 1970). O t h e r c o m m o n r e p i c h n i a l traces (Palaeophycus)
are t u b u l a r , u s u a l l y u n b r a n c h e d b u r r o w s that r u n slightly o b l i q u e t o b e d -
ding, preserved either as convex hyporeliefs or trough-like c o n c a v e epire-
liefs. (A h y p o r c l i c f is p r e s e r v e d on t h e b a s e or sole of a s e d i m e n t a r y b e d ;
e p i r c l i e f s are p r e s e r v e d o n t h e u p p e r s u r f a c e o f a bed.) O s g o o d d e s c r i b e d
t h r e e f o r m s o f Palaeophycus a n d i n d i c a t e d that m o d e r n c o u n t e r p a r t s are
produced by infaunal burrowing of polychaete worms or molluscs.
P a s c i c h n i a are m e a n d e r i n g traces m a d e b y the g r a z i n g activities o f d e -
posit feeders ( a n i m a l s that feed on p a r t i c u l a t e o r g a n i c matter either on or
w i t h i n the b o t t o m s e d i m e n t ) . A l t h o u g h n o a c t u a l m e a n d e r i n g traces a r e
f o u n d in t h e C i n c i n n a t i a n , o n e vcrv p e c u l i a r trace q u e s t i o n a b l y referred to
Paleodictyon, is classified by O s g o o d as a p a s c i c h n i a ] trace ( F i g u r e 14.1D).
T h i s trace is a r e m a r k a b l y r e g i d a r n e t w o r k of ridges as a c o n v e x hyporelief.
It r e s e m b l e s the impression of a h o n e y c o m b or c h i c k e n wire. O s g o o d de-
s c r i b e d o n l v t w o k n o w n s p e c i m e n s from t h e C i n c i n n a t i a n , and discussed
the m a n y v a r y i n g interpretations that workers h a v e p r o p o s e d for the origin

204 A Sea without Fish

of Paleodictyon from o t h e r localities. A l t h o u g h it m a y be t h e i m p r e s s i o n of a
patterned object b e i n g rolled a l o n g the s u b s t r a t u m , O s g o o d c o n c l u d e d that
the C i n c i n n a t i a n Paleodictyon is a trace fossil. S e i l a c h e r (1977) interpreted
patterned traces of this t y p e to represent c o m p l e x burrow systems rather than
g r a z i n g patterns. T h e s e b u r r o w n e t w o r k s are u s e d b y s o m e i n f a u n a l o r g a n -
isms for the " f a n n i n g " of m i c r o b e s b e n e a t h the sea floor.
F o d i n i c l m i a are ( c e d i n g t r a c e s o f d e p o s i t f e e d e r s o p e r a t i n g f r o m a
fixed b u r r o w . The m o s t c o m m o n C i n c i n n a t i a n f o d i n i c h n i a are t h e variet-
ies of the b r a n c h i n g b u r r o w Chondrites as d e s c r i b e d by O s g o o d . Chon-
drites, t y p e - A a p p e a r i n g o n t h e v e r t i c a l e d g e s o f b e d s o f f i n e - g r a i n e d car-
b o n a t e s , r e s e m b l e s n a r r o w rootlets (0.8 m m a v e r a g e d i a m e t e r ) , s o m e t i m e s
p e n e t r a t i n g t h e e n t i r e t h i c k n e s s o f a b e d (P'igure 14.3A). O n b e d d i n g
p l a n e s , this form o c c u r s as a c i r c u l a r p a t t e r n of c l o s e l y s p a c e d h o l e s . Chon-
drites, t y p e - B c a n b e s e e n o n b o t h b e d d i n g p l a n e s a n d v e r t i c a l s e c t i o n s .
C o m p a r e d t o Chondrites, t y p e - A , t y p e - B h a s t u b e s o f g r e a t e r d i a m e t e r ( 1 - 4
m m ) , a n d b r a n c h e s that p r o p a g a t e t o a g r e a t e r e x t e n t h o r i z o n t a l l y t h a n
vertically ( F i g u r e 14.3B). Chondrites, t y p e - C o c c u r s a s d e n s e l y i n t e r w o v e n
radiating branches, either parallel or oblique to b e d d i n g (Figure 14.3C).
A n o t h e r c o m m o n a n d i n t e r e s t i n g C i n c i n n a t i a n t r a c e classified b y O s -
g o o d as f o d i n i c l m i a , or possibly d o m i c h n i a , is Trichophycus venosum M i l l e r
( F i g u r e 14.1F). The m o s t c o m m o n e x p r e s s i o n o f this t r a c e , f o u n d t h r o u g h -
out the C i n c i n n a t i a n , is a shallow elongate depression with r o u n d e d ends
o n the u p p e r s u r f a c e o f calcisiltite b e d s . B e d s c o v e r e d w i t h t h e s e s c o o p - l i k e
depressions, oriented in r a n d o m fashion, give the a p p e a r a n c e of overlap-
p i n g tracks o f large birds; h e n c e l o c a l c o l l e c t o r s refer t o t h e s e f e a t u r e s a s
"turkey tracks." O s g o o d ' s c a r e f u l s t u d v o f " t u r k e v t r a c k s " r e v e a l e d that t h e y
represent o n l y a part of a U - s h a p e d b u r r o w s t r u c t u r e w i t h b r a n c h e s in t h e
vertical p l a n e . T h e d e p r e s s i o n s are often f i l l e d w i t h t h i n l a m e l l a e o f f i n e r -
g r a i n e d s e d i m e n t that i n d i c a t e t h e b u r r o w i n g o r g a n i s m d u g t h r o u g h a
m u d layer until it r e a c h e d c o a r s e r silt, w h e r e u p o n it t u r n e d b a c k toward
the s u r f a c e . A f t e r c o m p l e t i n g its d e e p e s t p e n e t r a t i o n t h e b u r r o w i n g o r g a n -
ism r e p o s i t i o n e d itself b y w o r k i n g u p w a r d , f o r m i n g t h e s t a c k e d l a m e l l a e
and d i g g i n g new t u n n e l s u p w a r d , possibly r e a c h i n g t h e s u r f a c e . I n m o s t
c a s e s s u b s e q u e n t e r o s i o n r e m o v e d t h e softer m u d , l e a v i n g t h e m o r e resis-
tant silt w i t h o n l y the basal floor of the b u r r o w as t h e " t u r k e y track." by this
i n t e r p r e t a t i o n , O s g o o d r e j e c t e d t h e f a s c i n a t i n g earlier idea p u t forth b y
R o u s s e a u H . F l o w e r (1955) that " t u r k e y t r a c k s " r e p r e s e n t e d " t o u c h d o w n "
i m p r e s s i o n s left b y n a u t i l o i d c e p h a l o p o d s . The large size of the "turkey
tracks" (width 2.5-3.5 c m
) i n d i c a t e s t h e a c t i v i t y o f a l a r g e o r g a n i s m , b u t its
e x a c t identity r e m a i n s c o n j e c t u r a l . F i n e striations p a r a l l e l t o t h e l e n g t h o f
the d e p r e s s i o n s w e r e f o r m e d b y a p p e n d a g e s o f a n a r t h r o p o d o r e h a e t a e o f
a p o l y c h a e t e w o r m ( e h a e t a e are bristles e x t e n d i n g f r o m t h e b o d y s e g m e n t s
o f a w o r m ) . B e c a u s e t h e b u r r o w s are n o t b i l o b e d , t h e y w e r e m o s t likelv n o t
p r o d u c e d b y a trilobite w i t h p a i r e d a p p e n d a g e s . " T u r k e y t r a c k s " m a y w e l l
represent the onlv e v i d e n c e we h a v e of a rather l a r g e , s o f t - b o d i e d i n f a u n a l
o r g a n i s m that was q u i t e c o m m o n o n t h e C i n c i n n a t i a n sea f l o o r .
D o m i c h n i a are p e r m a n e n t d w e l l i n g t r a c e s f o r m e d b y b e n t h i c a n i m a l s
feeding by predation, scavenging, or suspension feeding. The "U-tube"

Type-Cincinnatian Trace Fossils 205

 Diplocraterion i s t h e m o s t c o m m o n d o m i c h n i a l trace f o s s i l o f the C i n c i n -
n a t i a n . O s g o o d (1970) r e c o g n i z e d t h r e e i e l m o s p c c i e s o f Diplocraterion. A s
s e e n on v e r t i c a l joint s u r f a c e s , / ) . cf. luniformis ( B l a n c k e n h o r n ) has a p l a i n
I l-shape w i t h a r o u n d e d b a s e . N a r r o w , e l o n g a t e d slots on the u p p e r s u r f a c e
of a b e d r e p r e s e n t t h e b a s e of t h e 11. In t h e m o r e c o m m o n / ) . cincinnatien-
sis ( O s g o o d ) t h e U e x p a n d s w i t h d e p t h in different w a y s ( f i g u r e s 14.3A,
14.4 \i. f a i n t dow n w a r d - c u r v i n g striations (Spreiten) c o n n e c t the a r m s of
t h e U . T h e s e s o - c a l l e d p r o t r u s i v e Spreiten m a y represent the d o w n w a r d
propagation of the burrow as the organism grew. Alternatively, the organ-
ism ma}' h a v e b u r r o w e d d e e p e r i n o r d e r t o m a i n t a i n a c o n s t a n t d e p t h b e -
low a constantly e r o d i n g sediment-water interface. T h e reason for the basal
e x p a n s i o n i s u n c l e a r . S o m e e x p a n s i o n s o c c u r a t a n i n t e r f a c e with c o a r s e r
m a t e r i a l , yet o t h e r s e x p a n d w i t h i n t h e silt-sized layer ( f i g u r e 14.4A). T h i s
led O s g o o d (1970) to suggest that t h e o r g a n i s m may h a v e s e n s e d a c h e m i c a l
c h a n g e i n t h e s e d i m e n t b e f o r e e n c o u n t e r i n g a c h a n g e i n g r a i n size. W e
s p e c u l a t e that t h e e x p a n s i o n o f t h e U m i g h t h a v e d e v e l o p e d after t h e
w o r m - l i k e o r g a n i s m b u r r o w e d t o its preferred d e p t h . Lateral e x p a n s i o n o f
t h e U e n a b l e d t h e w o r m t o g r o w i n l e n g t h w h i l e m a i n t a i n i n g its d e p t h .
T h e third Diplocraterion i c h n o s p e c i e s , D. biclavata ( M i l l e r ) , has a pair
o f short e x t e n s i o n s d e v e l o p e d a t t h e b a s e o f t h e U a s b l i n d p o u c h e s . T h i s
f o r m also o c c u r s a s c o n c a v e e p i r c l i e f s i n w h a t a r e c o m m o n l y c a l l e d " d u m b -
b e l l s " ( f i g u r e s 1 4 . 4 B , C ) . O s g o o d s h o w e d that d u m b b e l l s r e p r e s e n t the
basal p e n e t r a t i o n o f t h e D - t u b e w i t h p a i r e d e x t e n s i o n s into a c o a r s e r s u b -
s t r a t u m t h a t u s u a l l y r e m a i n s after e r o s i o n s t r i p p e d off t h e u p p e r parts o f
t h e U . A l l t h r e e i c h n o s p e c i e s o f Diplocraterion c a n b e f o u n d i n t h e s a m e
b e d , s u g g e s t i n g t h a t a s i n g l e t y p e o f o r g a n i s m c o u l d h a v e p r o d u c e d all
three expressions. T h e t r a c c m a k i n g organism was probablv some type of
w o r m that u s e d t h e U - t u b e as its d w e l l i n g s t r u c t u r e , w h i l e s u s p e n s i o n feed-
ing near the sediment-water interface.
M a n y b o r i n g s found i n C i n c i n n a t i a n stromatoporoids, corals, b r y o z o a n s ,
b r a c h i o p o d s , m o l l u s c s , a n d h a r d g r o u n d s represent d o m i c h n i a . Pits on the
u n d e r s i d e of a R i e h m o n d i a n stromatoporoid c o n t a i n the t r a c c m a k i n g bivalve
Corallidomus scobina Pojeta and P a l m e r as t h e oldest-known c a s e of hard
substrate b o r i n g by a p e l e c y p o d (see F i g u r e 9.8A; Pojeta and P a l m e r 1976).
T h e s a m e e l o n g a t e pits, now referred to the i c h n o s p e c i e s Petroxestes pera, are
also f o u n d in c a l c a r e o u s n o d u l e s a n d the bases of b r y o z o a n s ( W i l s o n and
P a l m e r 1988). C i n c i n n a t i a n r u g o s e corals of the g e n u s Grewiugkia c o m m o n l y
show b o r i n g s c a l l e d Trypanites (see f i g u r e 6.5K; f l i a s 1986). A l t h o u g h it is
usually impossible to d e t e r m i n e w h e t h e r the b o r i n g s o c c u r r e d d u r i n g the life
of the c o r a l , Klias f o u n d s o m e b o r i n g s e n c a s e d w i t h i n swellings of the coral
septa. F l i a s inferred that a b o r i n g w o r m penetrated the septa of a living coral,
resulting in the secretion of the septal s w e l l i n g . In other eases borings did not
c o m e into c o n t a c t with the living coral polyps, hut were probablv located on
corals in life position so as to e x p o s e the w o r m s to a m b i e n t c u r r e n t flow.
A distinctive t y p e of b o r i n g c a l l e d Sanctum laurentiensis f rickson and
B o u c h a r d o c c u r s a s isolated, r o u n d t u n n e l s o n t h e interior o f b r a n c h i n g
C i n c i n n a t i a n b r y o z o a n s ( K r i c k s o n and B o u c h a r d 2003). T h e interior o f the
excavation is e l o n g a t e or sack-shaped and 1 m i m e d . T h e trace-maker was likely

206 A Sea without Fish

 Figure 1 4 . 2 . A. Cubich-
nia and sderite impres-
sions of trilobite Isotelus,
Rusophycus carleyi (J. F.
James), CMC IP 46411,
Maysvillian, Corryville
Formation, Clermont Co.,
Ohio, x 0.6. Note impres-
sions of cephalic margin,
genal spines, pleurae,
pygidial margin, as well
as coxae (medial paired
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 IP
37574, Maysvillian, Cor-
ryville Formation, Cler-
mont Co., Ohio, x
1.4. Right: Reverse side
of specimen shown in B,
cubichnia Rusophycus
pudicum Hall, convex
hyporelief, identifying
trilobite as tracemaker.
This exceptional speci-
men is one of the very
few known with both
trace and tracemaker
preserved together.

Type-Cinannatian Trace Fossils 207

Figure 14.3. A. Vertical joint surface showing two commonly associated trace fossils: at left, domichnia,
Diplocraterion cincinnatiensis Osgood; at right, fodinichnia, Chondrites, type-A, CMC IP 37607, Edenian,
Kope Formation, Cincinnati, Ohio, x 1.3. B. Fodinichnia, Chondrites, type-B, CMC IP 37623, Maysvillian,
Corryville Formation, Clermont Co., Ohio, x 0.9. C. Fodinichnia, Chondrites, type-C, CMC IP 37678, Rich-
mondian, Whitewater Formation, Wayne Co., Indiana, x 1.2. D. Cubichnia of sea star, Asteriacites stelli-
forme (Miller and Dyer), CMC IP uncatalogued, Maysvillian, PCorryville Formation, Cincinnati, Ohio, x 7.

208 A Sea without Fish

Figure 14.4. A. Vertical joint surface showing domichnia, Diplocraterion cincinnatiensis Osgood, CMC IP,
uncatalogued, Edenian, Kope Formation, Campbell Co., Kentucky, x 0.8. B. Reconstruction of Diplocrate-
rion biclavata (Miller), showing U-tube and spreiten constructed in upper shale (dashed pattern) with base
of U and paired lateral extensions in underlying calcisiltite (stippled pattern). When shale is eroded away, a
"dumbbell" impression remains on the upper surface of the calcisiltite, as shown in lower diagram. Modified
from Osgood (1970). C. Domichnia Diplocraterion biclavata (Miller), "dumbbell" on upper surface, CMC
IP 37657, Maysvillian, Corryville Formation, Clermont Co., Ohio, (from Osgood [1970, plate 61, figure 3]), x
0.4. D. bioimmuration structure in bryozoan, Catellocaula vallata Palmer and Wilson, University of Cincin-
nati collection, Edenian, Point Pleasant Formation, Bracken Co., Kentucky, x.2.3. E. Cubichnia of pele-
cypods, Lockeia siliquaria U. P. James, CMC IP 37597, Edenian, Kenton Co., Kentucky, x 0.5. F. Palaeoscia
floweri Caster, holotype, CMC IP 24079, Maysvillian, Corryville Formation, Clermont Co., Ohio. Although
originally interpreted as the impression of a porpitid jellyfish, this impression was considered by Osgood
(1970) to be either a feeding trace or of inorganic origin as a result of rotatory sweeping of an agglutinated
tube, x 0.6. B, C reprinted by permission of the Paleontological Research Institution.

Type-Cincinnatian Trace Fossils 209

 a c r u s t a c e a n that took a d v a n t a g e of the b r y o z o a n host for protection as well
as for an elevated f e e d i n g position. E x c a v a t i o n of the interior of the b r v o z o a n
b r a n c h e s probably r e d u c e d the structural integrity of the colonv and rendered
it m o r e susceptible to b r e a k a g e d u r i n g storms. Trypanites borings on b r y o z o -
ans u s u a l l y o c c u r in clusters and w e r e probably f o r m e d w h e n d e a d , broken
c o l o n i e s w e r e e x p o s e d on the sea floor ( E r i c k s o n and B o u c h a r d 2003).
A n o t h e r type of trace found in C i n c i n n a t i a n brvozoan colonies is not
a c t u a l l y a b o r i n g b u t rather r e c o r d s t h e p r e s e n c e of a s o f t - b o d i e d o r g a n i s m
that lived a s a n e n d o s y m b i o n t w i t h i n t h e b r v o z o a n s k e l e t o n , t e r m e d b i o -
c l a u s t r a t i o n b y P a l m e r a n d W i l s o n (lcjHS) ( f i g u r e 1 4 . 4 0 ; s e e F i g u r e 11.6E).
A f t e r s e t t l e m e n t by a larva of t h e e n d o s y m b i o n t o n t o the l i v i n g colonv,
b r v o z o a n z o o e c i a grew 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 s h a p e ,
r e s u l t i n g in d i s t i n c t i v e pits a r r a n g e d in r o w s , n a m e d Catellocaula vallata
by P a l m e r a n d 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 t h e s e pits is
l i n e d b y z o o e c i a l w a l l s . T h e m o r p h o l o g y o f the pits and their a r r a n g e m e n t
in rows s u g g e s t e d a c o l o n i a l , s t o l o n i f e r o u s o r g a n i s m , m o s t likely a t u n i c a t e .
T a p a n i l a (2005) has p r o p o s e d that a new b e h a v i o r a l c a t e g o r y , I m p e d i c h n i a ,
b e u s e d for s u c h c a v i t i e s that l o c a l l y i n h i b i t the n o r m a l skeletal g r o w t h o f
t h e host. Catellocaula vallata r e p r e s e n t s o n e of the oldest k n o w n e x a m p l e s
o f this e n d o s y m b i o t i c b e h a v i o r .
C u b i c h n i a are t e m p o r a r y traces m a d e b y m o b i l e a n i m a l s . A l t h o u g h they
are often regarded as " r e s t i n g " traces, there is also the possibly that s o m e c u -
b i c h n i a represent f e e d i n g or predation b u r r o w s . Trilobite "resting" traces,
represented by three i c h n o s p e c i e s of Rusophycus, are a m o n g the m o s t c o m -
m o n c u b i e h n i a l traces in the C i n c i n n a t i a n . O s g o o d (1970) r e c o g n i z e d R.
pudicum I lall as the trace of Flexicalymene on the basis of its size and a few
e x c e p t i o n a l o c c u r r e n c e s of the t r a c c n i a k e r preserved with the trace directly
b e n e a t h it ( f i g u r e 14.2B). O c c a s i o n a l l y clusters of R. pudicum are found that
suggest the activity of several trilobites, a l t h o u g h a single individual c o u l d also
p r o d u c e m u l t i p l e b u r r o w s in close proximity. R. carleyi O. f. James) is a large
b u r r o w attributable to the largest C i n c i n n a t i a n trilobite Isotelus. In a few ex-
c e p t i o n a l s p e c i m e n s , casts of the c e p h a l i c and pvgidial m a r g i n , genal spines,
and p l e u r a e are present a l o n g with impressions formed by the w a l k i n g legs
( f i g u r e 14.2A; O s g o o d 1970; Brandt et al. 1995). hi e v e n rarer s p e c i m e n s , in-
tersection by the trilobite trace of a w o r m burrow suggests that the burrow was
f o r m e d for the p u r p o s e s of predation ( F i g u r e 14.2A; Brandt et al. 1995; Fortey
a n d O w e n s 1999). S m a l l , ovoid Rusophycus (R. cryptolithi) are consistent with
formation by the trilobite Cryptolithus ( O s g o o d 1970). O t h e r c u b i e h n i a l traces
for w h i c h the t r a c c n i a k e r is identifiable are Lockeia siliquaria U. P. James,
f o r m e d by shallow-burrow i n g p e l e c v p o d s ( f i g u r e 14.4F) and Asteriacites stel-
liforme ( M i l l e r a n d O v e r ) f o r m e d by sea stars ( F i g u r e 14.3D; O s g o o d 1970).

Ichnofacies and S e i l a c h e r (1964) p r o p o s e d that t h e relative a b u n d a n c e of trace fossils b e l o n g -

Paleoenvironmental i n g to the b e h a v i o r a l c a t e g o r i e s varied a c c o r d i n g to the s u b m a r i n e e n v i r o n -

Interpretation m e n t , c h i e f l y in relation to d e p t h . Trace fossil a s s e m b l a g e s ( i c h n o f a c i e s )

d o m i n a t e d b y d o m i c h n i a ] and c u b i e h n i a l traces c h a r a c t e r i z e shallow, near-
shore m a r i n e e n v i r o n m e n t s b e c a u s e c o n d i t i o n s o f h i g h t u r b u l e n c e c a u s e

210 A Sea without Fish

vagile o r g a n i s m s and s u s p e n s i o n feeders to seek shelter in b u r r o w s . As d e p t h
increases w i t h i n the shallow n e a i shore / o n e , c o n d i t i o n s of l o w e r water m o v e -
m e n t p e r m i t food p a r t i c l e s to settle, a n d thus deposit f e e d i n g activity in-
creases, resulting in f o d i n i e h n i a l traces. In d e e p sea e n v i r o n m e n t s t h e r e is
little n e e d for p e r m a n e n t shelter, and so d w e l l i n g a n d resting traces are n o t
f o r m e d ; instead, o r g a n i s m s tend to g r a z e the s e d i m e n t s u r f a c e for f o o d , creat-
ing the m e a n d e r i n g p a s c i c h n i a l traces. K e p i c l m i a l traces are f o u n d in all
s u b m a r i n e e n v i r o n m e n t s , b e c a u s e o r g a n i s m s are a l w a y s g o i n g s o m e w h e r e
a n d l e a v i n g trails, n o matter w h a t the setting! S e i l a c h e r ' s o r i g i n a l c o n c e p t o f
trace fossil facics distribution has b e e n w i d e l y tested, s u b s t a n t i a t e d , and ex-
panded to include assemblages characterizing non-marine environments
and p a r t i c u l a r substrata s u c h a s h a r d g r o u n d s a n d w o o d ( B r o m l e y 1990). C a n
t h e i c h n o f a c i e s c o n c e p t b e a p p l i e d t o C i n c i n n a t i a n trace f o s s i l s , a n d what
c a n this tell us a b o u t the Late O r d o v i c i a n m a r i n e e n v i r o n m e n t ?
O s g o o d (1970) listed thirty i c h n o g e n e r a a n d forty-four i c h n o s p e c i e s
from t h e C i n c i n n a t i a n . R e c e n t a d d i t i o n s a n d revisions b r i n g t h e total t o
thirty-four i c h n o g e n e r a and fortv-seven i c h n o s p e c i e s ( H o l l a n d 2005; T a p a -
nila 200^). The f o l l o w i n g list s h o w s the distribution of t h e s e i c h n o s p e c i e s .

Cubichnia 5
Domichnia 6
Repichnia 17
Kodinichnia 9
Pascichnia 1
[mpedichnia 2
Borings 2
incertae sedis 5

T h e h i g h diversity i n the c a t e g o r i e s d o m i c h n i a , c u b i c h n i a , a n d fo-

d i n i c h n i a , c o m p a r e d t o the s i n g l e (and rare) o c c u r r e n c e o f p a s c i c h n i a , led
S e i l a c h e r (1964) a n d O s g o o d (1970) to c o n c l u d e that the C i n c i n n a t i a n as a
w h o l e c o i n c i d e s w ith the Cruziana i c h n o f a c i e s , r e f l e c t i n g a s h a l l o w , o p e n
m a r i n e e n v i r o n m e n t . O s g o o d s u g g e s t e d that the C i n c i n n a t i a n m a r i n e e n -
v i r o n m e n t w a s m i d e e p e r M i a n about 35 i n .
V e r y little work has b e e n d o n e t o refine p a l e o e n v i r o n m e n t a l a n a l y s i s
o f the C i n c i n n a t i a n u s i n g trace f o s s i l s . O s g o o d n o t e d that a " s u b a s s o c i a -
t i o n " of three trace fossils, Diplocraterion, Chondrites, a n d Trichophycus is
found in s i n g l e b e d s n e a r the top of t h e K o p c F o r m a t i o n a n d r e c u r s in the
C o r r y v i l l e a n d a g a i n in the Liberty F o r m a t i o n s , s u g g e s t i n g a r e c u r r e n c e
of s i m i l a r c o n d i t i o n s . I l i g h r e s o l u t i o n s t r a t i g r a p h i c w o r k has d e m o n s t r a t e d
that t h e s e Diplocraterion b e d s in t h e u p p e r K o p c are w i d e l v t r a c e a b l e o v e r
the G r e a t e r C i n c i n n a t i r e g i o n ( J e n n e t t e a n d P r y o r 1993; Brett e t a l . 2001b),
but there has b e e n no d e t a i l e d study of t h e trace fossil a s s o c i a t i o n .

"Traces" of Inorganic or Indeterminate Origin

S e v e r a l s e d i m e n t a r y s t r u c t u r e s i n C i n c i n n a t i a n strata w e r e n a m e d a s " f u -
c o i d s " but are likely t o h a v e a n i n o r g a n i c o r i g i n o r r e m a i n p r o b l e m a t i c .

Type-Cincinnatian Trace Fossils 211

 T h e s e are u s u a l l y p r e s e r v e d a s h y p o r e l i e f s . O n e s u c h s t r u c t u r e , "Blastophy-
cus," p r o b a b l y r e p r e s e n t s c a s t i n g s of i m p r e s s i o n s left by e n r o l l e d trilobites
a n d c u r r e n t s c o u r a r o u n d t h e m ( O s g o o d 1970). A n o t h e r , "Dystactophycus,"
i s a f a n - s h a p e d p a t t e r n o f f i n e c o n c e n t r i c ridges a n d g r o o v e s , a n d w a s o n c e
t h o u g h t t o b e a n a l g a l f r o n d . O s g o o d c o n c l u d e d that i t f o r m e d b y rotation
of a c r i n o i d s t e m as it w a s b u r i e d . At several C i n c i n n a t i a n l o c a l i t i e s , per-
fectly circular impressions of c o n c e n t r i c rings o c c u r on the upper surface
o f a b e d ( F i g u r e 1 4 . 4 F ) . C a s t e r i n t e r p r e t e d t h e s e t o b e b o d y fossil m o l d s o f
a p o r p i t i d jellyfish a n d d e s c r i b e d t h e m as Palaeoscia floweri (see C a s t e r
1942). H o w e v e r , O s g o o d e x a m i n e d a d d i t i o n a l s p e c i m e n s a n d c o n s i d e r e d
t h a t t h e s e c o n c e n t r i c r i n g s c o u l d h a v e f o r m e d u n d e r t h e i n f l u e n c e o f cur-
rents b y r o t a t i o n a l s w e e p i n g o f s o m e k i n d o f o r g a n i c d w e l l i n g t u b e e m b e d -
d e d i n t h e s e d i m e n t . A l t e r n a t i v e l y , s o m e l i v i n g p o l y c h a e t e s create a f e e d i n g
t r a c e t h a t r e s e m b l e s Palaeoscia, a n d t h u s O s g o o d relegated these most
p 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 " to incertae sedis. S t a n l e y (1986) s u p p o r t e d
O s g o o d ' s a s s e s s m e n t t h a t Palaeoscia is a t r a c e fossil, b u t c o n t r o v e r s y c o n -
t i n u e s o v e r i n t e r p r e t a t i o n of Palaeoscia a n d s i m i l a r c o n c e n t r i c ring-like
s t r u c t u r e s i n t h e g e o l o g i c a l r e c o r d ( E w i n g a n d D a v i s 1967, 274-275). B e l l
et al. (2001) listed C i n c i n n a t i a n Palaeoscia as a jellyfish, a n d asserted that
O s g o o d w a s i n c o r r e c t in r e g a r d i n g it as a t r a c e fossil, b u t g a v e no basis for
this e v a l u a t i o n .

212 A Sea without Fish

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 fourth
stage, 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
in modern usage are
shown, but dozens of
different named divisions
have been proposed over
the years and are not
shown. The names cur-
rently used by the Ohio
and Kentucky geological
surveys differ, and
dashed vertical lines indi-
cate these "stateline
stratigraphic divisions."
The Indiana Geological
Survey recognizes the
Kope, Whitewater, and
Saluda Formations, and
assigns all strata between
the Kope and Whitewa-
ter to the Bull Fork For-
mation (not shown).
Type-Cincinnatian strata
have also been divided
into six units that reflect
cycles of sedimentation
reflect falls in sea level that drained the seas from the Cincinnati area, result-
produced by the rise and
ing in no deposition of sediments. The actual duration of these unconformi-
fall of sea level. These
ties is poorly known.
depositional sequences,
numbered C1-C6, are
bounded by unconformi-
ties shown in gray that

214 A Sea without Fish

 15
PALEOGEOGRAPHY AND
PALEOENVIRONMENT
Steven M. Holland

E a r t h scientists r e c o n s t r u c l c o n d i t i o n s d u r i n g t h e a n c i e n t past f r o m a w i d e
variety o f c h i c s from m i n e r a l s , r o c k s , a n d f o s s i l s . A l t h o u g h n o p l a c e o n
E a r t h today i s e x a c t l y like t h e C i n c i n n a t i area d u r i n g t h e L a t e O r d o v i c i a n ,
c o m p a r i s o n s w i t h m o d e r n e n v i r o n m e n t s offer v a l u a b l e i n s i g h t s into t h e
interpretation o f t h e s e c l u e s . O f m o d e r n e n v i r o n m e n t s , t h e Persian G u l f
is perhaps the most similar to the Late O r d o v i c i a n of the eastern United
States in t e r m s of its c l i m a t e , t h e s i z e of the s e d i m e n t a r y b a s i n , the gcntlv
d i p p i n g sea floor, t h e m i x o f c a r b o n a t e s e d i m e n t a n d clay, a n d t h e o c c u r -
r e n c e o f storms that rework a n d d e p o s i t s e d i m e n t .

" W h e n Cincinnati Was in the Southern H e m i s p h e r e " Geography

( T i t l e of a p r e s e n t a t i o n by K e n n e t h E, C a s t e r for t h e C i n c i n n a t i
Historical S o c i e t y , 1974)

G l o b a l g e o g r a p h y d u r i n g the O r d o v ician has b e e n r e c o n s t r u c t e d primarily

t h r o u g h s t u d i e s o f the m a g n e t i c p r o p e r t i e s o f O r d o v i c i a n r o c k s . \ \ h e n
s e d i m e n t s a r e d e p o s i t e d , i r o n - b e a r i n g m i n e r a l s t e n d t o a l i g n w i t h the
Earth's m a g n e t i c f i e l d . A s the s e d i m e n t s b e c o m e c e m e n t e d t o g e t h e r a n d
u n d e r g o lithifieation t o b e c o m e r o c k s , t h e s e m i n i a t u r e m a g n e t s are l o c k e d
in place. By careful m e a s u r e m e n t of the orientations of these iron-bearing
m i n e r a l s , g e o p h y s i c i s t s r e c o n s t r u c t the l a t i t u d e a t w h i c h t h e s e d i m e n t s
were originally deposited.
Four large continents d o m i n a t e d the globe d u r i n g the O r d o v i c i a n
(Plate 1). S t r e t c h i n g from the S o u t h Pole into t h e n o r t h e r n h e m i s p h e r e ,
Gondvv ana w a s t h e largest o f t h e s e c o n t i n e n t s a n d c o n s i s t e d o f m o d e r n dav
S o u t h A m e r i c a , A f r i c a , A r a b i a , A n t a r c t i c a , A u s t r a l i a , India, s o u t h e a s t Asia,
a n d parts o f C h i n a . L a u r e n t i a i n c l u d e d m o s t o f p r e s e n t - d a y N o r t h A m e r -
ica, e x c e p t for New E n g l a n d a n d M a r i t i m e ( ' a n a d a , parts o f t h e s o u t h e a s t -
ern U n i t e d States, a n d t h e west c o a s t o f t h e U n i t e d States a n d C a n a d a , all
of which were added to Laurentia d u r i n g subsequent tectonic collisions.
L a u r e n t i a straddled t h e e q u a t o r d u r i n g the O r d o v i c i a n a n d w a s rotated 45"
c l o c k w i s e from its present-day o r i e n t a t i o n . As a result, C i n c i n n a t i w a s situ-
ated a t 20-25 s o u t h o f t h e e q u a t o r for t h e e n t i r e L a t e O r d o v i c i a n . T o t h e
east o f L a u r e n t i a a l o n g the e q u a t o r sat t h e c o n t i n e n t o f S i b e r i a - K a z a k h s t a n ,
w h i c h consisted o f present-day p o r t i o n s o f c e n t r a l A s i a . T h e f o u r t h c o n t i -
n e n t , B a l t i c a , lav to the s o u t h at r o u g h l y 60 a n d w a s c o m p o s e d of n o r t h e r n
Europe and Scandinavia.

215
 T h r e e major o c e a n s separated these continents. T h e I a p e t u s O c e a n
separated L a u r e n t i a and S i b e r i a - K a z a k h s t a n from B a l t i c a t o the s o u t h . T h e
P a l e o t e t h y s O c e a n lay b e t w e e n G o n d w a n a a n d t h e c o n t i n e n t s o f B a l t i c a
a n d S i b e r i a - K a z a k h s t a n t o t h e w e s t . T h e m a s s i v e P a n t h a l a s s i c O c e a n cov-
e r e d a l m o s t all o f t h e N o r t h e r n H e m i s p h e r e a n d w o u l d have d w a r f e d to-
day's Pacific O c e a n .
G l o b a l sea level w a s h i g h d u r i n g the O r d o v i c i a n , and a l t h o u g h its
p o s i t i o n is d i f f i c u l t to c o n s t r a i n , t h e c u r r e n t c o n s e n s u s is that it was 100 to
200 m e t e r s h i g h e r t h a n p r e s e n t - d a y sea level (see F i g u r e 1.3). S e v e r a l factors
c o n t r i b u t e d to s u c h a h i g h p o s i t i o n of sea l e v e l . Rates of sea floor s p r e a d i n g
w e r e h i g h f o l l o w i n g t h e b r e a k u p o f a n older, L a t e P r o t e r o z o i c s u p e r c o n t i -
n e n t c a l l e d R o d i n i a , c a u s i n g t h e a v e r a g e e l e v a t i o n o f the sea floor t o b e
higher than normal. This r a i s i n g o f t h e " b o t t o m o f the b u c k e t " forced
o c e a n waters to spill o n t o t h e c o n t i n e n t s . In a d d i t i o n , the lack of p o l a r ice
c a p s in m o s t of t h e O r d o v i c i a n a l s o w o u l d h a v e raised sea level relative to
t o d a y b e c a u s e w a t e r i n m o d e r n g l a c i a l ice c a p s s u c h a s A n t a r c t i c a a n d
G r e e n l a n d is p r o d u c e d from snow generated by evaporation from the
o c e a n . B e c a u s e o f this h i g h sea l e v e l , l o w - l y i n g areas o n the c o n t i n e n t s
w e r e f l o o d e d w i t h o c e a n w a t e r s , m u c h like the f l o o d i n g o f the present-day
c o n t i n e n t a l s h e l v e s , b u t t o a m u c h greater e x t e n t . D u r i n g m u c h o f the
O r d o v i c i a n , most o f L a m e n t i a w a s s u b m e r g e d , w i t h the e x c e p t i o n o f parts
o f t h e C a n a d i a n S h i e l d , a low m o u n t a i n r a n g e r u n n i n g from M i n n e s o t a
toward C o l o r a d o , t h e O z a r k r e g i o n o f M i s s o u r i , a n d the u p l i f t i n g l a c o n i c
M o u n t a i n s a l o n g t h e s o u t h e a s t e r n e d g e o f L a u r e n t i a (Plate 1).
The eastern U n i t e d States was d i v i d e d into several g e o g r a p h i c regions
d u r i n g t h e L a t e O r d o v i c i a n (Plate 12). E x t e n d i n g from central K e n t u c k y
n o r t h w a r d into O h i o a n d I n d i a n a w a s a shallow m a r i n e c a r b o n a t e area
k n o w n a s the L e x i n g t o n Platform. W a t e r d e p t h s o n the L e x i n g t o n Platform
d e e p e n e d t o the n o r t h . T h e L e x i n g t o n Platform was b o u n d e d o n the west
by a d e e p e r water t r o u g h k n o w n as the S e b r e e T r o u g h . Far to the east rose
t h e T a c o n i c M o u n t a i n s , p r o d u c e d by the collision of Laurentia with a small
plate or island are, s u c h as the m o d e r n - d a y islands of Japan and the A l e u -
tians. This c o l l i s i o n w a r p e d t h e s o u t h e a s t e r n m a r g i n of L a u r e n t i a to form a
d e e p water trough c a l l e d the A p p a l a c h i a n Basin that separated the l a c o n i c
M o u n t a i n s from the L e x i n g t o n Platform. Into this t r o u g h , s e d i m e n t s shed
from the e r o d i n g T a c o n i c M o u n t a i n s built a Series of northwest ward-advanc-
i n g deltas c a l l e d the Q u e e n s t o n D e l t a . " R e d - b e d " tidal f l a t s e d i m e n t s typical
of the Q u e e n s t o n c a n be seen in the latest O r d o v i c i a n strata on the eastern
side o f t h e C i n c i n n a t i A r c h i n A d a m s C o u n t y , O h i o .

Climate The c l i m a t e o f t h e L a t e O r d o v i c i a n was w a r m , with m u c h m o r e e v e n t e m -

peratures from the p o l e to the e q u a t o r than seen today. H i g h c o n c e n t r a t i o n s
of c a r b o n d i o x i d e in t h e a t m o s p h e r e (referred to as the partial pressure of
carbon dioxide, or pCO2,) caused these w a r m conditions. C o m p u t e r models
and l i m i t e d g e o c h e m i c a l data from O r d o v i c i a n soil deposits suggest that lev-
els of a t m o s p h e r i c pCO2, w e r e nearly sixteen times greater than today, c o m -
pared to the 20 p e r c e n t i n c r e a s e in pCO2,witnessed in the past half-century.

216 A Sea without Fish

 W a r m t e m p e r a t u r e s a t the p o l e s i n h i b i t e d t h e f o r m a t i o n o f i c e c a p s ,
s u c h a s today's c o n t i n e n t a l g l a c i e r s o n A n t a r c t i c a a n d G r e e n l a n d a n d the
sea ice o v e r the A r c t i c O c e a n . I n t h e last m i l l i o n years o f the O r d o v i c i a n ,
a t m o s p h e r i c p C O , levels d r o p p e d precipitously, t r i g g e r i n g t h e rapid g r o w t h
2

of p o l a r g l a c i e r s a n d a g e o l o g i c a l l y brief 160 m e t e r g l o b a l sea level fall. T h i s

fall in sea level d r a i n e d t h e seas f r o m t h e C i n c i n n a t i a r e a , p r o d u c i n g an
e r o s i o n a l division b e t w e e n t h e O r d o v i c i a n a n d S i l u r i a n strata c a l l e d a n
unconformity.
In a d d i t i o n to this e n d - O r d o v i c i a n fall in sea l e v e l , e v i d e n c e for six
c y c l e s o f g l o b a l sea level c h a n g e i s p r e s e r v e d i n the O r d o v i c i a n n e a r C i n -
c i n n a t i ( f i g u r e 15.1). T h e e v i d e n c e for t h e s e c y c l e s c o m e s f r o m p a c k a g e s
o f rock k n o w n a s d e p o s i t i o n a l s e q u e n c e s , w h i c h are b o u n d e d b y u n c o n f o r -
m i t i e s , o r s u r f a c e s that record t h e e r o s i o n a n d w e a t h e r i n g o f s e d i m e n t s .
E a c h d e p o s i t i o n a l s e q u e n c e b e g i n s w i t h a relatively t h i n interval o f r o c k
that records l o c a l d e e p e n i n g o f t h e o c e a n s a n d e n d s w i t h a m u c h t h i c k e r
interval o f rock that r e c o r d s p r o g r e s s i v e s h a l l o w i n g o f t h e o c e a n s . T h e s e
s a m e s e q u e n c e s c a n b e r e c o g n i z e d across the U n i t e d States a n d i n E s t o n i a .
The fact that t h e s e s e q u e n c e s are n o t just l o c a l f e a t u r e s is s t r o n g e v i d e n c e
that they reflect g l o b a l sea level c h a n g e s rather t h a n l o c a l t e c t o n i c c h a n g e s .
I n t h e C i n c i n n a t i a r e a , t h e s e si\ d e p o s i t i o n a l s e q u e n c e s a l s o c o n t a i n e v i -
d e n c e of shorter-term v a r i a t i o n s in sea l e v e l , but it is c u r r e n t l y u n c l e a r
w h e t h e r t h e s e represent g l o b a l o r r e g i o n a l c h a n g e s i n sea l e v e l .
f r o m s t u d i e s o f t o d a y s g e o g r a p h y a n d c l i m a t e , a s well a s d i r e c t e v i -
d e n c e from O r d o v i c i a n strata, g e o l o g i s t s h a v e r e c o n s t r u c t e d t h e O r d o v i -
c i a n c l i m a t e o f the C i n c i n n a t i area. Today, r e g i o n s n e a r 30 n o r t h a n d
south of the e q u a t o r are c h a r a c t e r i z e d by d e s e r t s , w h i c h f o r m as air r i s i n g
n e a r the e q u a t o r d e s c e n d s a n d f o r m s a series of h i g h p r e s s u r e c e l l s that
inhibit rainfall. I n O r d o v i c i a n rocks o f the C i n c i n n a t i a r e a , d i r e c t e v i d e n c e
of such a semi-arid climate can be seen in the finely l a m i n a t e d d o l o m i t i c
tidal flat deposits of c e n t r a l K e n t u c k y . S i m i l a r d e p o s i t s o c c u r in s e m i - a r i d
regions today, w h e r e h i g h tides d e p o s i t t h i n l a m i n a e o f s e d i m e n t , b u t h i g h
salinity a n d t e m p e r a t u r e e x c l u d e a n i m a l s that m i g h t burrow a n d d i s r u p t
the s e d i m e n t . D o l o m i t e itself c a n h a v e a variety of o r i g i n s , b u t the t e x t u r e
and o c c u r r e n c e o f d o l o m i t e i n C i n c i n n a t i a n r o c k s m a t c h e s that f o u n d i n
m o d e r n s e m i - a r i d s e t t i n g s w i t h h i g h rates o f e v a p o r a t i o n . Trade w i n d s
c h a r a c t e r i s t i c o f s u b t r o p i c a l l a t i t u d e s w o u l d h a v e b l o w n w e s t w a r d across
the C i n c i n n a t i area d u r i n g t h e O r d o v i c i a n .
The C i n c i n n a t i area w a s s u b j e c t e d t o f r e q u e n t h u r r i c a n e s , w h i c h p r o -
d u c e d the d i s t i n c t i v e a l t e r n a t i o n s o f l i m e s t o n e a n d m u d s t o n e , often c a l l e d
s h a l e , a l t h o u g h few true s h a l e s exist in t h e C i n c i n n a t i area ( F i g u r e 15.2; see
f i g u r e 4.6). A l t h o u g h there is c o n s i d e r a b l e v a r i a t i o n in i n d i v i d u a l s t o r m
deposits as a result of d i f f e r e n c e s in s e d i m e n t supply, w a t e r d e p t h , p r o x i m -
ity t o t h e h u r r i c a n e , a n d the s t r e n g t h o f t h e h u r r i c a n e , m o s t o f t h e s e d e -
posits c o n t a i n a t least s o m e o f t h e c h a r a c t e r i s t i c f e a t u r e s p r o d u c e d b y
storms. S t o r m - g e n e r a t e d deposits are well k n o w n not o n l y from s e d i m e n t
cores on m o d e r n continental shelves, but also t h r o u g h o u t the geologic
record. S t o r m b e d s t y p i c a l l y h a v e a n e r o s i o n a l b a s e , w h i c h reflects p r o g r e s -
sively i n t e n s i f y i n g c u r r e n t s a n d w a v e s as the storm a p p r o a c h e s . This e r o -

Paleogeography and Paleoenvironment 217

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 S T A G E III. C A L M C O N D I T I O N S , P O S T - S T O R M
fine-grained bottom sed-
iments into suspension.
Stronger wave and cur-
rent forces can displace
benthic organisms, and
suspended sediment can
clog feeding and respira-
tory mechanisms of or-
ganisms, and even
smother entire benthic
communities. Mobile or-
ganisms can escape if
burial is not too severe,
but storms can be lethal
for many immobile ben-
thic organisms. Following S T A G E II. S T O R M C O N D I T I O N S
a storm, barren sediment
can cover the bottom
until benthic communi-
ties again develop from
larval settlement or im-
migration. Modified after
Hinterlong (1981). Cour-
tesy of Wayne D. Martin.

S T A G E I. C A L M C O N D I T I O N S , P R E - S T O R M

sional b a s e is often o v e r l a i n by a s h e l l - r i c h l i m e s t o n e , in w h i c h the size of

shell f r a g m e n t s d e c r e a s e s u p w a r d s , w h i c h m a y i n turn b e o v e r l a i n b y l a m i -
n a t e d siltstone a n d b u r r o w e d m u d s t o n e . S u c h a n u p w a r d d e c r e a s e i n shell
a n d s e d i m e n t g r a i n size i s c a l l e d n o r m a l g r a d i n g b y s e d i m e n t o l o g i s t s a n d
reflects d e p o s i t i o n d u r i n g t h e w a n i n g p h a s e o f a s t o r m w h e n s t o r m - g e n e r -
ated w a v e s and c u r r e n t s w e a k e n e d a n d d e p o s i t e d the s e d i m e n t t h e y car-
ried. L a m i n a t e d siltstone m a y d i s p l a y h o r i z o n t a l p l a n a r l a m i n a t i o n s , h u m -
m o c k y c r o s s - l a m i n a t i o n , o r w a v e - r i p p l e l a m i n a t i o n , all o f w h i c h c a n b e
produced by strong storm-generated waves and currents.

218 A Sea without Fish

 C y c l i c a l c h a n g e s in the c h a r a c t e r of storm b e d s are well d e v e l o p e d in
s o m e C i n c i n n a t i a n deposits, such as the K o p e f o r m a t i o n . At a broad s c a l e ,
these roughly meter-thick c y c l e s consist of a m u d s t o n e - r i c h unit a n d a l i m e -
stone-rich unit (see F i g u r e 4.6). T h e m u d s t o n e - r i c h unit consists of 35 cm
beds o f n o r m a l l y graded m u d s t o n e , with u n c o m m o n thin l a m i n a t e d siltstone
b e d s . L i m e s t o n e - r i c h units consist of shelly l i m e s t o n e b e d s , with a lesser
a m o u n t o f thin m u d s t o n e and siltstone b e d s . T h e alternation b e t w e e n these
t w o units w a s originally t h o u g h t to reflect c h a n g e s in sea level, b u t r e c e n t
studies suggest that these c y c l e s m a y instead reflect c h a n g e s in the a v e r a g e
f r e q u e n c y and intensity of h u r r i c a n e s over tens of t h o u s a n d s of years.

C o m p a r e d to man) modern carbonate settings, O r d o v i c i a n limestone of Oceanography

the C i n c i n n a t i area i s u n u s u a l i n several r e g a r d s . M o s t m o d e r n a n d a n -
cient w a r m water c a r b o n a t e d e p o s i t s c o n t a i n a w i d e v a r i e t y o f g r a i n t y p e s ,
i n c l u d i n g skeletal g r a i n s (the shells o f o r g a n i s m s ) , o o i d s ( s m a l l , s p h e r o i d a l ,
c o n c e n t r i c a l l y l a m i n a t e d grains), p e l o i d s (ovoid g r a i n s p r o d u c e d p r i m a r i l y
as fecal pellets), a n d intraclasts ( p i e c e s of s e m i - c e m e n t e d c a r b o n a t e sedi-
m e n t that h a v e b e e n e r o d e d and r e d e p o s i t e d ) . I n the t y p e - C i n c i n n a t i a n ,
o o i d s are a b s e n t , p e l o i d s are u n c o m m o n , a n d intraclasts o c c u r s p a r i n g l y
and o n l y in p a r t i c u l a r h o r i z o n s . In c o n t r a s t , skeletal g r a i n s of b r a c h i o p o d s ,
b r y o z o a n s , e c h i n o d e r m s , m o l l u s c s , and trilobites d o m i n a t e m o s t l i m e s t o n e
i n the C i n c i n n a t i a n (see f i g u r e 4.2). M o s t m o d e r n w a r m w a t e r c a r b o n a t e
deposits c o n t a i n a b u n d a n t l i m e m u d , c a l l e d m i c r i t e , p r o d u c e d p r i m a r i l y
by the p h o t o s y n t h c t i c a c t i v i t i e s of a l g a e . In c o m p a r i s o n , most l i m e s t o n e in
the C i n c i n n a t i area c o n t a i n s o n l y m i n o r a m o u n t s o f m i c r i t e . S e d i m e n t s i n
m o d e r n w a r m water c a r b o n a t e e n v i r o n m e n t s are p r o n e t o u n d e r g o c e m e n -
tation w i t h i n a few c e n t i m e t e r s below the s e d i m e n t s u r f a c e , a n d if c u r r e n t s
or w a v e s strip a w a y the o v e r l y i n g u n c e m e n t e d s e d i m e n t , this e x p o s e s a
hard c o n c r e t e - l i k e s u r f a c e on the sea floor, k n o w n as a h a r d g r o u n d . I lard-
g r o u n d s c a n b e i m p o r t a n t substrata u p o n w h i c h e n c r u s t i n g o r g a n i s m s
such as bryozoans and corals may attach. A l t h o u g h hardgrounds do o c c u r
i n the t y p e - C i n c i n n a t i a n , t h e y are relatively u n c o m m o n c o m p a r e d t o
w a r m water settings b o t h today a n d i n t h e past. T h e f e a t u r e s that t y p i f y
l i m e s t o n e o f the C i n c i n n a t i a r e a a b u n d a n t skeletal g r a i n s , a lack o f o o i d s
and p e l o i d s , m i n i m a l m i c r i t e , a n d u n c o m m o n h a r d g r o u n d s a r e t y p i c a l
of c a r b o n a t e s d e p o s i t e d t o d a y in c o o l t e m p e r a t e to p o l a r waters.
The presence of cool water carbonates at tropical latitudes at first
s e e m s like a p a r a d o x , but s u c h c o n d i t i o n s o c c u r today w h e r e c o a s t a l u p -
w e l l i n g b r i n g s c o o l water up to the s u r f a c e f r o m d e p t h s of less t h a n 200
meters. T h e s e c o o l e r waters also c o n t a i n a b u n d a n t n u t r i e n t s , w h i c h g e n e r -
ate p h o s p h a t e deposits. I n d e e d , the t y p e - C i n c i n n a t i a n is rich in p h o s p h a t e ,
p a r t i c u l a r l y in strata of M a y s v i l l i a n a g e . P h o s p h a t e is also f o u n d in a b u n -
d a n c e i n U p p e r O r d o v i c i a n strata n e a r N a s h v i l l e , T e n n e s s e e , s u g g e s t i n g
that the entire c a r b o n a t e p l a t f o r m f r o m C i n c i n n a t i to N a s h v i l l e w a s a site
of u p w e l l i n g of cool, nutrient-rich water d u r i n g m u c h of the Late O r d o v i -
c i a n . U p p e r O r d o v i c i a n r o c k s o f t h e C i n c i n n a t i area c o n t a i n a g r e a t e r
a m o u n t o f l i m e m u d , m o r e h a r d g r o u n d s , a n d less p h o s p h a t e , w h i c h c o l -

Paleogeography and Paleoenvironment 219

 l e c t i v e l y s u g g e s t a d e c r e a s e in t h e i n t e n s i t y of u p w e l l i n g in t h e latest
Ordovician.
A d d i t i o n a l e v i d e n c e f r o m the rich fossil f a u n a s supports the interpreta-
tion of c o o l waters, f o l l o w e d by a return to w a r m water in the latest O r d o v i -
c i a n . D u r i n g t h e L a t e O r d o v i c i a n , t h e western U n i t e d States and C a n a d a
straddled t h e e q u a t o r . T h e i r c a r b o n a t e s e d i m e n t s are typical o f m o d e r n
w a r m w a t e r settings, so their f a u n a s are interpreted to reflect w a r m water
c o n d i t i o n s . T h e s e areas c o n t a i n a b u n d a n t corals a n d stromatoporoids, with
a diverse array of b r a c h i o p o d s a n d trilobites. In particular, c o l o n i a l r u g o s a n
and t a b u l a t e corals (for e x a m p l e , Tetradium), solitary corals (Grewingkia,
Streptelasma), several brachiopods (Glyptorthis, Plaesiomys, Rhynchotrema,
Hiscobeccus, Lepidocyclus, Holtedahlina, and Leptaena, for e x a m p l e ) , trilo-
bites (Ceraurinus), a n d diverse c e p h a l o p o d s are characteristic of this w a r m
water f a u n a . T h e s e o r g a n i s m s are a b s e n t from E d e n i a n a n d M a y s v i l l i a n
strata in t h e C i n c i n n a t i a r e a , b u t a p p e a r in the R i c h m o n d i a n as the l i m e -
stones b e g i n to reflect a return to w a r m water, low-nutrient c o n d i t i o n s .
T y p e - C i n c i n n a t i a n rocks differ from typical c a r b o n a t e platform deposits
in a n o t h e r s i g n i f i c a n t a s p e c t : t h e a b u n d a n c e of t e r r i g e n o u s m u d , that is, clay
p r o d u c e d b y t h e w e a t h e r i n g o f silica-rich m i n e r a l s s u c h a s feldspar. T h e
earliest influx o f this m u d closely c o i n c i d e s w i t h t h e b e g i n n i n g o f nutrient-
r i c h , c o o l water deposits a t t h e b a s e o f t h e L e x i n g t o n L i m e s t o n e , w h i c h
u n d e r l i e s t y p e - C i n c i n n a t i a n strata. T h e arrival o f c o o l water, nutrients, and
siliciclastic m u d a p p e a r s to h a v e b e e n triggered by the uplift of the Laconic
M o u n t a i n s to t h e east. F i n e - g r a i n e d t e r r i g e n o u s clay and silt w e r e supplied
b y the Q u e e n s t o n D e l t a , a s s u c h s e d i m e n t s c o u l d easily have stayed sus-
p e n d e d in t h e water across t h e A p p a l a c h i a n Basin until d e p o s i t i o n in the
C i n c i n n a t i area. T h e s e m u d s w e r e best able t o a c c u m u l a t e i n relatively
c a l m e r d e e p water e n v i r o n m e n t s that w e r e less d i s t u r b e d b y storms.
T h e Persian G u l f is similar in many respects to eastern L a u r e n t i a d u r i n g
the O r d o v i c i a n . B o t h w e r e d e v e l o p e d as foreland basins, that is, d e e p water
t r o u g h s adjacent to uplifting m o u n t a i n s . B o t h possess a c a r b o n a t e platform
that dips gradually into d e e p water. T h e s e d i m e n t a r y basins are similar in size
a n d c l i m a t e , and b o t h sit at subtropical latitudes p r o n e to storms. L a r g e deltas
supplied b y a b u n d a n t terrigenous s e d i m e n t a d v a n c e d into both basins. T h e
Persian G u l f is notably different in that it lacks u p w e l l i n g of c o o l , nutrient-rich
water, w h i c h u n d e r s c o r e s that no m o d e r n setting is exactly a n a l o g o u s to C i n -
c i n n a t i d u r i n g the O r d o v i c i a n . S o m e have s u g g e s t e d the m o d e r n B a h a m a
platform was similar to the C i n c i n n a t i area d u r i n g the O r d o v i c i a n , but the
B a h a m a platform is flat-topped rather t h a n gently d i p p i n g , is not a foreland
basin adjacent to uplifting m o u n t a i n s that feed a b u n d a n t terrigenous sedi-
m e n t to a d v a n c i n g deltas, a n d lacks u p w e l l i n g of c o o l , nutrient-rich waters.

Marine F o u r m a j o r s e d i m e n t a r y e n v i r o n m e n t s w e r e p r e s e n t d u r i n g the L a t e O r d o -

Environments of v i c i a n o f t h e C i n c i n n a t i A r c h ( F i g u r e 15.3). T o d a y , d i s t i n c t i v e features o f

t h e r o c k s a n d fossils c h a r a c t e r i z e t h e s e e n v i r o n m e n t s . E a c h o f t h e s e e n v i -
t h e Cincinnati Arch
ronments is interpreted based on distinctive rock types and sedimentary
s t r u c t u r e s that are f o u n d in s i m i l a r s e t t i n g s today.

220 A Sea without Fish

 T i d a l flat e n v i r o n m e n t s t o d a y are flat, n e a r l y f e a t u r e l e s s a r e a s t h a t Figure 1 5 . 3 . The four
form b e t w e e n the low tide l i n e a n d the h i g h tide l i n e . T h e s e a r e a s are principal sedimentary

c o v e r e d daily by tides, b u t are s u b j e c t e d to e x t r e m e v a r i a t i o n s in salinity environments of the

type-Cincinnatian. Cin-
and t e m p e r a t u r e on a d a i l y basis.
cinnatian seas generally
I n U p p e r O r d o v i c i a n r o c k s o f the C i n c i n n a t i A r c h , tidal flat e n v i r o n -
deepened northward
m e n t s are p r e s e r v e d a s l a m i n a t e d t o b u r r o w e d d o l o m i t e a n d d o l o m i t i c
from shallow water envi-
l i m e s t o n e c o n t a i n i n g s m a l l a m o u n t s o f c l a y ( F i g u r e 15.4A). T h e p r e s e n c e ronments in central Ken-
o f d o l o m i t e s u g g e s t s s t r o n g levels o f e v a p o r a t i o n , w h i c h w o u l d h a v e d r a w n tucky to deeper water
m a g n e s i u m - r i c h brines t h r o u g h fine-grained limestone and converted it to environments in Ohio
d o l o m i t e . A l t h o u g h s u c h c o n d i t i o n s form t o d a y i n relatively arid s e t t i n g s , and Indiana. The bound-
the lack of o t h e r e v a p o r i t e m i n e r a l s s u c h as h a l i t e or g y p s u m in t h e s e tidal aries between these four

flat facies a r g u e s m o r e for a s e m i - a r i d e n v i r o n m e n t . W a v e - f o r m e d ripple- environments correspond

to sea level, fairweather
m a r k s attest to the shallow w a t e r e n v i r o n m e n t in w h i c h t h e s e r o c k s w e r e
wave base, and storm
d e p o s i t e d , and d e s i c c a t i o n c r a c k s ( " m u d c r a c k s " ) i n d i c a t e the d r y i n g a n d
wave base. Wave base
s h r i n k i n g o f the m u d w h e n i t w a s e x p o s e d d u r i n g p r o l o n g e d low tides.
reflects the depth at
S o m e o f t h e s e strata c o n t a i n n u m e r o u s c l o s e l y s p a c e d p l a n a r l a m i n a e that
which waves can move
record t h e d e p o s i t i o n o f i n d i v i d u a l layers o f c a r b o n a t e m u d d u r i n g i n c o m - sediment on the sea floor
i n g tides a n d storms o n the h i g h e s t part o f t h e tidal flat, w h i c h r e m a i n e d and this depth increases
a b o v e a v e r a g e h i g h tide. I n p l a c e s , t h e s e l a m i n a e are p e n e t r a t e d b y s h o r t , with the height and pe-
vertical b u r r o w s , w h i c h s u g g e s t a r e a s s o m e w h a t l o w e r o n t h e tidal flat riod of waves, both of
w h e r e b u r r o w i n g o r g a n i s m s w o u l d not h a v e b e e n k i l l e d b y d r y i n g o u t a n d which increase during

o v e r h e a t i n g d u r i n g low tides. Elsewhere in t h e C i n c i n n a t i r e g i o n , tidal flat storms. The locations of

these environments
deposits lack p l a n a r l a m i n a t i o n a n d are t h o r o u g h l y b u r r o w e d b y soft-bod-
changed over thousands
ied o r g a n i s m s s u c h a s p o l y c h a e t e w o r m s a n d a r t h r o p o d s . S u c h extensive-
to millions of years, with
b u r r o w i n g w o u l d r e q u i r e m o r e f r e q u e n t a n d p e r s i s t e n t s u b m e r g e n c e dur-
environments shifting
i n g a tidal c y c l e , as on t h e l o w e s t p o r t i o n s of t h e tidal flat. M a n y of t h e s e
northward (as shown in
b u r r o w s are filled with the d i s t i n c t i v e g r e e n iron m i n e r a l g l a u c o n i t e . Tidal figure) during times of
flat deposits are m o s t c o m m o n i n c e n t r a l K e n t u c k y , b u t s o m e tidal f l a t s slowly rising sea level and
a d v a n c e d as far n o r t h w a r d as s o u t h e r n I n d i a n a , as c a n be s e e n in e x p o s u r e s retreating southward
o f the S a l u d a D o l o m i t e n e a r M a d i s o n , I n d i a n a . during times of rapidly
rising sea level. Falls in
M o s t tidal flat d e p o s i t s in the C i n c i n n a t i r e g i o n are u n f o s s i l i f e r o u s ,
sea level resulted in the
b u t l o c a l l y t h e b u r r o w e d d e p o s i t s c o n t a i n a sparse f a u n a o f b r y o z o a n s , a n d
draining of the seas from
m o r e rarely o s t r a c o d s , b r a c h i o p o d s , s t r o m a t o p o r o i d s , a n d r u g o s a n a n d
the Cincinnati area.
tabulate c o r a l s . This restricted g r o u p o f o r g a n i s m s p r e s u m a b l y w o u l d h a v e
b e e n c a p a b l e of t o l e r a t i n g fluctuations in salinity a n d t e m p e r a t u r e . Preser-

Paleogeography and Paleoenvironment 221

Figure 1 5 . 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 shallow
subtidal environ-
ment. C. Outcrop pho-
tograph of interbedded
limestone and mudstone
deposited in a deep sub-
tidal environment, with
the limestone beds re-
cording deposition dur-
ing hurricanes. D. Out-
crop photograph of
mudstone with thin beds
of limestone and silt-
stone, all deposited in an
offshore environment.

v a t i o n of t h e s e fossils is t y p i c a l l y p o o r , as a result of d o l o m i t i z a t i o n , w h i c h
tends to d e s t r o y fine d e t a i l s .
S h a l l o w s u b t i d a l e n v i r o n m e n t s t o d a y a r e shallow marine environ-
m e n t s below the l o w tide l i n e , b u t a b o v e fair w e a t h e r w a v e base, the d e p t h
t o w h i c h w a x e s c a n stir the s e d i m e n t d u r i n g c a l m w e a t h e r . Fair w e a t h e r
Behind the history of w a v e b a s e is t y p i c a l l y o n l y a f e w m e t e r s on coasts p r o t e c t e d from large
every sedimentary rock
o c e a n i c w a v e s . I n m o d e r n c a r b o n a t e s e t t i n g s , s h a l l o w subtidal e n v i r o n -
there lurks an ecosys-
m e n t s are a d j a c e n t to tidal flats a n d are c h a r a c t e r i z e d by intense b u r r o w i n g
tem, but what one
by soft-bodied organisms such as w o r m s and arthropods.
first sees is an environ-
ment of deposition. Shallow subtidal deposits are f o u n d in the type-Cincinnatian a n d are
l i k e w i s e c h a r a c t e r i z e d b y h i g h l y b u r r o w e d shallow m a r i n e deposits that
E d w a r d S. D e e v e y
g r a d e u p w a r d s into tidal flat deposits, i n d i c a t i n g that the t w o w e r e deposited
1965, 592
in laterally a d j a c e n t e n v i r o n m e n t s . In the t y p e - C i n c i n n a t i a n , shallow sub-
tidal deposits consist of n o d u l a r to very thin w a v y - b e d d e d shelly l i m e s t o n e
a n d fossiliferous m u d s t o n e ( F i g u r e 1 5 . 4 B ) . B e c a u s e o f the t h i n n e s s and w a v i -
ness of the l i m e s t o n e b e d s , t h e s e rocks w e a t h e r to a characteristic rubble of
fist-sized l i m e s t o n e n o d u l e s . This distinctive b e d d i n g results from the per-

222 A Sea without Fish

vasive b u r r o w i n g of the s e d i m e n t by soft-bodied o r g a n i s m s . A l t h o u g h storms
c e r t a i n l y reworked the s e d i m e n t and d e p o s i t e d the c h a r a c t e r i s t i c well-sorted
layers of shells overlain by layers of m u d that are preserved in s o m e p l a c e s ,
s u b s e q u e n t b u r r o w i n g m i x e d these l a y e r s , p r o d u c i n g p o d s o f shell-rich a n d
shell-poor material. Preferential c e m e n t a t i o n o f these c h u r n e d s e d i m e n t s
p r o d u c e d p o c k e t s of w e l l - c e m e n t e d shells material s u r r o u n d e d by n o n - c e -
m e n t e d z o n e s rich in clay.
S h a l l o w subtidal l i m e s t o n e in t h e C i n c i n n a t i a r e a is l o c a l l y r i c h in
p h o s p h a t e , p a r t i c u l a r l y i n t h e M a y s v i l l i a n . M u c h o f this p h o s p h a t e o c c u r s
a s infillings o f b r y o z o a n z o o e c i a , the p o r o u s s k e l e t o n s o f e c h i n o d e r m s , a n d
the larval shells of pelecypods, g a s t r o p o d s (such as Cyclora), a n d m o n o p l a -
cophorans. The p r e s e n c e o f this p h o s p h a t e i n d i c a t e s large a m o u n t s o f d e -
c a y i n g o r g a n i c matter w i t h i n t h e s e d i m e n t . B y d i s s o l v i n g p i e c e s o f s h a l l o w
subtidal l i m e s t o n e i n v i n e g a r o r d i l u t e h y d r o c h l o r i c a c i d , o n e c a n see t h e
rich f a u n a p r e s e r v e d b y this p h o s p h a t i z a t i o n . S h a l l o w s u b t i d a l r o c k s are
broadly d i s t r i b u t e d o v e r t h e C i n c i n n a t i A r c h a n d o c c u r f r o m t h e s o u t h e r n
e d g e o f the O r d o v i c i a n o u t c r o p belt i n s o u t h e r n K e n t u c k y t o the n o r t h e r n
l i m i t o f O r d o v i c i a n rocks i n c e n t r a l O h i o a n d I n d i a n a . T h e B e l l e v u e , M t .
A u b u r n , O r e g o n i a , a n d W h i t e w a t e r F o r m a t i o n s all a c c u m u l a t e d within
shallow subtidal e n v i r o n m e n t s .
Shallow s u b t i d a l rocks a r e e x c e e d i n g l y fossiliferous i n m o s t p l a c e s ,
r e f l e c t i n g the a b u n d a n c e o f life i n this shallow m a r i n e habitat. M o s t c o m -
m o n l y , shallow s u b t i d a l r o c k s are p a c k e d with large brachiopods, s u c h as
Platystrophia, Hebertella, a n d Rafinesquina. M a n y of t h e s e have t h i c k or
c o a r s e l y r i b b e d s h e l l s , p r e s u m a b l y for p r o t e c t i o n a g a i n s t w a v e s a n d c u r -
rents. Platystrophia, in p a r t i c u l a r , has a g r e a t l y t h i c k e n e d p e d i c l e v a l v e
n e a r the h i n g e , w h i c h w o u l d h a v e i n c r e a s e d t h e stability o f t h e shell o n t h e
sea floor. D i s a r t i c u l a t i o n , b r e a k a g e , a n d a b r a s i o n o f t h e s e shells a r e w i d e -
spread a n d attest t o the d a m a g i n g effects o f w a v e s a n d c u r r e n t s . L a r g e
b r y o z o a n s a r e often a b u n d a n t a n d i n c l u d e b r a n c h i n g , e n c r u s t i n g , sheet-
like, a n d m a s s i v e f o r m s . As is t r u e for the brachiopods, t h e s e r o b u s t b r y o -
z o a n skeletons reflect the intensity o f w a v e s a n d c u r r e n t s i n this shallow
water e n v i r o n m e n t . M o l l u s c s are p r e s e n t in shallow s u b t i d a l r o c k s , p a r t i c u -
larly t h e byssally attached pelecypods Ambonychia a n d Caritodens, t h e
carnivorous cephalopod Treptoceras, a n d the g a s t r o p o d s Lophospira a n d
Cyclonema. Cyclonema is c o m m o n l y a s s o c i a t e d with c r i n o i d s , on w h i c h it
m a y have b e e n a parasite. Lophospira has b e e n i n t e r p r e t e d as a s c a v e n g e r .
C r i n o i d s and trilobites o c c u r , but m o s t s p e c i m e n s a r e d i s a r t i c u l a t e d rather
t h a n w h o l e , p r e s u m a b l y o w i n g not o n l y t o w a v e s a n d c u r r e n t s , b u t also
burrowing organisms.
Deep s u b t i d a l e n v i r o n m e n t s t o d a y are t h o s e that lie b e l o w fair
w e a t h e r w a v e base, but a b o v e the w a v e b a s e o f all b u t t h e most p o w e r f u l
storms or h u r r i c a n e s , w h i c h w o u l d have e x t e n d e d to d e p t h s of a several
tens of meters. In t h e s e s e t t i n g s , t h e s e d i m e n t a r y d e p o s i t s are c h a r a c t e r i z e d
b y the a l t e r n a t i o n o f s a n d y a n d shelly b e d s d e p o s i t e d d u r i n g s t o r m s and
muddy b e d s that reflect quiet water d e p o s i t i o n d u r i n g w e a k s t o r m s or dur-
i n g p e r i o d s b e t w e e n s t o r m s . D e e p s u b t i d a l e n v i r o n m e n t s are a d j a c e n t t o
a n d slightly d e e p e r t h a n s h a l l o w s u b t i d a l e n v i r o n m e n t s .

Paleogeography and Paleoenvironment 223

 A s o n m o d e r n s h e l v e s , storm d e p o s i t s are the m o s t c o n s p i c u o u s feature
of the d e e p subtidal e n v i r o n m e n t in the t y p e - C i n c i n n a t i a n , with roughly
e q u a l p r o p o r t i o n s o f t h i n t o m e d i u m - b e d d e d shelly l i m e s t o n e , l a m i n a t e d
siltstones, a n d m u d s t o n e ( F i g u r e 15.4C). B u r r o w i n g is m u c h less i n t e n s e
here than in d e e p subtidal environments, resulting in thicker and more
laterally c o n t i n u o u s l i m e s t o n e b e d s . B e d s o f siltstone are c o m m o n l y rip-
pled or display internal planar or h u m m o c k y lamination generated by
s t r o n g storm c u r r e n t s a n d w a v e s . A t t i m e s , s t o r m s o c c u r r e d w i t h sufficient
f r e q u e n c y that t h e y c o m m o n l y e r o d e d t h r o u g h t h e m u d s t o n e layer c a p -
p i n g t h e d e p o s i t f r o m t h e p r e v i o u s s t o r m , s u c h that the shelly layer f r o m
o n e s t o r m w a s d e p o s i t e d d i r e c t l y o n t h e s h e l l y b e d o f t h e p r e v i o u s storm.
T h i s p h e n o m e n o n , k n o w n a s a m a l g a m a t i o n , p r o d u c e s thick layers o f l i m e -
s t o n e w i t h s u b t l e i n t e r n a l e r o s i o n s u r f a c e s t h a t s e p a r a t e i n d i v i d u a l storm
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 a s m u l t i - e v e n t b e d s . I n s o m e
c a s e s , a o n e - f o o t t h i c k b e d o f l i m e s t o n e m a y r e c o r d h a l f a d o z e n storm
e v e n t s . D e e p s u b t i d a l r o c k s are a s b r o a d l y d i s t r i b u t e d o v e r t h e C i n c i n n a t i
A r c h as shallow subtidal rocks. The F a i r v i e w , C o r r y v i l l e , S u n s e t , a n d L i b -
erty F o r m a t i o n s a c c u m u l a t e d i n d e e p s u b t i d a l e n v i r o n m e n t s .
D e e p subtidal rocks of the t y p e - C i n c i n n a t i a n contain an abundant
and diverse fauna. Preservation is c o m m o n l y better than in shallow sub-
tidal r o c k s , w i t h less o v e r a l l d i s a r t i c u l a t i o n , b r e a k a g e , a n d a b r a s i o n , sug-
g e s t i n g less e x p o s u r e t o t h e d a m a g i n g effects o f w a v e s a n d c u r r e n t s . M a n y
brachiopod genera m a y be p r e s e n t , i n c l u d i n g Rafinesquina, Strophomena,
Leptaena, Hiscobeccus, Platystrophia, Plectorthis, Glyptorthis, and Plaesio-
mys. A l l h a v e shells t h a t are t h i n n e r a n d finer-ribbed t h a n t h o s e in t h e
s h a l l o w s u b t i d a l . B r y o z o a n s c a n b e a b u n d a n t a n d also t e n d t o b e t h i n n e r
a n d less m a s s i v e t h a n i n t h e s h a l l o w s u b t i d a l . M o l l u s c s are c o m m o n , w i t h
s i m i l a r f o r m s as in t h e s h a l l o w s u b t i d a l , as w e l l as t h e byssally a t t a c h e d
pelecypod Modiolopsis. Crinoids (Glyptocrinus, Pycnocrinus, and Iocrinus)
a n d e d r i o a s t e r o i d s (Carneyella, Isorophus, a n d Streptaster) are l o c a l l y a b u n -
d a n t , a n d b e d s a n d p o c k e t s c o n t a i n i n g f u l l y a r t i c u l a t e d s p e c i m e n s are n o t
u n u s u a l . T r i l o b i t e s (Isotelus a n d Flexicalymene) are c o m m o n b o t h as indi-
v i d u a l sclerites a n d a s a r t i c u l a t e d s p e c i m e n s . The frequency of articulated
c r i n o i d s a n d trilobites s u g g e s t s early b u r i a l a n d less f r e q u e n t d i s t u r b a n c e
by waves, currents, and b u r r o w i n g organisms. In the uppermost C i n c i n -
n a t i a n , solitary c o r a l s (Grewingkia a n d Streptelasina), a n d the encrusting
t a b u l a t e c o r a l , Protaraea, are c o n s p i c u o u s a d d i t i o n s to t h e d e e p s u b t i d a l
f a u n a . T r a c e fossils are c o m m o n i n t h e siltstone b e d s . Chondrites a n d
Trichophycus were the burrows of deposit feeding worms or arthropods.
T h e U - s h a p e d b u r r o w s o f Diplocraterion w e r e t h e h o m e s o f a p o l y c h a e t e
w o r m , b u t w h e t h e r it w a s a s u s p e n s i o n f e e d e r , a s t a t i o n a r y d e p o s i t f e e d e r ,
o r a n a m b u s h c a r n i v o r e i s u n c e r t a i n , a s m o d e r n e x a m p l e s o f all s u c h U -
t u b e b u i l d e r s are k n o w n . Paleophycus r e c o r d s t h e h o r i z o n t a l b u r r o w i n g o f
another scavenging or deposit feeding w o r m .
T h e t h i c k , t a b u l a r l i m e s t o n e b e d s o f t h e d e e p s u b t i d a l are well suited
a s b u i l d i n g s t o n e s . M a n y old q u a r r i e s w e r e e s t a b l i s h e d i n d e e p subtidal
r o c k s a n d m a n y o f t h o s e s t o n e s c a n n o w b e f o u n d i n old b u i l d i n g f o u n d a -
t i o n s a n d r o c k w a l l s . T w o i n t e r v a l s o f d e e p s u b t i d a l rocks f r e q u e n t e d b y

224 A Sea without Fish

q u a r r y m e n w e r e the River Q u a r r y B e d s ( n o w c a l l e d t h e Point Pleasant
F o r m a t i o n ) and the Hill Q u a r r y B e d s (now c a l l e d the F a i r v i e w F o r m a t i o n ) .
A l t h o u g h the River Q u a r r y B e d s n e a r C i n c i n n a t i a r e n o w largely u n d e r
the O h i o River, w h o s e level w a s raised d u r i n g t h e c o n s t r u c t i o n o f d a m s ,
they c a n still b e s e e n n e a r Point P l e a s a n t , O h i o , a l o n g t h e crest o f the
C i n c i n n a t i A r c h . M a n y o f t h e H i l l Q u a r r i e s c a n still b e s e e n i n t h e bluffs
south o f the U n i v e r s i t y o f C i n c i n n a t i a n d f l a n k i n g t h e M i l l C r e e k V a l l e y .
O f f s h o r e e n v i r o n m e n t s o n m o d e r n c o a s t s lie below t h e w a v e base o f
m o s t s t o r m s , but a r e s o m e t i m e s a f f e c t e d b y the most severe s t o r m s a n d
extend to d e p t h s of several t e n s of m e t e r s . In t h e s e m o d e r n s e t t i n g s , d e p o s i -
tion i s d o m i n a t e d b y m u d s , w h i c h c a n a c c u m u l a t e w h e n c u r r e n t s a n d
waves are w e a k . Rare, e x c e p t i o n a l l y s t r o n g s t o r m s a r e c a p a b l e o f m o v i n g
shells and s e d i m e n t s e v e n a t t h e s e d e p t h s a n d p r o d u c e t h i n storm b e d s ,
a l t h o u g h these m a k e u p a m i n o r i t y o f the d e p o s i t s . O f f s h o r e e n v i r o n m e n t s
are adjacent t o a n d s o m e w h a t d e e p e r t h a n d e e p s u b t i d a l s e t t i n g s .
In the t y p e - C i n c i n n a t i a n , o f f s h o r e r o c k s c o n t a i n a greater p r o p o r t i o n
o f m u d s t o n e ( c o m m o n l y n e a r two-thirds), but i n o t h e r regards are q u i t e
s i m i l a r to the deep s u b t i d a l ( F i g u r e 15.4D). The less f r e q u e n t o c c u r r e n c e
of storm b e d s in offshore d e p o s i t s i n d i c a t e s less f r e q u e n t d i s t u r b a n c e by
s t o r m - g e n e r a t e d w a v e s a n d c u r r e n t s . As a result, a m a l g a m a t i o n is m u c h
less c o m m o n in t h e o f f s h o r e , and m o s t l i m e s t o n e b e d s are s i n g l e - e v e n t
beds and record the passage of a single h u r r i c a n e . Many of the thick mud-
stone layers f o u n d in t h e offshore are also the result of storm d e p o s i t i o n , as
i n d i c a t e d by the p r e s e n c e of m u l t i p l e 2-3 cm f i n i n g - u p w a r d m u d s t o n e
b e d s , e a c h with a slightly silty interval at its b a s e . E a c h of t h e s e t h i n layers
r e c o r d s the m i n o r d i s t u r b a n c e of the sea floor by a h u r r i c a n e , w i t h s e t t l i n g
o f silt and t h e n c l a y f o l l o w i n g the s t o r m . F r e q u e n t l y , s u c h m u d layers w o u l d
b l a n k e t the b o t t o m , s m o t h e r t h e f a u n a l i v i n g o n the sea floor, a n d p r e s e r v e
a r t i c u l a t e d c r i n o i d s a n d trilobites. O f f s h o r e r o c k s o c c u r a s tar s o u t h a s
n o r t h - c e n t r a l K e n t u c k y but e x t e n d b e y o n d the n o r t h e r n l i m i t o f O r d o v i -
cian exposures in central O h i o and Indiana. The K o p e a n d W a y n e s v i l l e
F o r m a t i o n s largely reflect offshore s e t t i n g s .
O f f s h o r e strata c o n t a i n a n a b u n d a n t a n d diverse f a u n a , c h a r a c t e r i z e d
b y s m a l l , t h i n , a n d d e l i c a t e fossils, s u g g e s t i n g g e n e r a l l y q u i e t w a t e r c o n d i -
tions, e x c e p t d u r i n g rare, severe s t o r m s . C o m m o n b r a c h i o p o d s i n c l u d e t h e
highly gregarious Dalmanella a n d Sowerbyella, and the burrowing inar-
ticulates Pseudolingula a n d Leptobolus, w h i c h are sometimes found pre-
served inside their vertical b u r r o w s . A l t h o u g h sheet-like b r y o z o a n s d o o c -
c u r , t h i n b r a n c h i n g f o r m s a n d flat d i s c - s h a p e d f o r m s are m o r e c o m m o n .
M o l l u s c s are d i v e r s e a n d a b u n d a n t in a few w i d e l y t r a c e a b l e h o r i z o n s ,
which are conspicuously poor in brachiopods and bryozoans. C o m m o n
m o l l u s c s i n c l u d e the byssally a t t a c h e d pelecypods Ambonychia a n d Modi-
olopsis, the b u r r o w i n g p e l e c y p o d s Deceptrix and Rhytimya, t h e scavenging
gastropods Lophospira a n d Liospira, the m o n o p l a c o p h o r a n Sinuites, a n d
the cephalopods Treptoceras a n d Cameroceras. T r i l o b i t e s are n u m e r o u s
and f r e q u e n t l y fully a r t i c u l a t e d . T h e b u r r o w e r s Flexicalymene, Gravicaly-
mene, Cryptolithus, a n d Isotelus are the m o s t c o m m o n , b u t the spiny s w i m -
m i n g Acidaspis c a n b e a b u n d a n t in s o m e c r i n o i d - r i c h layers. S u g g e s t i v e of

Paleogeography and Paleoenvironment 225

 d e e p w a t e r s e t t i n g s is t h e p r e s e n c e of t h e blind trilobites Cryptolithus and
Triarthrus in offshore strata. C r i n o i d s are also n u m e r o u s and are f r e q u e n t l y
articulated. The most c o m m o n genera are Cincinnaticrinus a n d Fxteno-
crinus, w h o s e ossicles m a y c o m p r i s e e n t i r e b e d s o f l i m e s t o n e . G i v e n t h e
t e n s o f k i l o m e t e r s o v e r w h i c h s u c h b e d s c a n b e t r a c e d , the n u m b e r o f cri-
noid individuals must have b e e n astronomical. As in the d e e p subtidal,
t r a c e fossils are n u m e r o u s in b e d s of siltstone. Chondrites, Diplocraterion,
Trichophycus a n d Paleophycus are all c o m m o n . T h e trilobite b u r r o w Ruso-
phycus is a l s o c o m m o n , a n d e x a m p l e s of Rusophycus m a d e by Isotelus and
Cryptolithus h a v e b e e n r e p o r t e d , but o n e s p r o d u c e d b y the c a l y m e n i d s
Gravicalymene a n d F l e x i c a l y m e n e are m u c h m o r e c o m m o n .

226 A Sea without Fish

228 A Sea without Fish
 16
LIFE IN THE CINCINNATIAN SEA

The Ecological Theater and the Evolutionary Play is a b o o k of f a s c i n a t i n g es- The Ecological Theater
says a b o u t the c o m p l e x interactions b e t w e e n the e n v i r o n m e n t and the o r g a n - and the Evolutionary Play
isms i n h a b i t i n g it. It was written by the e c o l o g i s t G. E. H u t c h i n s o n in 1965. G. E. H u t c h i n s o n 1965
Hutchinson's title, an extrapolation of the S h a k e s p e a r e a n m e t a p h o r , provides
a useful a n a l o g y by w h i c h to v i e w the C i n c i n n a t i a n as a Series of acts in t h e
e v o l u t i o n a r y play. In c h a p t e r s 5-14 of o u r b o o k we i n t r o d u c e d t h e cast of
characters, the players on the stage, of the L a t e O r d o v i c i a n sea that c o v e r e d
Figure 16.1. A. Internal
the C i n c i n n a t i A r c h region. H a v i n g read these chapters, the reader s h o u l d be
mold of nautiloid, Trep-
able to r e c o g n i z e the c h a r a c t e r s and know s o m e t h i n g of their r o l e s i n par- toceras duseri (Hall and
ticular their m o d e s of life and f e e d i n g habits. Whitfield), with crinoid
In scientific t e r m s , this is the r e a l m of a u t e c o l o g y , the r e l a t i o n s h i p of Xenocrinus baeri
organisms of a single species with the e n v i r o n m e n t , assessing the influence (Meek) preserved within
body chamber. Richard
o f c h e m i c a l , p h y s i c a l , a s w e l l a s b i o l o g i c a l factors. G i v e n that w e are d e a l -
Arnold Davis collection,
ing w i t h o r g a n i s m s l o n g s i n c e e x t i n c t , p e r h a p s w e s h o u l d add the prefix for
Waynesville Formation,
"ancient" and enter the a n c i e n t realm of paleoautecology.
Adams Co., Ohio, col-
For m a r i n e o r g a n i s m s , m a j o r i n o r g a n i c factors i n c l u d e t e m p e r a t u r e , lected by Thomas T.
salinity, o x y g e n c o n t e n t o f the water, n a t u r e o f t h e sea b o t t o m , w a t e r m o v e - Johnson. B. Ophiuroid,
m e n t ( b o t h c u r r e n t s and w a v e - i n d u c e d t u r b u l e n c e ) , a n d t u r b i d i t y (sedi- Taeniaster spinosus
m e n t c o n t e n t o f t h e water). C h a n g e s i n hydrostatic pressure a s s o c i a t e d w i t h (Billings), MUGM 28187,

water d e p t h do not exert a m a j o r i n f l u e n c e e x c e p t in o r g a n i s m s l i k e fish preserved on internal

mold of nautiloid,
with air b l a d d e r s or a i r - b r e a t h i n g , d i v i n g m a r i n e vertebrates. However, all
Waynesville Formation,
o f t h e o t h e r factors listed c a n vary s i g n i f i c a n t l y w i t h w a t e r d e p t h , a n d ,
Butler Co., Ohio, scale in
h e n c e , d e p t h itself o f t e n is c o r r e l a t e d w i t h c h a n g e s in t h e d i s t r i b u t i o n of
mm. C. Trilobites, Aci-
s p e c i e s . B i o l o g i c a l factors i n c l u d e t h e m o d e o f f e e d i n g , food availability, daspis sp., preserved on
and a host of potential i n t e r a c t i o n s w i t h o r g a n i s m s of o t h e r s p e c i e s s u c h as internal mold of nau-
predator-prey i n t e r a c t i o n s , c o m m e n s a l i s m , a n d p a r a s i t i s m . tiloid, ?Treptoceras sp.,
In c h a p t e r 15, S t e v e n M. H o l l a n d d e s c r i b e d t h e s t a g e s e t t i n g s for t h e CMC IP 2257, Cincinna-

C i n c i n n a t i a n play, the p a l e o g e o g r a p h y a n d e n v i r o n m e n t s o f t h e L a t e O r - tian, vicinity of Cincinnati,

Ohio, x 0.9. D. Trilo-
d o v i c i a n sea in w h i c h the o r g a n i s m s lived. In order to u n d e r s t a n d t h e e n -
bites, Flexicalymene
tire play, we n o w m u s t c o n s i d e r t h e ( C i n c i n n a t i a n players as an a s s e m b l e d
meeki (Foerste), pre-
cast o n that stage. How did the cast c h a n g e w i t h e a c h a c t , a n d how did t h e
served on internal mold
players interact with o n e a n o t h e r a s t h e plot u n f o l d e d ? of nautiloid, ?Trepto-
R e a s s e m b l y o f the cast o f c h a r a c t e r s and the interplay b e t w e e n t h e m ceras sp., OSU 50329,
brings us to the s u b d i s c i p l i n e of s y n e c o l o g y (or, in this c a s e , p a l e o s y n e c o l - Cincinnatian, vicinity of
ogy), t h e relationships o f the a n i m a l s o f the m a n y C i n c i n n a t i a n s p e c i e s that Cincinnati, Ohio, x 2.6.
C, D from Davis et al.
lived together w i t h their c h e m i c a l , p h y s i c a l , and b i o l o g i c a l e n v i r o n m e n t . I n
(2001, figures 2, 5), and
order to d e t e r m i n e how extinct s p e c i e s w e r e i n f l u e n c e d by o t h e r s p e c i e s and
reprinted by permission
their e n v i r o n m e n t , a p a l e o e c o l o g i s t m u s t first g r a p p l e w i t h a very difficult
of Blackwell Publishing.
question: what species a c t u a l l y lived t o g e t h e r at a g i v e n t i m e in t h e past?

229
 W h a t cast w a s o n stage for a g i v e n act? A n e c o l o g i s t c a n observe a n d s a m p l e
l i v i n g o r g a n i s m s i n the f i e l d , b u t t h e p a l e o e c o l o g i s t m u s t d e a l w i t h assem-
b l a g e s of d e a d r e m a i n s preserved in s e d i m e n t a r y rock. Factors a f f e c t i n g fossil
p r e s e r v a t i o n d i s c u s s e d in c h a p t e r 1 are of t h e u t m o s t i m p o r t a n c e here. A r e
t h e s e a s s e m b l a g e s representative of a c t u a l life a s s e m b l a g e s or are t h e y d e a t h
a s s e m b l a g e s r e p r e s e n t i n g m i x t u r e s of o r g a n i s m s that lived at different t i m e s
o r p l a c e s a n d a c c u m u l a t e d g r a d u a l l y o v e r t i m e (time-averaged assemblages)
o r s u d d e n l y i n s o m e q u i c k event? H o w m u c h i n f o r m a t i o n i s m i s s i n g from
the fossil record b e c a u s e of preservational bias? Fossil a s s e m b l a g e s are biased
in favor of o r g a n i s m s w i t h preservable r e m a i n s w i t h hard parts like shells,
skeletons, and e x o s k e l e t o n s , a n d t h e y are biased a g a i n s t o r g a n i s m s l a c k i n g
h a r d parts. C r i t e r i a s u c h as t h o s e p r e s e n t e d in T a b l e 1 in chapter 1 c a n be
a p p l i e d t o a n s w e r t h e s e q u e s t i o n s . T h r o u g h o u t this b o o k , e x a m p l e s o f C i n -
c i n n a t i a n fossils p r e s e r v e d in life position or in d i r e c t association with o r g a n -
isms o f o t h e r s p e c i e s p r o v i d e e v i d e n c e b y w h i c h t o d i s t i n g u i s h life assem-
b l a g e s f r o m d e a t h a s s e m b l a g e s . K e e p i n g t h e s e issues i n m i n d , w e c a n
p r o c e e d t o e x a m i n e h o w fossil a s s e m b l a g e s vary t h r o u g h t h e C i n c i n n a t i a n
a n d w h a t this reveals a b o u t C i n c i n n a t i a n p a l e o s y n e c o l o g y .

E v e r s i n c e s o m e o f t h e e a r l i e s t s t u d i e s o f C i n c i n n a t i a n fossils a n d strata,
p a l e o n t o l o g i s t s h a v e r e c o g n i z e d that a s s e m b l a g e s o f o r g a n i s m s o f e x t i n c t
species c h a n g e markedly through the section and show a close relationship
t o t h e c h a r a c t e r o f t h e r o c k (see c h a p t e r 5). T h e q u o t a t i o n f r o m N i c k l e s
(1902) in c h a p t e r 4 d e m o n s t r a t e s that he r e c o g n i z e d the p a l e o e n v i r o n m e n -
tal s i g n i f i c a n c e o f t h e l i t h o l o g i e s a n d their a s s o c i a t e d fossil f a u n a s . P a l e o n -
tologists c o m p i l e d lists o f fossils c h a r a c t e r i s t i c o f e a c h f o r m a t i o n . Fossils o f
o r g a n i s m s o f s o m e s p e c i e s o c c u r i n m a n y f o r m a t i o n s and thus h a v e l o n g
s t r a t i g r a p h i c r a n g e s ; fossils of o t h e r s p e c i e s are restricted to s i n g l e f o r m a -
t i o n s o r t h i n n e r i n t e r v a l s w i t h i n a f o r m a t i o n . E l i z a b e t h D a l v e (1948) c o m -
p i l e d f a u n a l lists for e a c h C i n c i n n a t i a n r o c k - u n i t b a s e d o n m a n y p r e v i o u s
s t u d i e s , b u t h e r p a p e r i s n o t w i d e l y a v a i l a b l e . T h e w e l l - k n o w n biostrati-
g r a p h i c z o n a t i o n o f t h e C i n c i n n a t i a n that i s still w i d e l y u s e d ( C a s t e r e t al.
1955; D a v i s 1985, 1992) e x p r e s s e s t h e s e f a u n a l c h a n g e s . U s i n g this k i n d o f
i n f o r m a t i o n o n fossil d i s t r i b u t i o n o n e c o u l d p r e d i c t w h a t s p e c i e s m i g h t b e
f o u n d in a g i v e n f o r m a t i o n , b u t t h e a s s e m b l a g e s of s p e c i e s in a f o r m a t i o n
a n d their relative a b u n d a n c e w e r e less w e l l k n o w n .
B e g i n n i n g i n t h e late 1960s, t h e b u r g e o n i n g s u b d i s c i p l i n e o f p a l e o -
e c o l o g y f o c u s e d t h e a t t e n t i o n o f p a l e o n t o l o g i s t s o n a s s e m b l a g e s o f fossils
o c c u r r i n g together and their relationship to the e n c l o s i n g sedimentary
m a t r i x . T o w h a t e x t e n t d i d fossil a s s e m b l a g e s r e p r e s e n t e c o l o g i c a l c o m -
munities c o m p a r a b l e to present-day m a r i n e b e n t h i c c o m m u n i t i e s ? M a r i n e
ecologists r e c o g n i z e d c o m m u n i t i e s as recurrent assemblages of species
inhabiting particular environments characterized by particular water
d e p t h , b o t t o m t y p e , t e m p e r a t u r e , salinity, o r o t h e r c h e m i c a l o r p h y s i c a l
a t t r i b u t e s . T h e y d e l i m i t e d a s s e m b l a g e s o n t h e basis o f statistical analysis o f
s a m p l e s r e c o v e r e d f r o m t h e sea floor. C o m m u n i t i e s w e r e n a m e d for d o m i -
nant or characteristic species.

230 A Sea without Fish

 T h e e x i s t i n g f a u n a l lists o f C i n c i n n a t i a n fossils w e r e i n a d e q u a t e for
this t y p e o f a n a l y s i s , s o p a l e o e c o l o g i s t s c o l l e c t e d n e w s a m p l e s i n w h i c h t h e
a b u n d a n c e a s well a s s i m p l e o c c u r r e n c e o f fossil s p e c i e s w a s r e c o r d e d f r o m
censuses of bed surfaces or bulk samples of rock. O n e of the p i o n e e r i n g
s t u d i e s of this t y p e w a s that of Peter Bretsky (1970), w h o r e c o g n i z e d a series
o f fossil c o m m u n i t i e s o f C i n c i n n a t i a n a g e in the A p p a l a c h i a n Basin.
Bretsky's c o m m u n i t i e s w e r e c h a r a c t e r i z e d b y b r a c h i o p o d s , m o l l u s c s , a n d
a n i m a l s o f o t h e r taxa a n d w e r e d i s t r i b u t e d a c c o r d i n g t o d e p t h i n p a r a l l e l
b a n d s c l o s e t o t h e s h o r e l i n e o f t h e s a m e e p i c o n t i n e n t a l sea t h a t e x t e n d e d
w e s t w a r d t o the C i n c i n n a t i A r c h r e g i o n . O t h e r s t u d i e s f o c u s e d o n c o m -
munity p a l e o e c o l o g y of the C i n c i n n a t i A r c h region, such as the work of
F o x (1962, 1968), L o r e n z (1973), O l d r o y d (1978), a n d H a r r i s a n d M a r t i n
(1979) (see c h a p t e r 8). S u b s e q u e n t l y , u s e o f t h e t e r m fossil c o m m u n i t y b e -
c a m e less f r e q u e n t , b e c a u s e i n c r e a s e d u n d e r s t a n d i n g o f t a p h o n o m i c p r o -
c e s s e s s h o w e d that m o s t o f t h e t i m e - a v e r a g e d a s s e m b l a g e s o f fossils are n o t
directly comparable to present-day c o m m u n i t i e s .
In recent q u a n t i t a t i v e studies of C i n c i n n a t i a n fossil a s s e m b l a g e s , fossil
s p e c i m e n s o f i n d i v i d u a l s f r o m e x t i n c t taxa (either s p e c i e s o r g e n e r a ) are
treated as v a r i a b l e s t a k e n from s a m p l e s restricted to a s i n g l e b e d s u r f a c e or
d i s a g g r e g a t e d from a t h i n b e d of l i m e s t o n e or s h a l e . In e a c h s a m p l e , the
a b u n d a n c e o f fossil s p e c i m e n s o f e a c h t a x o n i s c o u n t e d a n d t a b u l a t e d .
In t h e study of fossil a s s e m b l a g e s in t h e C1 ( K o p e F o r m a t i o n ) d e p o s i -
tional s e q u e n c e , H o l l a n d , M i l l e r , M e y e r , a n d D a t t i l o (2001) c o l l e c t e d s a m -
ples from every fossiliferous b e d t h r o u g h a s e v e n t y m e t e r s t r a t i g r a p h i c
s e c t i o n 1 9 4 9 s a m p l e s in all. In e a c h s a m p l e , the relative a b u n d a n c e of fossils
was r e c o r d e d in the field as rare (1-2 s p e c i m e n s per 1000 cm 2
of b e d d i n g
surface), c o m m o n (3-10 s p e c i m e n s p e r 1000 c m ) , or a b u n d a n t (>10 s p e c i -
2

m e n s per 1000 c m ) . Fossils w e r e identified to g e n u s , a n d i n c l u d e d b r a c h i o -

2

pods, c r i n o i d s , trilobites, p e l e c y p o d s , c e p h a l o p o d s , a n d g a s t r o p o d s . S o m e
distinctive b r y o z o a n s w e r e identified to g e n u s , w h e r e a s others w e r e classified
on the basis of c o l o n y m o r p h o l o g y as t h i n bifoliate (<5 m m ) , thick bifoliate
(>5 m m ) , t h i n r a m o s e or b r a n c h i n g (<5 m m ) , t h i c k r a m o s e (>5 m m ) , or e n -
crusting. Fifty-seven taxa w e r e t a b u l a t e d f r o m t h e 1949 s a m p l e s . A f t e r re-
m o v a l of all taxa o c c u r r i n g in just a single s a m p l e , as well as the r e m o v a l of
all s a m p l e s c o n t a i n i n g s p e c i m e n s of just a s i n g l e t a x o n , the final dataset
formed a matrix of forty-six taxa or c o l o n y forms a n d 1337 s a m p l e s .
Large datasets of this type c a n be a n a l y z e d u s i n g several kinds of c o m -
puter-driven, m u l t i v a r i a t e statistical t e c h n i q u e s . I n t h e w o r k o f H o l l a n d ,
Miller, M e y e r , and D a t t i l o (2001), a t e c h n i q u e c a l l e d d e t r e n d e d c o r r e s p o n -
d e n c e analysis ( D C A ) w a s u s e d , a l t h o u g h o t h e r t e c h n i q u e s , for e x a m p l e ,
cluster analysis, factor analysis, a n d p o l a r o r d i n a t i o n , p r o d u c e s i m i l a r results.
D C A c a l c u l a t e s a n u m e r i c a l s c o r e for e a c h t a x o n a n d e a c h s a m p l e a n d plots
t h e m a l o n g g r a p h i c a l axes that reflect similarity of taxa as g r o u p e d in s a m p l e s
or similarity of s a m p l e s based on their taxa. Ecologists h a v e f o u n d that t e c h -
niques like D C A are very useful t o e x a m i n e h o w taxa are arrayed a l o n g these
axes, w h i c h c o m m o n l y c o r r e s p o n d t o s o m e e n v i r o n m e n t a l p a r a m e t e r o r
gradient. I n the analysis o f the K o p e F o r m a t i o n , n u m e r i c a l v a l u e s o f taxa
a l o n g o n e DCA axis displayed a shift from l o w e r v a l u e s in the l o w e r parts of

Life in the Cincinnatian Sea 231

 t h e s e c t i o n , toward h i g h e r v a l u e s m o v i n g u p - s e c t i o n . As noted in chapters 4
and 15, analyses of l i t h o l o g i c features s u c h as the shale-to-limestone ratio and
b e d d i n g t h i c k n e s s d e m o n s t r a t e s that water d e p t h d e c r e a s e d from the base to
t h e top o f the f o r m a t i o n . The D C A s h o w e d that t h e c o m p o s i t i o n o f fossil
a s s e m b l a g e s also reflects this trend a n d may e v e n provide a m o r e sensitive
m e a s u r e o f d e p t h c h a n g e s t h a n the c h a r a c t e r o f the rock reveals.
In fossil a s s e m b l a g e s from the l o w e r K o p e ( d e e p e r water), the m o s t
a b u n d a n t fossils are the slender crinoids Ectenocrinus and Cincinnaticri-
nus, the small, thin-shelled b r a c h i o p o d Sowerhyella, a n d t h e trilobites
Cryptolithus a n d Acidaspis (see F i g u r e s 11.5,11.6). Higher in the K o p e , the
larger b r a c h i o p o d Dalmanella, branching bryozoans, and t h e trilobites
Flexicalymene a n d Isotelus b e c o m e t h e m o s t a b u n d a n t taxa. The brachiopods Zygospira a n
Dalmanella a s s e m b l a g e at h i g h e r levels. At the lop of the K o p e , the larger
concavo-convex brachiopods Rafinesquina a n d Strophomena are t h e m o s t
abundant brachiopods, a n d t h e larger c r i n o i d Glyptocrinus r e p l a c e s t h e
s m a l l e r c r i n o i d s . I n a n earlier s t u d y o f t h e K o p e t o F a i r v i e w t o B e l l e v u e
Formations, D i e k m e y e r (1998) f o u n d s i m i l a r t r a n s i t i o n s f r o m taxa o f
s m a l l e r , m o r e d e l i c a t e a n i m a l s i n t h e K o p e t o larger, m o r e t h i c k - s h e l l e d
a n d r o b u s t a n i m a l s (Platystrophia, m o r e m a s s i v e bryozoans) in the overly-
i n g F a i r v i e w . S h e i n t e r p r e t e d this to be a result of a c o n t i n u a t i o n of the
s h a l l o w i n g initiated d u r i n g t h e d e p o s i t i o n o f t h e K o p e s e q u e n c e .

R e c e n t research by H o l l a n d a n d P a t z k o w s k y (2007) provided similar fau-

nal analyses for e a c h o f the C i n c i n n a t i a n d e p o s i t i o n a l s e q u e n c e s , C 1 - C 6 . I n
F i g u r e 16.3 we present a t i m e e n v i r o n m e n t d i a g r a m for the entire C i n c i n n a -
tian based on the work by Holland and Patzkowsky. In this d i a g r a m we show
the major taxa of depth-related a s s e m b l a g e s in e a c h s e q u e n c e . T h e transition
from taxa of smaller, m o r e delicate a n i m a l s in d e e p e r parts of a s e q u e n c e to
taxa of larger, m o r e robust o r g a n i s m s in the s h a l l o w e r parts that was found in
the C1 s e q u e n c e by Holland, Miller, Meyer, and Dattilo (2001) was c o n f i r m e d
a n d f o u n d to be repeated in the s u c c e e d i n g C2 and C3 s e q u e n c e s . In other
w o r d s , in e a c h of these s e q u e n c e s the fossil a s s e m b l a g e s reflect a depth gradi-
ent from d e e p e r to s h a l l o w e r water. they also identified a s e c o n d e n v i r o n -
m e n t a l g r a d i e n t c o r r e s p o n d i n g to the n a t u r e of the substratum. A n i m a l s of
taxa a l o n g o n e e n d of this g r a d i e n t c h a r a c t e r i z e soft substrata and tend to be
smaller and thinner-shelled, w h e r e a s a n i m a l s of taxa arrayed toward the other
e n d of the g r a d i e n t c h a r a c t e r i z e firm substrata, tend to be larger and m o r e
robust, and c o m m o n l y are attached or e n c r u s t i n g in g r o w t h habit.

The R i c h m o n d i a n Invasion

W i t h i n t h e u p p e r d i v i s i o n o f t h e C i n c i n n a t i a n , the R i c h m o n d i a n S t a g e ,
the stable p a t t e r n s of d i s t r i b u t i o n of fossil a s s e m b l a g e s f o u n d in the lower
C i n c i n n a t i a n a r e d i s r u p t e d a n d r e o r g a n i z e d b y t h e influx o f o r g a n i s m s o f
many new taxa, i n c l u d i n g s p e c i e s , g e n e r a , a n d classes. This influx is t e r m e d
the R i c h m o n d i a n Invasion. T h e s e i n v a d e r s did not r e p l a c e p r e - e x i s t i n g
taxa b u t instead i n c r e a s e d C i n c i n n a t i a n diversity to its h i g h e s t level. There
i s a n initial p h a s e o f t h e R i c h m o n d i a n Invasion w i t h i n t h e C 4 s e q u e n c e ,

232 A Sea without Fish

hut the influx c u l m i n a t e s w i t h i n t h e C 5 s e q u e n c e w h e r e fossils o f o v e r f i f t y The invasion was not
new g e n e r a o f c o r a l s , brachiopods, b r y o z o a n s , m o l l u s c s , trilobites, and limited to particular fa-

e c h i n o d e r m s a p p e a r ( H o l l a n d 1997; H o l l a n d a n d P a t z k o w s k y 2007; f i g u r e cies, trophic groups,

or life-habit groups;
16.3; see c h a p t e r s S and 9). many of the new taxa w e r e n o t p r e s e n t d u r i n g
rather the Riehmondian
the Edenian and Maysvillian Stages of the C i n c i n n a t i a n , a l t h o u g h s o m e
Invasion was a major
n e w c o m e r s represent s p e c i a t i o n w i t h i n l o n g - r a n g i n g C i n c i n n a t i a n taxa
ecological revolution
s u c h as the brachiopods Platystrophia a n d Strophomena (Holland 1997).
affecting all aspects of
N e w taxa a p p e a r e d i n all d e p o s i t i o n a l e n v i r o n m e n t s across the s p e c t r u m the Cincinnatian seas.
of the C i n c i n n a t i a n depth gradient, and the a n i m a l s o c c u p y the entire
Steven M. Holland
r a n g e o f f e e d i n g t y p e s a n d life habits ( H o l l a n d 1997).
1997, 320
In the C4 s e q u e n c e , s o m e e l e m e n t s of the older, depth-related assem-
b l a g e s , s u c h as Hebertella a n d Platystrophia, are p r e s e n t in the shallow
subtidal z o n e , a n d Rafinesquina and Zygospira in t h e d e e p e r s u b t i d a l z o n e
( f i g u r e 16.3). In t h e C5 s e q u e n c e , a Dalmanella b r a c h i o p o d a s s e m b l a g e is
a g a i n p r e s e n t in the offshore e n v i r o n m e n t s , h u t s p e c i m e n s of Zygospira
also are present. S p e c i m e n s of Rafinesquina, Platystrophia, a n d b r a n c h i n g
b r y o z o a n s o c c u p y the d e e p e r s u b t i d a l z o n e , b u t t h e Hebertella-Platystro-
phia a s s e m b l a g e i s g o n e f r o m t h e s h a l l o w s u b t i d a l z o n e , w h e r e a n a s s e m -
b l a g e of c o l o n i a l c o r a l s a p p e a r s for the first t i m e . The d e p t h g r a d i e n t is
re-established in t h e C5 s e q u e n c e , but it is m u c h m o r e c r o w d e d w i t h taxa
and has a different c h a r a c t e r t h a n in o l d e r s e q u e n c e s ( H o l l a n d a n d P a t z -
k o w s k y 2007). T h e C 6 s e q u e n c e i s r e p r e s e n t e d o n l y b y s h a l l o w w a t e r a s -
s e m b l a g e s i n c l u d i n g c o r a l s a n d robust b r a c h i o p o d s a n d b r y o z o a n s .
T h e r e has b e e n considerable debate a b o u t the causes of the R i e h m o n d i a n
Invasion. B e c a u s e m a i n o f the invasive taxa o c c u r i n the M i d d l e a n d U p p e r
O r d o v i c i a n of the western United States a n d C a n a d a as far north as the pres-
ent-day A r c t i c , it is most likely that the invasion originated from the west and
northwest, tropical, w a r m water latitudes d u r i n g the L a t e O r d o v i c i a n (see
chapter 15). T h e invasion was a large-scale i m m i g r a t i o n e v e n t rather t h a n an
evolutionary hurst w i t h i n the C i n c i n n a t i A r c h region itself ( H o l l a n d 1997).
R e t u r n i n g to the theater analogy, we find that many players (species)
i n the R i e h m o n d i a n f i n a l a c t o f the C i n c i n n a t i a n a p p e a r e d i n earlier acts
r e c o r d e d b y p r e - C i n c i n n a t i a n f o r m a t i o n s o f the A p p a l a c h i a n r e g i o n . T h e s e
include brachiopods like Leptaena, Glyptorthis, a n d Plaesiomys ( H o l l a n d
1997), as w e l l as s t r o m a t o p o r o i d s a n d c o r a l s l i k e Tetradium. L i k e w i s e t h e
s c e n e d u r i n g p r e - C i n c i n n a t i a n t i m e featured w i d e s p r e a d l i m e s t o n e d e p o s i -
tion in K e n t u c k y s u g g e s t i n g a w a r m e r w a t e r e n v i r o n m e n t c o n d u c i v e to
carbonate deposition.
A s the E d e n i a n act o f the C i n c i n n a t i a n play o p e n e d , t h e s c e n e c h a n g e d
with the influx of m u d s derived from t e c t o n i c activity offstage in t h e T a c o n i c
region of the A p p a l a c h i a n o r o g e n i c belt, a n d c o o l e r waters s w e p t in as c i r c u -
lation patterns c h a n g e d ( H o l l a n d 1997). Many players m a d e their exits, to
return o n l y w h e n the s c e n e a g a i n altered i n R i e h m o n d i a n t i m e

F u r t h e r u n d e r s t a n d i n g of the Cincinnatian e v o l u t i o n a r y play also d e p e n d s The Cincinnatian

o n the interplay b e t w e e n c h a r a c t e r s . H o w t h e m e m b e r s o f t h e cast inter- Ecosystems: A Sea
acted with o n e a n o t h e r w a s o f f u n d a m e n t a l i m p o r t a n c e i n d e t e r m i n i n g t h e
w i t h o u t Fish!

Life in the Cincinnatian Sea 233

 plot a n d o u t c o m e o f t h e play. I n t e r a c t i o n s b e t w e e n o r g a n i s m s are o f m a j o r
i m p o r t a n c e i n e c o l o g y a n d lead t o t h e c o n c e p t o f a n e c o s y s t e m .
For a n e c o l o g i s t , a n e c o s y s t e m e n c o m p a s s e s all t h e c h e m i c a l , physical,
a n d b i o l o g i c a l a s p e c t s o f t h e e n v i r o n m e n t , i n c l u d i n g the s o u r c e s o f e n e r g y
a n d n u t r i e n t s e n t e r i n g t h e e n v i r o n m e n t a n d the w a y living o r g a n i s m s u s e
this e n e r g y to survive a n d r e p r o d u c e . The S u n is the p r i m a r y e n e r g y s o u r c e
for t h e vast majority o f e c o s y s t e m s o n E a r t h , t h e o n l y k n o w n e x c e p t i o n s b e -
i n g the r e c e n t l y d i s c o v e r e d d e e p sea vents, w h e r e h y d r o t h e r m a l fluids rich
in n u t r i e n t s sustain m i c r o b i a l life that is, in t u r n , the basis for u n i q u e e c o -
systems. In shallow seas like that of the C i n c i n n a t i a n , we c a n a s s u m e that
p l a n k t o n i c a s w e l l a s b e n t h i c a l g a e h a r n e s s e d solar e n e r g y a s t h e p r i m a r y
p r o d u c e r s . I n d i v i d u a l s o f all o f the a n i m a l g r o u p s c o n s t i t u t e d c o n s u m e r s ,
f e e d i n g either directly on t h e p r i m a r y p r o d u c e r s or on other c o n s u m e r s . In
order t o u n d e r s t a n d t h e n a t u r e o f C i n c i n n a t i a n e c o s y s t e m s , w e must under-
stand how t h e c o n s u m e r s w e r e interrelated in w h a t ecologists call a f o o d
c h a i n or, m o r e realistically, a f o o d w e b . H o w did O r d o v i c i a n m a r i n e ecosys-
t e m s c o m p a r e t o t h o s e o f t h e present-day s h a l l o w sea? D i d the n a t u r e o f the
interrelationships a m o n g o r g a n i s m s a n d the form of the food w e b play a role
i n d e t e r m i n i n g t h e diversity a n d a b u n d a n c e o f o r g a n i s m s i n the C i n c i n n a -
tian sea? In o u r a n a l o g y , did t h e interplay a m o n g c h a r a c t e r s a c t u a l l y deter-
m i n e the cast o n stage d u r i n g a n y p a r t i c u l a r act?
For a n e c o l o g i s t , i n t e r p r e t a t i o n o f a food w e b r e q u i r e s d e t a i l e d k n o w l -
e d g e o f t h e f e e d i n g habits a n d diets o f o r g a n i s m s o f s p e c i e s l i v i n g t o g e t h e r
a t a g i v e n t i m e . A n e c o l o g i s t c a n d i r e c t l y o b s e r v e predator-prey interactions
a n d c a n s a m p l e g u t c o n t e n t s o f a n i m a l s a n d their d r o p p i n g s . A n e c o l o g i s t
c a n g a u g e t h e flow o f e n e r g y t h r o u g h a n e c o s y s t e m b y m e a s u r i n g t h e c a -
loric c o n t e n t of organisms at different levels in the food c h a i n . However, a
p a l e o e c o l o g i s t w o r k i n g w i t h fossils p r e s e r v e d i n r o c k c a n n o t o b s e r v e o r
s a m p l e t h e l i v i n g e c o s y s t e m d i r e c t l y a n d thus faces s o m e severe l i m i t a t i o n s
in r e c o n s t r u c t i n g a " f o s s i l " e c o s y s t e m . It often is hard to d e t e r m i n e w i t h
certainty w h e t h e r all fossils p r e s e r v e d in a s i n g l e b e d w e r e alive at the s a m e
t i m e , b e c a u s e m a n y fossil a s s e m b l a g e s are t i m e - a v e r a g e d a m a l g a m a t i o n s
of organisms of many generations. Moreover, organic remains can be
m o v e d into a n a r e a o r o u t o f a n a r e a , for e x a m p l e , b y c u r r e n t s . D e s p i t e
t h e s e l i m i t a t i o n s , it is rather s u r p r i s i n g to find h o w m u c h i n f o r m a t i o n the
fossil r e c o r d o f t h e C i n c i n n a t i a n sea d o e s p r o v i d e , i n f o r m a t i o n that c a n b e
used to answer many questions about the nature of the ecosystems.
T h e m o s t d i r e c t e v i d e n c e for i n t e r a c t i o n s b e t w e e n o r g a n i s m s o f C i n -
cinnatian species c o m e s from a compilation of close associations a m o n g
d i f f e r e n t t y p e s o f o r g a n i s m s . A s s o c i a t i o n s are essential b e c a u s e the\ c a n
p r o v i d e e v i d e n c e for predator-prey r e l a t i o n s h i p s as w e l l as e v i d e n c e that
t w o d i f f e r e n t s p e c i e s lived a t t h e s a m e t i m e a n d possibly a f f e c t e d o n e a n -
other in various ways. Besides predator-prey relationships, other c o m m o n
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 i n c l u d e host-parasite a s s o c i a t i o n s , c o m p e t i t i v e
i n t e r a c t i o n s , m u t u a l i s m s , or c o m m e n s a l i s m s . In competitive interactions,
i n d i v i d u a l s o f b o t h a s s o c i a t e d s p e c i e s are i n s o m e w a y i n h i b i t e d o r h a r m e d
by t h e a s s o c i a t i o n ; in a m u t u a l i s t i c i n t e r a c t i o n , b o t h i n d i v i d u a l s derive
s o m e benefit from the association; in a c o m m e n s a l i s t i c interaction, one

234 A Sea without Fish

s p e c i e s derives s o m e b e n e f i t , w h e r e a s t h e " h o s t " i s u n a f f e c t e d b y t h e pres-
e n c e of the symbiont. In s o m e cases a l i v i n g a n i m a l o f o n e s p e c i e s m i g h t
b e associated w i t h n o n - l i v i n g r e m a i n s o f a n o t h e r s p e c i e s , a s i n t h e c a s e o f
h e r m i t crabs o c c u p y i n g shells o f d e a d snails ( D a v i s , M a p e s , a n d K l o f a k
1999; D a v i s , F r a a y e , a n d H o l l a n d 2001).
W e c o m p i l e d a v a i l a b l e i n f o r m a t i o n o n a s s o c i a t i o n s o f fossils o f C i n c i n -
n a t i a n s p e c i e s into t w o s u m m a r y tables. Table 2 s h o w s p o t e n t i a l predator-
prey a s s o c i a t i o n s d e r i v e d from d i r e c t fossil e v i d e n c e , a n d Table 3 s h o w s all
other associations reported a m o n g individuals of C i n c i n n a t i a n species or
other groups.

Predator-Prey Interactions

The list of potential predator-prey associations in t h e C i n c i n n a t i a n is quite

short (Table 2), and the n a t u r e of the e v i d e n c e is variable. T h r o u g h o u t the
fossil record there are rare but n o t a b l e eases of fossilized s t o m a c h c o n t e n t s
that are the strongest e v i d e n c e for the diet of an e x t i n c t a n i m a l . The only
possible instance of this in the C i n c i n n a t i a n is the o c c u r r e n c e of o s t r a c o d e s
preserved w i t h i n the coralla of the r u g o s e corals Grewingkia and Streptel-
asma (Elias 1984). The o s t r a c o d e s are m o s t l y articulated a n d located n e a r or
w i t h i n the alar and c a r d i n a l fossulae o f t h e coral's c a l i c e . T h e fossulae are
e x p a n d e d regions b e t w e e n the septa that probably f u n c t i o n e d in water c i r c u -
lation w i t h i n the p o l y p a n d thus w e r e related to i n g e s t i o n of food and/or
ejection of waste. A l t h o u g h it is possible that the coral ingested t h e o s t r a c o d e s
as prey, Elias favored an alternative h y p o t h e s i s that the o s t r a c o d e s entered the
c a l i c e w h e n the p o l y p b e c a m e d e t a c h e d from t h e side o f t h e c a l i c e and
e v e n t u a l l y b e c a m e trapped b e n e a t h n e w l y secreted t a b u l a e . In this s c e n a r i o ,
the ostracodes m i g h t have lived s y m b i o t i c a l l y w i t h i n the coral c a l i c e .
The most direct e v i d e n c e of a predator-prey relationship is the remarkable
s p e c i m e n of a sea star Promopalaeaster preserved with its a r m s w r a p p e d a r o u n d
a s p e c i m e n of p e l e c y p o d tentatively assigned to g e n u s Cumeamya (see F i g u r e
12.15E; Blake and C u e n s b u r g 1994). This s p e c i m e n is probably o n e of the most
extraordinary fossils ever to be found in the C i n c i n n a t i a n . It provides strong
e v i d e n c e that s o m e C i n c i n n a t i a n sea stars had a c q u i r e d the ability to o p e n
p e l e c y p o d s seen in living asteriid sea stars, e v e n t h o u g h the present-day forms
are not direct d e s c e n d a n t s of Promopalaeaster (Blake a n d C u e n s b u r g 1994).
T h e predatory b e h a v i o r o f s o m e i n d i v i d u a l s o f C i n c i n n a t i a n trilobite
s p e c i e s was d i s c u s s e d in c h a p t e r 11. Rusophycus t r a c e fossils that intersect
b u r r o w s s u g g e s t that i n d i v i d u a l s o f t h e trilobite Isotelus preyed on some
b u r r o w i n g , p r o b a b l y s o f t - b o d i e d o r g a n i s m s . B a b c o c k (2003) i n f e r r e d that
both eurypterids and cephalopods w e r e p o t e n t i a l p r e d a t o r s o n trilobites
d u r i n g the O r d o v i c i a n , a l t h o u g h t h e r e i s n o d i r e c t e v i d e n c e a v a i l a b l e .
Specimens of C i n c i n n a t i a n brachiopods such as Rafinesquina and
several o t h e r g e n e r a p r o v i d e e v i d e n c e for p r e d a t i o n i n t h e f o r m o f c h a r a c -
teristic b r e a k a g e a n d shell repair ( A l e x a n d e r 1986). A l e x a n d e r c o n s i d e r e d
possible predators a m o n g sea stars, e u r y p t e r i d s , g a s t r o p o d s , a n d n a u t i l o i d
c e p h a l o p o d s . H e c o n c l u d e d that t h e b e a k s o f n a u t i l o i d s w e r e m o s t likely
t o h a v e inflicted the t y p e o f d a m a g e f o u n d i n t h e b r a c h i o p o d s .

Life in the Cincinnatian Sea 235

 C i n c i n n a t i a n crinoids also show e v i d e n c e o f d a m a g e and regeneration
of the c a l y x , a r m s , a n d c o l u m n that is very likely the result of predation
(Ausich a n d B a u m i l l e r 1993; D o n o v a n and S c h m i d t 2001; B a u m i l l e r and
Gahm 2004). However there is no e v i d e n c e as to the specific predator respon-
sible. W e s p e c u l a t e that n a u t i l o i d s m i g h t b e the m o s t likely culprits, i n v i e w
o f their a b u n d a n c e a n d potential b e h a v i o r .

O t h e r Interspecific Interactions

Table 3 s h o w s that v i r t u a l l y all the m a j o r i n v e r t e b r a t e g r o u p s f o u n d in the

C i n c i n n a t i a n h a v e r e c o r d e d a s s o c i a t i o n s o f i n d i v i d u a l s o f taxa b e l o n g i n g
t o a w i d e v a r i e t y o f g r o u p s . T h e t y p e o r n a t u r e o f the a s s o c i a t i o n s r a n g e s
f r o m t h e u s e of e i t h e r a l i v i n g h o s t or d e a d r e m a i n s as a s u b s t r a t u m for
e n c r u s t a t i o n , b o r i n g , o r h a b i t a t i o n ( F i g u r e 16.1), t o possible c o m m e n s a l i s m
or p a r a s i t i s m . To t h e e x t e n t that a host w a s , by d e f i n i t i o n , l i v i n g at the t i m e
o f t h e a s s o c i a t i o n , i t a p p e a r s that i n t e r a c t i o n s b e t w e e n i n d i v i d u a l s o f C i n -
cinnatian species were very c o m m o n . Detailed discussion of the nature of
t h e a s s o c i a t i o n s listed c a n b e f o u n d i n t h e c h a p t e r s c o n c e r n i n g t h e t a x o -
n o m i c g r o u p o f e a c h host.
We p r e s e n t Table 3 w i t h a s i g n i f i c a n t c a v e a t . In c a s e s in w h i c h a fossil
s p e c i m e n includes an organism of o n e species attached to an individual of
a n o t h e r s p e c i e s , it is n o t a l w a y s c l e a r w h e t h e r b o t h a n i m a l s w e r e alive at
t h e t i m e of a t t a c h m e n t . It is p o s s i b l e that the a t t a c h e d o r g a n i s m fastened
o n t o t h e o t h e r after t h e hitter's d e a t h a n d , for e x a m p l e , was u s i n g the e m p t y
shell as a hard s u b s t r a t u m . An a d d e d c o m p l i c a t i o n is that v a r i o u s biologists
a n d p a l e o n t o l o g i s t s h a v e n o t b e e n c o n s i s t e n t a s t o w h i c h t e r m s are u s e d for
w h i c h associations. Take t h e t e r m " e p i z o a , " for e x a m p l e . For a n association
p r o p e r l y t o b e t e r m e d " e p i z o i s m , " b o t h p a r t i e s m u s t h a v e b e e n alive a t t h e
t i m e o f t h e a s s o c i a t i o n b o t h host a n d g u e s t . H o w e v e r , the term " e p i z o a "
n o t u n c o m m o n l y has b e e n u s e d i n c a s e s i n w h i c h the " h o s t " clearly was
d e a d at t h e t i m e of a t t a c h m e n t a n d in c a s e s in w h i c h it is u n c l e a r w h e t h e r
t h e " h o s t " w a s alive o r d e a d a t t h e t i m e o f t h e a s s o c i a t i o n . ( D a v i s , M a p e s ,
a n d K l o f a k 1999; D a v i s , F r a a y e , a n d H o l l a n d 2001.)
M o s t of t h e c a s e s cited in Table 3 h a v e b e e n g a t h e r e d from p u b l i s h e d
reports. The a u t h o r s of those reports h a v e not a l w a y s b e e n able to d e t e r m i n e
or state w h e t h e r b o t h " h o s t " a n d " g u e s t " w e r e alive at the t i m e of a given
association. In cases w h e r e the t y p e of association is r e c o r d e d in Table 3 as
e p i z o i c , e n d o z o i c , o r o n e i n v o l v i n g b o r i n g , the putative host o r g a n i s m m a y
or may n o t h a v e b e e n l i v i n g at t h e t i m e of the a s s o c i a t i o n ; the entry in the
table d e p e n d s on the i n f o r m a t i o n a n d interpretations presented in the origi-
nal p u b l i c a t i o n s . O n the o t h e r h a n d , i n c a s e s entered a s c o m m e n s a l o r para-
sitic, there is e v i d e n c e that t h e host w a s , in fact, l i v i n g at the t i m e of the as-
s o c i a t i o n . B i o i m m u r a t i o n is also i n d i c a t i v e of interaction b e t w e e n a living
host a n d o r g a n i s m s o f a n associated s p e c i e s , b e c a u s e the associate m o d i f i e d
the g r o w t h of the host in s o m e w a y as to leave e v i d e n c e that the associate was
present. T h e r e are n o k n o w n d o c u m e n t e d c a s e s i n the C i n c i n n a t i a n o f ei-
ther c o m p e t i t i v e or m u t u a l i s t i c interactions, L i d d e l l and Brett (19S2) reported

236 A Sea without Fish

e v i d e n c e for c o m p e t i t i o n b e t w e e n associated s p e c i e s o f e n c r u s t i n g b r y o z o -
ans from the M i d d l e S i l u r i a n o f I n d i a n a .
C o l o n i e s o f different s p e c i e s o f b r y o z o a n s e n c r u s t i n g c r i n o i d c a l y c e s
s o m e t i m e s f o r m e d raised m a r g i n s w h e r e t h e y g r e w into c o n t a c t w h e n e n -
crusting crinoid calyces. The raised m a r g i n s i n d i c a t e s o m e k i n d o f "stand-
off" i n w h i c h c o l o n i e s o f e a c h s p e c i e s w e r e u n a b l e t o overgrow t h e o t h e r a n d
both w e r e thus inhibited. B r y o z o a n s e n c r u s t i n g b r a c h i o p o d s i n the C i n c i n -
natian m i g h t also b e e x p e c t e d t o p r o v i d e this k i n d o f e v i d e n c e a l t h o u g h n o
i n s t a n c e s h a v e yet b e e n reported. E v i d e n c e for m u t u a l b e n e f i t to t w o associ-
ated species is e v e n m o r e difficult to establish from fossil material.

Cincinnatian Guilds

As a m e a n s of characterizing the complexity of an ecosystem, ecologists

d e v e l o p e d the c o n c e p t of a g u i l d . As a p p l i e d to n a t u r a l e c o s y s t e m s , a g u i l d
is d e f i n e d as "a g r o u p of s p e c i e s that exploit the s a m e class of e n v i r o n m e n t a l
resources in a similar way T h i s term g r o u p s t o g e t h e r s p e c i e s , w i t h o u t regard
t o t a x o n o m i c p o s i t i o n , that o v e r l a p s i g n i f i c a n t l y i n t h e i r n i c h e r e q u i r e -
m e n t s " ( R o o t 1967, 335). T h e p a l e o e c o l o g i s t R i c h a r d B a m b a c h a p p l i e d t h e
g u i l d c o n c e p t to fossil c o m m u n i t i e s a n d e c o s y s t e m s in order to e x p l a i n h o w
the e c o l o g i c a l s t r u c t u r e a n d c o m p l e x i t y o f e c o s y s t e m s has c h a n g e d o v e r
e v o l u t i o n a r y t i m e ( B a m b a c h 1 9 8 3 , 1 9 9 3 ; B u s h a n d B a m b a c h 2004).
B a m b a c h (19S3) first c o m p a r e d the major a d a p t i v e strategies of o r g a n -
isms c o m p r i s i n g the three m a r i n e " E v o l u t i o n a r y f a u n a s " : C a m b r i a n , P a l e o -
zoic, and M e s o z o i c - C e n o z o i c (Sepkoski 1981). B a m b a c h classified adaptive
strategies a c c o r d i n g t o the m o d e o f s p a c e u t i l i z a t i o n a n d f e e d i n g habit. S u b -
s e q u e n t l y D r o s e r and S h e e h a n (1997) c a l l e d these broad c a t e g o r i e s " m e g a -
g u i l d s " and assigned b e n t h i c taxa to m e g a g u i l d s for t h e entire O r d o v i c i a n .
they found that the patterns established for e p i f a u n a a n d i n f a u n a d u r i n g the
O r d o v i c i a n r e m a i n e d stable for the rest o f the P a l e o z o i c Era. W e c o n s t r u c t e d
m e g a g u i l d d i a g r a m s for major g r o u p s f o u n d i n the C i n c i n n a t i a n , m o d i f i e d
s o m e w h a t from those o f D r o s e r a n d S h e e h a n a n d i n c l u d i n g p e l a g i c g r o u p s
(Table 4a). M o s t of the g r o u p s are at t h e class level, b u t s o m e are orders. It
should be r e c o g n i z e d that this c o m p i l a t i o n is c u m u l a t i v e for the entire C i n -
c i n n a t i a n ; thus the total n u m b e r o f g u i l d s e x c e e d s t h e total that w o u l d b e
e x p e c t e d in a single c o n t e m p o r a n e o u s a s s e m b l a g e .
C o m p a r i s o n o f m e g a g u i l d s t r u c t u r e o f t h e C i n c i n n a t i a n w i t h that o f t h e
M e s o z o i c - C e n o z o i c reveals major contrasts ( T a b l e 4b). A m o n g the e p i f a u n a ,
the M e s o z o i c - C e n o z o i c i n c l u d e s m a n y s u s p e n s i o n f e e d i n g taxa a s d o e s the
C i n c i n n a t i a n , but present-day g r o u p s have r e p l a c e d m a n y o f those that w e r e
present d u r i n g the C i n c i n n a t i a n . P e l e c y p o d s b e c a m e d o m i n a n t , w h e r e a s
brachiopods a s s u m e d a m i n o r role. The b r y o z o a n s d o m i n a n t in the P a l e o -
zoic, the t r e p o s t o m e s , b e c a m e e x t i n c t a n d t h e c h e i l o s t o m e s b e c a m e d o m i -
nant. Stalked echinoderms (crinoids, blastoids, a n d cystoids) suffered major
e x t i n c t i o n s at t h e e n d of the P a l e o z o i c , a n d stalked c r i n o i d s s u r v i v e d o n l y in
d e e p water. The n u m b e r o f taxa o f m o b i l e e p i f a u n a l a n i m a l s , s u c h a s gastro-
p o d s , c r u s t a c e a n s , a n d sea u r c h i n s i n c r e a s e d m a r k e d l y .
Exploitation of living space and food resources within the s e d i m e n t by

Life in the Cincinnatian Sea 237

Figure 16.2. Reconstruc- i n f a u n a l o r g a n i s m s s t a n d s o u t a s o n e o f the m a j o r c o n t r a s t s b e t w e e n the
tion of life on the Cincin- O r d o v i c i a n g u i l d s t r u c t u r e a n d that o f the p o s t - P a l e o z o i c . The diversifica-
natian sea floor. Drawing
tion o f b u r r o w i n g p e l e c y p o d s , g a s t r o p o d s , p o l y c h a e t e s , c r u s t a c e a n s , a n d
by Anneliese Caster and
e c h i n o i d s p o p u l a t e d b l o c k s o f t h e i n f a u n a l m e g a g u i l d s t r u c t u r e that w e r e
Elizabeth A. Dalve. From
relatively e m p t y d u r i n g t h e P a l e o z o i c ( T a b l e 4a).
von Engeln and Caster,
Geology, 1952, figure P e r h a p s t h e m o s t s t r i k i n g c o n t r a s t s b e t w e e n t h e O r d o v i c i a n and post-

347, McGraw-Hill, and P a l e o z o i c w e r e i n t h e p e l a g i c r e a l m , t h e o p e n w a t e r a b o v e t h e sea floor.

reprinted by permission D u r i n g C i n c i n n a t i a n t i m e few g r o u p s o c c u p i e d the w a t e r c o l u m n , and
of the McGraw-Hill a n i m a l s o f t h o s e g r o u p s differed g r e a t l v from t h o s e o f the p o s t - P a l e o z o i c
Companies. ( T a b l e 4b). It is a s t o u n d i n g to r e a l i z e that d u r i n g the C i n c i n n a t i a n there
really w a s , i n t h e r e g i o n o f C i n c i n n a t i , a sea w i t h o u t f i s h , w h e r e n a u t i l o i d
c e p h a l o p o d s a n d s o m e trilobites a n d e u r y p t e r i d s w e r e t h e o n l y large, ac-
tively s w i m m i n g o r g a n i s m s (Plates 13, 14; figure 16.2). O n l y later in t h e
P a l e o z o i c did fish b e g a n to proliferate, but m o d e r n b o n y fish did not diver-
sify a s h e r b i v o r e s a n d c a r n i v o r e s u n t i l t h e late M e s o z o i c a n d C e n o z o i c .
D u r i n g the C i n c i n n a t i a n a n d i n d e e d for m o s t o f t h e P a l e o z o i c , t h e shallow
m a r i n e e c o s y s t e m was vastly different from a n y t h i n g r e s e m b l i n g a present-
dav s e l l i n g . H o w c a n w e e x p l a i n t h e s e s t r i k i n g d i f f e r e n c e s ?

Was the Cincinnatian Sea a "Beggar's B a n q u e t " ?

Bambach (1993) a r g u e d that m a j o r c h a n g e s in t h e s t r u c t u r e of m a r i n e

e c o s y s t e m s , as seen in the fossil r e c o r d , are a r e f l e c t i o n of an i n c r e a s e in

238 A Sea without Fish

T H E p r i m a r y p r o d u c t i v i t y o f the world's o c e a n s d u r i n g t h e P h a n e r o z o i c
E o n . A s m e n t i o n e d earlier, w e a s s u m e that p r i m a r y p r o d u c t i o n i n t h e
C i n c i n n a t i a n sea was b a s e d o n m i c r o p h y t o p l a n k t o n a s w e l l a s s o m e b e n -
thic m a c r o a l g a e . It is a p a r a d o x that d e s p i t e t h e h i g h a b u n d a n c e of b e n t h i c
suspension feeding invertebrates in the C i n c i n n a t i a n fauna the only pre-
served m i c r o p h y t o p l a n k t o n i c o r g a n i s m s are t h e a c r i t a r c h s . W e r e a c r i t a r c h s
t h e o n l y s u s p e n d e d food s o u r c e for a b o t t o m f a u n a d o m i n a t e d by a b u n d a n t
and diverse s u s p e n s i o n feeders? W e c a n s p e c u l a t e that p e r h a p s o t h e r m i -
c r o p l a n k t o n i c a l g a e e x i s t e d o n e s that l a c k e d p r e s e r v a b l e o r g a n i c c e l l
w a l l s o r m i n e r a l i z e d tests. O r , p e r h a p s c l a y p a r t i c l e s s u s p e n d e d i n t h e water
c o l u m n served as substrata for b a c t e r i a that f o r m e d a " m a r i n e s n o w " that
n o u r i s h e d b e n t h i c s u s p e n s i o n feeders. B u t , s p e c u l a t i o n a s i d e , t h e fossil
record o f m a r i n e p l a n k t o n d e f i n i t e l y i n d i c a t e s that t h e diversity o f taxa o f
planktonic organisms in the C i n c i n n a t i region d u r i n g the C i n c i n n a t i a n
was a b o u t fifty s p e c i e s ( C o l b a t h 1979), 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
t h r e e - t o four h u n d r e d s p e c i e s o f a e r i t a r c h s , t h e n suffered a d e c l i n e d u r i n g
the late P a l e o z o i c a n d early M e s o z o i c . It t h e n b e g a n to i n c r e a s e d u r i n g t h e
Jurassic a n d C r e t a c e o u s , w i t h t h e a p p e a r a n c e o f p r e s e n t - d a y g r o u p s s u c h
as dinoflagellates, calcareous n a n n o p l a n k t o n , and diatoms (Tappan and
L o e b l i c h 1973). If t h e diversity of p l a n k t o n i c o r g a n i s m s is c o r r e l a t e d in
s o m e w a y w i t h a b u n d a n c e a n d p r o d u c t i v i t y , t h e n t h e fossil r e c o r d o f m a -
rine p l a n k t o n indicates a m a r k e d increase in productivity d u r i n g the
P h a n e r o z o i c ( B a m h a e h 1993). The C i n c i n n a t i a n s e a , d e s p i t e its s e e m i n g
organic a b u n d a n c e , may, in fact, have b e e n poor in food resources c o m -
pared to present-day s e t t i n g s .
A l t h o u g h the earliest spores of land plants date from the O r d o v i c i a n ,
terrestrial regions of the Earth r e m a i n e d essentially d e v o i d of w i d e s p r e a d
plant cover until the late D e v o n i a n ( B a m h a e h 1993). B a m h a e h s u g g e s t e d that
the rise of land plants had a p r o f o u n d effect on m a r i n e e c o s y s t e m s , b e c a u s e
rivers b e g a n to carry vast a m o u n t s of o r g a n i c matter a n d nutrients into the
sea. T h i s o r g a n i c matter provided a major new food r e s o u r c e for b e n t h i c
m a r i n e o r g a n i s m s and c o n t r i b u t e d to the great diversification of i n f a u n a l
guilds in the post-Paleozoic. ( T h e s e had b e e n poorly d e v e l o p e d in the C i n -
cinnatian.) Increased o r g a n i c input from the land also increased the supply
of nutrients to coastal m a r i n e e n v i r o n m e n t s , in turn e n h a n c i n g overall m a -
rine productivity. B a m h a e h a r g u e d that i n c r e a s i n g levels of m a r i n e productiv-
ity enabled an increase in m a r i n e biomass, diversity, and f u n c t i o n a l versatility
that is reflected in the fossil record of p o s t - P a l e o z o i c m a r i n e e c o s y s t e m s .
The vastly different, s e e m i n g l y a l i e n w o r l d o f t h e C i n c i n n a t i a n sea
c a n thus h e u n d e r s t o o d a s t h e p r o d u c t o f a p e r i o d i n E a r t h history w h e n
the c a p a c i t y of t h e sea to s u p p o r t lite was a m e r e f r a c t i o n of w h a t it b e c a m e
through subsequent Phanerozoic time. The m o d e s o f life, b o d y s i z e , a n d
overall a b u n d a n c e o f m a r i n e life i n t h e C i n c i n n a t i a n o f t h e C i n c i n n a t i
r e g i o n w e r e all l i m i t e d b y t h e a m o u n t a n d q u a l i t y o f food r e s o u r c e s avail-
able. The " b e g g a r ' s b a n q u e t " m a y h a v e d e f i n e d t h e r u n n i n g plot o f t h e
C i n c i n n a t i a n play.
T h e u l t i m a t e o u t c o m e o f this plot i s h i d d e n f r o m v i e w i n t h e C i n c i n -
nati region b e c a u s e the e v i d e n c e o f t h e c l o s i n g a c t s o f t h e O r d o v i c i a n r e -

Life in the Cincinnatian Sea 239

Figure 1 6 . 3 . Time-envi-
ronment diagram for the
Cincinnatian 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,
SWB = storm wave base.
Letter-codes indicate ma-
jor fossils characteristic of
assemblages found in
each sequence and envi-
ronment, defined as fos-
sils occurring in >20% of
samples from each set-
ting. Letter-codes as fol-
lows: corals: Gr = Gew-
ingkia, Sp =
Streptelasma, Te = Tet-
radium, Pr - Protaraea;
bryozoans: bf = bifoliate
forms, en = encrusting
forms, nr = thin ramose
forms, ra = ramose forms;
brachiopods: Da = Dal-
manella, He = Heber-
c o r d w a s o b l i t e r a t e d b y s u b s e q u e n t e r o s i o n . T h e m i s s i n g script, a s read
tella. Hi = Hiscobeccus,
Ht = Holtedahlina, Lp = e l s e w h e r e , tells u s that s w e e p i n g e n v i r o n m e n t a l c h a n g e s b r o u g h t a b o u t
Leptaena, PI = Plat- m a s s e x t i n c t i o n s o f m a n y O r d o v i c i a n players w o r l d w i d e . The causes of
ystrophia, Pp = Plat- these extinctions are debated. T h e y apparently were c o m p l e x and seem to
ystrophia ponderosa, Pt h a v e i n v o l v e d a g l o b a l ice a g e , a l o n g w i t h c h a n g e s i n sea level a n d o c e a n
= Plectorthis, Ra =
c h e m i s t r y ( H a l l a m a n d W i g n a l l 1997). T h e shallow seas that a g a i n c o v e r e d
Rafinesquina, Re = Ret-
t h e r e g i o n o f t h e C i n c i n n a t i A r c h d u r i n g t h e e n s u i n g S i l u r i a n act w e r e
rorsirostra, Rn = Rhyn-
p o p u l a t e d w i t h m a n y f a m i l i a r players a n d roles, yet t h e cast o f c h a r a c t e r s
chotrema, So = Sower-
byella, St = r e p r e s e n t s a c l e a r l y d i f f e r e n t t r o o p of players on a new stage setting.
W h e n w e s e a r c h t h e seas o f t h e p r e s e n t day for a n y t h i n g r e s e m b l i n g
Strophomena, Zy = Zy- the C i n c i n n a t i a n e c o s y s t e m s , a revival o f t h e C i n c i n n a t i a n play, w e are

240 A Sea without Fish

hard pressed t o f i n d m a i n g o o d a n a l o g u e s . O f c o u r s e , e x t i n c t i o n has s w e p t gospira; trilobites; ca =
a w a y v i r t u a l l y all t h e taxa o f t h e o r g a n i s m s that i n h a b i t e d t h e C i n c i n n a t i a n calymenid, Ct = Cryptoli-

sea, i n c l u d i n g e n t i r e m a j o r g r o u p s s u c h a s g r a p t o l i t e s , c o n u l a r i i d s , trilo- thus, Is = Isotelus; ostra-

c o d e s = os; g a s t r o p o d s =
bites, e u r y p t e r i d s , a n d e d r i o a s t e r o i d s . H o w e v e r , t o d a y w e c a n f i n d rather
ga; p e l e c y p o d s ; Am =
restricted r e g i o n s w h e r e t h e sea f l o o r i s c o v e r e d w i t h b r y o z o a n s , s u c h a s t h e
Ambonychia, by = inde-
s h e l f off S o u t h A u s t r a l i a ( B r a d s t o c k a n d C o r d o n 1983; H a g e m a n e t al.
terminate pelecypod, Ca
2000). N e w Z e a l a n d , isolated i n t h e s o u t h w e s t e r n P a c i f i c , h a r b o r s t h e = Carotidens, mo = modi-
greatest diversity o f l i v i n g b r a c h i o p o d s ; e l s e w h e r e t h e s e a n i m a l s are b u t omorphid pelecypod; cri-
m i n o r c o m p o n e n t s of shallow water c o m m u n i t i e s . C l o s e r to the area of the noids. Ci = Cincinnaticri-
C i n c i n n a t i a n , t h e S a n Juan Islands in t h e P a c i f i c N o r t h w e s t s u p p o r t a di- nus, Ec = Ectenocrinus,

verse a n d a b 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 a s Gl = Glyptocrinus, Xe =

Xenocrinus; graptolites =
e c h i n o d e r m s , b r y o z o a n s , b r a c h i o p o d s , solitary c o r a l s , a n d s p o n g e s . H o w -
gra. Italicized fossils are
ever, t h e s e relicts are a m e r e f r a c t i o n of a m o r e d i v e r s e f a u n a rich in m o l -
those invading region
l u s c s , c r u s t a c e a n s , a n d f i s h a l l g r o u p s that proliferated i n t h e p o s t - P a l e o -
during the Richmondian
z o i c . P r e s e n t - d a y t r o p i c a l reefs far s u r p a s s t h o s e o f t h e O r d o v i c i a n i n invasion. Information de-
diversity a n d a b u n d a n c e , a n d yet t h e r e are c e r t a i n reef-related habitats t h a t rived from Holland and
mirror, t o s o m e e x t e n t , t h e P a l e o z o i c . I n A u s t r a l i a ' s G r e a t B a r r i e r Reef, Patzkowsky (2007); see
d e e p e r , soft s e d i m e n t b o t t o m s l o c a t e d b e l o w t h e c o r a l - d o m i n a t e d shallow this publication for distri-
bution of additional
reefs h a v e a P a l e o z o i c a s p e c t , rich in a l g a e , s p o n g e s , solitary c o r a l s , b r y o -
fossils.
z o a n s , a n d c r i n o i d s ( M e s s i n g e t al. 2006). I n d i v i d u a l s o f t h e c h a m b e r e d
Nautilus, the o n l y l i v i n g d e s c e n d a n t s o f the n a u t i l o i d c e p h a l o p o d s o f t h e
P a l e o z o i c , also s u r v i v e at d e p t h s of 100 m b e l o w t h e tops of t h e reefs of t h e
Pacific. L i k e w i s e , the o n l y l i v i n g s t a l k e d c r i n o i d s are restricted t o d e p t h s
greater t h a n 100 i n ( M e y e r 1997). C o l l e c t i v e l y , t h e s e e x a m p l e s o f p r e s e n t -
day survivors f r o m t h e past a n d P a l e o z o i c - l i k e c o m m u n i t i e s s h a r e t h e c h a r -
acteristics o f e x i s t e n c e i n s o m e isolated o r restricted s e t t i n g s w h e r e d o m i -
n a n c e b y the m o r e typical shallow m a r i n e f a u n a is, t o s o m e d e g r e e , r e l a x e d .
I t i s o n l y u n d e r s u c h a n o m a l o u s c o n d i t i o n s that o r g a n i s m s o f m a n y a n c i e n t
g r o u p s c a n f l o u r i s h a n d p r o v i d e u s w i t h a f l e e t i n g g l i m p s e o f the a n c i e n t
O r d o v i c i a n sea that o n c e c o v e r e d t h e C i n c i n n a t i r e g i o n .

Table 2. Predators and

Predator Prey References
Prey among Cincinnatian
Rugose corals ostracodes? Elias 1 9 8 4 Taxa

Nautiloids brachiopods Alexander 1 9 8 6

trilobites Babcock 2003

Unspecified crinoids Ausich and Baumiller 1993; D o n o -

van and Schmidt 2001; Baumiller
and Gahn 2004

Trilobites worms Babcock 2003; Brandt et al. 1995

Eurypterids trilobites Babcock 2003

Asteroids pelecypods Blake and Guensburg 1 9 9 4

Life in the Cincinnatian Sea 241

Table 3. Associations
ARRANGED BY "HOST"
among Individuals of Cin-
cinnatian Taxa Host Associate Type of References /
Association [Annotations]

PROTISTA

foraminifers bryozoan bioimmuration; Wilson, Palmer,

epizoism and Taylor (1994)

PORIFERA

stromatoporoids pelecypods boring Pojeta and Palmer

(1976);
Wilson and
Palmer (1988)

CNIDARIA

hydrozoan bryozoan encrustation (post- Wilson, Palmer,

mortem); and Taylor (1994)
dwelling inside
empty shell

corals algae, fungi boring Elias and Lee

(1993)

corals epizoism Elias (1983)

bryozoans encrustation: Bassler(1953)
epizoism

bryozoans encrustation: Elias (1983)

epizoism

worms boring Elias (1983)

(at least s o m e [Trypanites in
during life of host) Grewingkia]

worms boring Elias (1986)

BRACHIOPODA

brachiopods algae boring Kobluk and Risk

(1977)

corals epizoism Elias (1983)

post-mortem Shrake(1989)
brachiopods
attachment

encrustation; Nicholson (1875)

bryozoans
epizoism

bioimmuration; epi- Wilson, Palmer,

bryozoans
zoism and Taylor (1994)

bryozoans "boring"; Anstey and Wil-

presumably post- son (1996)
mortem
bryozoans "boring" Pohowsky (1974,
1978)

bryozoans post-mortem Shrake(1989)

attachment

242 A Sea without Fish

 ARRANGED BY "HOST"

Host Associate Type of References /

Association [Annotations]

bryozoans encrustation; ? Anstey and Wil-

epizoism son (1996)
[Corynotrypa on
interior of bra-
chiopod shell]

bryozoans encrustation; ? Anstey and Wil-

post-mortem son (1996)
[Cuffeyella on
interior of bra-
chiopod shell]

bryozoans encrustation Ulrich (1879)

bryozoans encrustation Ulrich (1883)

brachiopods epizoism Alexander and

Scharpf (1990)

Cornulites ? commensalism Morris and Rollins

(1971)

crinoids post-mortem Shrake (1989)

attachment

edrioasteroids epizoism Richards (1972)

epizoism or Meyer (1990)

commensalism

gastropods borings Fenton and Fen-

ton (1931)

borings Bucher (1938)

Sphenothallus epizoism Neal and Hanni-

bal (2000)

stromatopor- epizoism Richards (1972);

oids Alexander and
Scharpf (1990)

ECTOPROCTA

bryozoans corals epizoism Elias (1983)

brachiopods epizoism Richards (1972)

brachiopods aegism; epizoism Shrake (1989)

bryozoans encrustation; Nicholson (1875)

epizoism

bryozoans encrustation Ulrich (1879a)

bryozoans encrustation Ulrich (1883)

bryozoans aegism; epizoism Shrake (1989)

bryozoans bioimmuration; Wilson, Palmer,

epizoism and Taylor (1994)

Catellocaula bioclaustration; Palmer and Wil-

parasitism; son (1988)
epizoism

Life in the Cincinnatian Sea 243

 A R R A N G E D BY "HOST

Host Associate Type of References /

Association [Annotations]

Cornulites epizoism; ? Morris and Rollins

commensalism (1971)

cornulitids encrustation and Baird, Brett, and

intergrowth Frey(1989)

pelecypods boring Wilson and

Palmer (1988)

Sanctum boring Erickson and

Bouchard (2003)

Sphenothallus epizoism Bodenbender et

al. (1989)

trilobites aegism Shrake(1989)

"worms" borings Palmer and Wil-

Trypanites son (1988)

CORNUUTIDS

cornulitids bryozoans encrustation and Baird, Brett, and

intergrowth Frey (1989)

cornulitids bryozoans bioimmuration; Wilson, Palmer,

epizoism and Taylor (1994)

MOLLUSCA

Mollusca brachiopods epizoism Morris and Felton

gastropods (1993)

Mollusca Cornulites encrustation; S. A. Miller

gastropods ? commensalism (1874c)

Mollusca Cornulites commensalism Morris and Rollins

gastropods (1971);
Morris and Felton
(1993)

Mollusca Cornulites ? epizoism: ? Richards (1974)

gastropods commensalism

Mollusca trilobites ? endozoism ? Brandt (1993)

gastropods

Mollusca bryozoans encrustation Ulrich (1883)

monoplacoph-
Cornulites epizoism or Morris and Rollins
orans
commensalism (1971)

Mollusca brachiopods encrustation Davis and M a p e s

nautiloids (inartic.) (1996)

Mollusca bryozoans encrustation; Nicholson (1875)

nautiloids epizoism

244 A Sea without Fish

 ARRANGED BY "HOST"

Host Associate Type of References/

Association [Annotations]

Mollusca bryozoans epizoism; ? Frey (1988, 1989);

"nautiloids" commensalism Baird et al. (1989)
actinoceroids
bryozoans encrustation Davis and M a p e s
endoceroids
presumably (1996)
nautiloids
epizoism;
? commensalism

bryozoans encrustation Ulrich (1879a)

bryozoans encrustation U. P. James

(1884b)

bryozoans encrustation Ulrich (1883);

Bassler (1953)

bryozoans encrustation (post- Wilson, Palmer,

mortem); and Taylor (1994)
dwelling inside
empty shell

Cornulites ? epizoism Richards (1974)

? commensalism

hydrozoan encrustation (post- Wilson, Palmer,

mortem); and Taylor (1994)
dwelling inside
empty shell

stromatoporoid encrustation J. F. James (1886)

stromatoporoid encrustation Baird, Brett, and
Frey (1989)
trilobites ? incolenism Davis et al. (2001)

Mollusca Cornulites epizoism or Morris and Rollins

pelecypods commensalism (1971);
Morris and Felton
(2003)

ARTHROPODA

trilobites bryozoans epizoism Brandt (1996)

trilobites Cornulites epizoism Morris and Rollins
(1971); Brandt
(1996)

ECHINODERMATA

Echinodermata brachiopods aegism; epizoism Shrake (1989)

crinoids

Echinodermata brachiopods commensal Sandy (1996)

crinoids Zygospira

Echinodermata bryozoans encrustation Ulrich (1883)

crinoids

Echinodermata bryozoans encrustation: Bassler (1953)

crinoids epizoism

Life in the Cincinnatian Sea 245

 ARRANGED BY "HOST"

Host Associate Type of References/

Association [Annotations]

Echinodermata byroniids parasitism Warn (1974);

crinoids Welch (1976);
Malinky et al.
(2004)

Echinodermata Cornulites commensalism Morris and Rollins

crinoids (1971);
Morris and Felton
(1993, 2003);
Richards (1974)

Echinodermata gastropods commensalism Bowsher (1955);

crinoids Morris and Felton
(1993)

Echinodermata gastropods ? parasitism ? Baumiller and

crinoids Gahn (2002)

"WORMS"

(see also:
cornulitids)

246 A Sea without Fish

 PRIMARY
SUSPENSION HERBIVORE CARNIVORE PRODUCERS
chitinozoans (?) cephalopods acritarchs
conodonts eurypterids
graptolites
trilobites (pt)

SUSPENSION DEPOSIT HERBIVORE CARNIVORE

trilobites (pt) monoplacophorans monoplacophorans trilobites (pt)
gastropods gastropods asteroids
MOBILE ostracods
trilobites
ostracods
ophiuroids, asteroids (pt)

stromatoporoids (pt)
EPIFAUNA

ATTACHED tab., rugose corals (pt)

LOW bryozoans
craniate brachs
rhynch. brachs
bivalves
cornulitids SUSPENSION DEPOSIT CARNIVORE
edrioasteroids
cyclocystoids rostroconchs
SHALLOW
sponges PASSIVE
conulariids
ATTACHED Ungulate brachs bivalves
stromatoporoids (pt) polychaetes
ERECT SHALLOW bivalves polychaetes
tabulate corals (pt)
bryozoans ACTIVE Diplocraterion
rhombiferans
crinoids
DEEP
PASSIVE
rugose corals (pt)
RECLINING strophomenate brachs
tentaculitids DEEP
hyolithids ACTIVE
stylophorans

Table 4 A . Type-Cincinna-
tian Marine Guilds
Modified after Droser
and Sheehan (1997).

Life in the Cincinnatian Sea 247

 PRIMARY
SUSPENSION HERBIVORE CARNIVORE PRODUCERS
gastropods bony fish cephalopods dinoflagellates

PELAGIC
malacostracans mammals chondrichthyans coccolithophores
mammals bony fish diatoms
reptiles
mammals

SUSPENSION DEPOSIT HERBIVORE CARNIVORE

bivalves gastropods chitons gastropods
crinoids ostracods gastropods cephalopods
MOBILE ophiuroids (pt) malacostracans ostracods malacostracans
asteroids (pt) ophiuroids malacostracans asteroids
holothuroids asteroids (pt) echinoids
echinoids
holothuroids
EPIFAUNA

sponges
corals
ATTACHED bryozoans
LOW brachiopods
polychaetes
bivalves SUSPENSION DEPOSIT CARNIVORE
barnacles
gastropods bivalves bivalves
ATTACHED sponges SHALLOW bivalves
ERECT corals PASSIVE echinoids
bryozoans
crinoids lingulate brachs bivalves gastropods
corals (pt) SHALLOW bivalves polychaetes malacostracans
RECLINING
bivalves ACTIVE polychaetes echinoids polychaetes
<
INFAUNA

echinoids holothuroids

bivalves
DEEP
PASSIVE

bivalves bivalves polychaetes

DEEP polychaetes polychaetes
ACTIVE malacostracans

Table 4B. Mesozoic and

Cenozoic Marine Guilds
Modeled after Bambach
(1983).

248 A Sea without Fish

EPILOGUE: DIVING IN THE
CINCINNATIAN SEA

m a n y p a l e o n t o l o g i s t s , o u r s e l v e s i n c l u d e d , b e c a m e f a s c i n a t e d w i t h fossils
a n d e m b a r k e d o n scientific c a r e e r s l o n g b e f o r e w e e v e r e n c o u n t e r e d l i v i n g
m a r i n e a n i m a l s . For m a n y o f u s , t h e greatest thrill has b e e n o u r f i r s t e n -
c o u n t e r s w i t h l i v i n g representatives o f t h e a n i m a l g r o u p s w e k n e w f i r s t o n l y
as grey, lifeless f o r m s e n c a s e d in rock. B o t h of us h a v e b e e n p r i v i l e g e d to
e x a m i n e f i r s t h a n d l i v i n g relatives o f a n i m a l s o f o u r favorite g r o u p s o f fos-
s i l s c r i n o i d s for M e y e r a n d n a u t i l o i d c e p h a l o p o d s for D a v i s . O u r e x p e r i -
e n c e s h a v e f u e l e d a c u r i o s i t y that affects p r a c t i c a l l y a n y o n e w h o c o n t e m -
plates the fossil richness of the Cincinnatian or other comparable
fossiliferous strata. M a n y t i m e s , in the field, we stand on a C i n c i n n a t i a n
o u t c r o p w h e r e fossils a r e a b u n d a n t i n a l m o s t e v e r y r o c k , a n d w e w o n d e r :
w h a t did t h e C i n c i n n a t i a n sea a c t u a l l y l o o k like? H o w did t h e s e c r e a t u r e s
b e h a v e w h e n alive? It we c o u l d travel h a c k in t i m e to d i v e into t h e C i n c i n -
n a t i a n sea, what w o u l d w e see?
In his b o o k The Crucible of Creation, t h e p a l e o n t o l o g i s t S i m o n C o n -
w a y M o r r i s (1998) takes t h e r e a d e r on a j o u r n e y t h r o u g h t i m e in an i m a g i -
nary t i m e m a c h i n e that l a n d s o n t h e shores o f t h e C a m b r i a n sea i n w e s t e r n
C a n a d a o f 520 m i l l i o n years a g o . T h e t i m e m a c h i n e t h e n d e s c e n d s into
the sea and e n a b l e s t i m e t r a v e l i n g scientists to view t h e varied a n d b i z a r r e
a n i m a l s f o u n d a s f o s s i l s i n t h e f a m o u s B u r g e s s S h a l e . C o n w a y M o r r i s rec-
reated the e n v i r o n m e n t of the C a m b r i a n sea a n d t h e life w i t h i n it f r o m t h e
e v i d e n c e of the fossils and r o c k s , but he e m b e l l i s h e d the s c e n a r i o w i t h a
m e a s u r e o f s p e c u l a t i o n and fantasy.
W e r e we to travel b a c k to the Late O r d o v i c i a n , w o u l d we n e e d a d e e p -
d i v i n g s u b m e r s i b l e to e x p l o r e t h e C i n c i n n a t i a n w o r l d in a s i m i l a r , i m a g i -
nary journey? Based o n t h e e v i d e n c e f r o m t h e rocks a n d fossils (see c h a p t e r
15), t h e t i m e traveler t o C i n c i n n a t i i n t h e L a t e O r d o v i c i a n w o u l d h a v e t o
land o n t h e sea s u r f a c e , b e c a u s e n o dry l a n d w o u l d b e f o u n d . W i t h n o
f a m i l i a r l a n d m a r k s t h e O h i o R i v e r valley, t h e S u s p e n s i o n B r i d g e , o r
C a r e w T o w e r b r e a k i n g the h o r i z o n , w e w o u l d h e a t t r a c t e d t o t h e area
o n l y b y vast s h a l l o w s a p p e a r i n g a s v a r y i n g s h a d e s o f a q u a m a r i n e , w i t h o c -
c a s i o n a l shoals m a r k e d b y breakers.
O u r t i m e m a c h i n e t r a n s f o r m s into a s m a l l b o a t a s w e l a n d , a n d w e
c h e c k o u r position. T h e r e i s n o G l o b a l P o s i t i o n i n g S y s t e m o f satellites, s o
we use a sensitive d i p - n e e d l e that i n d i c a t e s that we are at 2 5 s o u t h l a t i t u d e ,
0

but w e h a v e n o reliable i n d i c a t o r o f l o n g i t u d e . T h e w a t e r feels c o o l , a b o u t

65F, so we n e e d wet suits for t h e dive. Breakers off to t h e s o u t h l o o k m e n -
acing, so we keep our distance and drop anchor where we can see the
b o t t o m . Let's s e e what's d o w n there!

249
 We d o n m a s k s and snorkels for a q u i c k r e c o n n o i t e r , a n d slip over the
side. We have a n c h o r e d over very shallow water, and the b o t t o m appears only
a m e t e r or so b e n e a t h us. As we take a closer look at the b o t t o m , we n o t i c e
that it is irregular, with low m o u n d s separated by p a t c h e s that are m o r e level.
T h e m o u n d s are a c t u a l l y c l u m p s o f large, ribbed b r a c h i o p o d s , Platystrophia
ponderosa. L i v i n g a n i m a l s with articulated shells are i n t e r m i n g l e d with sepa-
rated v a l v e s , s o m e b r o k e n a n d w o r n s m o o t h . It is the e n v i r o n m e n t that,
m i l l i o n s o f w a r s i n t h e f u t u r e , will b e preserved a s the M t . A u b u r n M e m b e r
o f the G r a n t L a k e L i m e s t o n e . W e e a s i l y s c o o p u p a s a m p l e o f s p e c i m e n s o f
Platystrophia b e c a u s e they h a v e no p e d i c l e a t t a c h m e n t s .
C l e a r l y t h e r e is m u c h of interest to see h e r e , but we return to the boat
b e c a u s e snorkel d i v i n g i s not a d e q u a t e for p r o l o n g e d e x p l o r a t i o n . W e h a v e
n o t b e e n a b l e t o h o l d o u r b r e a t h d u r i n g o u r dives a s l o n g a s w e n o r m a l l y
d o w e h a d t o c o m e u p q u i c k l y , g a s p i n g for air. A c h e e k o f o u r air q u a l i t y
m o n i t o r r e v e a l s t h e r e a s o n : t h e L a t e O r d o v i c i a n a t m o s p h e r e has o n l y a
f r a c t i o n of t h e o x y g e n c o n t e n t of p r e s e n t - d a y air, p e r h a p s as little as 1 per-
cent. Fortunately we have brought a l o n g s o m e sophisticated diving gear
that w i l l let us fill o u r d i v i n g c y l i n d e r s w i t h c o m p r e s s e d air in w h i c h we
h a v e b o o s t e d t h e o x y g e n c o n t e n t to its p r e s e n t - d a y l e v e l , 21 p e r c e n t . B e -
c a u s e the rest of o u r c o m p r e s s e d g a s m i x t u r e is p r e d o m i n a n t l y n i t r o g e n ,
we still h a v e to f o l l o w t h e d i v e tables that tell us how l o n g we c a n d i v e at a
g i v e n d e p t h a n d r e s u r f a c e w i t h o u t s u f f e r i n g d e c o m p r e s s i o n sickness. W e
n e e d to be really c a r e f u l : we c o u l d not be farther a w a y from a r e c o m p r e s -
sion c h a m b e r !
E q u i p p e d with c o l l e c t i n g gear and cameras, we resume our dive using
s c u b a gear. T h e w a t e r clarity a c t u a l l y i s q u i t e g o o d : w e c a n see p e r h a p s 1 5
m e t e r s (50 feet) h o r i z o n t a l l y . H o w e v e r , t h e r e are p a t c h e s o f t h e m u d and
silt c o v e r i n g w i d e areas of t h e b o t t o m that c o u l d be easily stirred up if
s u r f a c e w a v e s p i c k e d u p . V i s i b i l i t y c o u l d d r o p sharply. A s w e settle t o the
b o t t o m right b e n e a t h t h e b o a t , w e c a n feel s o m e w a v e m o t i o n . A l t h o u g h
t h e sea floor a p p e a r s very flat, we c a n see that there is a g e n t l e slope t a i l i n g
off toward t h e n o r t h , s o w e s w i m slowly toward d e e p e r water. W e c o v e r a
lot o f d i s t a n c e , b u t t h e d e p t h c h a n g e s very little. W e see vast areas o f the
sea floor c o v e r e d w i t h b r y o z o a n s in d e n s e t h i c k e t s or folded s h e e t s like a
r u m p l e d c a r p e t . A s w e g o d e e p e r , m o r e b u s h - l i k e b r y o z o a n c o l o n i e s ap-
p e a r , s o m e very d e l i c a t e . It is a m a z i n g to see how s i m i l a r the c o l o n y forms
are to t h o s e of present-day b r y o z o a n s . But we know that, in the O r d o v i c i a n ,
we are l o o k i n g at t r e p o s t o m e s , n o t at a n i m a l s of the g r o u p s that thrive in
t h e seas of today. In p l a c e s t h e sea floor is a h a r d , l i m e s t o n e p a v e m e n t with
a c o n v o l u t e d s u r f a c e ; d e p r e s s i o n s are Idled w i t h fine silt or shelly s a n d .
B r y o z o a n s a n d b r a c h i o p o d s a r e a t t a c h e d t o the hard s u r f a c e s . T h e heart
skips a b e a t as we s p o t for t h e first t i m e a l i v i n g e d r i o a s t e r o i d e c h i n o d e r m ,
a l s o a t t a c h e d like a p r e s e n t - d a y b a r n a c l e to the h a r d g r o u n d . B e c a u s e e d r i o -
asteroids b e c a m e e x t i n c t i n the L a t e P a l e o z o i c , w e h a v e n e v e r b e e n able t o
e n v i s i o n t h e l i v i n g a n i m a l w i t h c o n f i d e n c e until now. A c l u m p o f tall,
pillar-like s p e c i m e n s o f Streptaster s h o w s that C o l i n S u m r a l l had the right
idea i n s u g g e s t i n g t h a t s o m e e d r i o a s t e r o i d s c o u l d inflate their t h e c a e a n d
t e l e s c o p e u p o r d o w n f r o m t h e s u b s t r a t u m . Fine l u b e f e e t e x t e n d i n g from

250 A Sea without Fish

the a m b u l a c r a ] g r o o v e s recall t h e r e c o n s t r u c t i o n s m a d e b y B r u c e B e l l .
W i t h a rock pick w e easily b r e a k off a p i e c e o f t h e h a r d g r o u n d w i t h e d r i o -
asteroids a t t a c h e d , a n d we b a g it for s t u d y in t h e l a b . T h e h a r d g r o u n d g i v e s
w a y to an area c o v e r e d w i t h t h i n - s h e l l e d Rafinesquina b a c h i o p o d s , f o r m i n g
shell p a v e m e n t s like t h o s e w e f o u n d i n r o c k u n i t s w i t h n a m e s l i k e C o r -
ryville, B e l l e v u e , a n d f a i r v i e w i n t h e distant f u t u r e f r o m w h i c h w e c a m e .
T h e c o n c a v o - c o n v e x b r a c h i o p o d s rest w i t h t h e c o n v e x v a l v e e i t h e r d o w n
o r u p , a n d m a i n are e n c r u s t e d w i t h s m a l l b r y o z o a n s o r e d r i o a s t e r o i d s .
Brachiopods of taxa l i k e Zygospira and pelecypods of g e n e r a like Cari-
todens are a t t a c h e d . O t h e r c l a m s , of taxa like Modiolopsis, p o k e up t h r o u g h
t h e s e d i m e n t b e t w e e n shells. S m a l l Flexicalymene a n d Acidaspis trilobites
g l i d e over the s u r f a c e , a n d h e r e a n d t h e r e a c r i n o i d has t h e s t e m c o i l e d
a r o u n d a b r y o z o a n . T h i s is a d i v e r s e habitat.
But the w a t e r m a s s a b o v e the sea f l o o r i s s u r p r i s i n g l y e m p t y c o m p a r e d
to the s c e n e in p r e s e n t - d a y shallow seas. Today, fish are e v e r y w h e r e in t h e
sea, f i l l i n g a w i d e variety o f e c o l o g i c a l roles. W h e r e are t h e f i s h i n this
Ordovician sea? O n l y small n a u t i l o i d c e p h a l o p o d s are j e t t i n g a b o u t ,
a g a i n s t a b a c k d r o p o f p u l s a t i n g jellyfish. A t c l o s e r a n g e , w e c a n p i c k o u t
very s m a l l strings s u s p e n d e d i n t h e w a t e r w i t h c l u m p s o f m i n u t e t e n t a c l e s
a r r a n g e d in vertical series. These are graptolites. C l o s e r to t h e b o t t o m s o m e
s m a l l , spiny trilobites, p r o b a b l y o d o n t o p l e u r i d s , s w i m a b o v e t h e b o t t o m for
short d i s t a n c e s . A l t h o u g h early, jawless f i s h i n h a b i t s o m e L a t e O r d o v i c i a n
seas e l s e w h e r e , t h e y h a d n o t y e t spread to t h e C i n c i n n a t i r e g i o n . It t r u l y is
a sea w i t h o u t f i s h ! W e d o not h a v e t o w o r r y a b o u t sharks e i t h e r , b e c a u s e
t h e y w i l l n o t e v o l v e u n t i l t h e D e v o n i a n m i l l i o n s o f years i n t h e f u t u r e .
O u r depth gauge shows that, as we s w i m farther away from the boat,
the b o t t o m g r a d u a l l y rises, a n d i t b e c o m e s littered w i t h b r a c h i o p o d s a n d
b r y o z o a n s . H e r e a n d t h e r e m o u n d s rise u p a s tall a s a m e t e r f r o m t h e bot-
t o m . A l t h o u g h t h e m o u n d s l o o k s o m e w h a t like c o r a l c o l o n i e s , c l o s e r in-
s p e c t i o n reveals that t h e y are h e a v i l y c a l c i f i e d c o l o n i c s o f b r y o z o a n s , w h o s e
f u z z y - l o o k i n g s u r f a c e s a r e m i l l i o n s o f o h - s o - t i n y t e n t a c l e s e x t e n d e d into
the water. They retract w h e n w e b r u s h a g a i n s t t h e s u r f a c e , a n d w e c a n see
t h e m i n u t e h o n e y c o m b - l i k e s k e l e t o n b e n e a t h . S o m e c o l o n i e s are r o u n d e d
and b o u l d e r - l i k e ; o t h e r s are b r a n c h i n g o r c o m p o s e d o f c o m p l e x l y f o l d e d
sheets. T h e p a t c h e s b e t w e e n b r y o z o a n s are littered w i t h b r o k e n f r a g m e n t s
of b r y o z o a n s a n d brachiopods of g e n e r a like Hebertella a n d Platystrophia,
including both living a n i m a l s and dead, disarticulated shells. H e r e a n d
there w e see b r y o z o a n c o l o n i e s a p p a r e n t l y t u r n e d u p s i d e d o w n ; w e k n o w
that b e c a u s e t h e r o u n d e d side o f e a c h i s o n t h e b o t t o m , o r t h e b r a n c h e s d o
not radiate u p w a r d f r o m a basal plate. T h e s e hefty, o v e r t u r n e d c o l o n i e s
s u g g e s t that c o n d i t i o n s c a n b e far m o r e t u r b u l e n t t h a n t h e p l a c i d c a l m i n
w h i c h , w i t h great g o o d l u c k , w e l a n d e d . T h e s e w a t e r s s o m e t i m e s m u s t b e
raked b y great storms. W e feel s t r o n g e r w a v e m o t i o n a n d s o o n w e are b r e a k -
i n g t h e s u r f a c e on this s h o a l . P a t c h e s of d e a d , t h i n - s h e l l e d Rafinesquina
b r a c h i o p o d s are t u r n e d o n their e d g e s a n d p a c k e d t i g h t l y t o g e t h e r b y w a v e
a c t i o n w h a t will be c a l l e d " s h i n g l e d Rafinesquina" b e d s in t h e far-distant
f u t u r e . B r y o z o a n b r a n c h e s a n d s h e e t s are s t a c k e d into p a t c h e s o f c o a r s e
rubble. This is the e n v i r o n m e n t of the B e l l e v u e L i m e s t o n e to be.

Epilogue 251
 A g u l l y leads us b a c k into d e e p e r water. H e r e we find vast areas c o v e r e d
w i t h b r a c h i o p o d s of taxa l i k e Rafinesquina a n d Strophomena or b r a n c h i n g
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 o f m u d a b o u t e q u a l i n a r e a t o
t h e s h e l l y p a t c h e s ; this m u s t b e t h e e n v i r o n m e n t o f t h e F a i r v i e w , w i t h
c o n t i n u o u s , e v e n b e d s that w i l l p r o d u c e a b o u t 5 0 p e r c e n t l i m e s t o n e a n d
50 p e r c e n t s h a l e .
In t h e d i s t a n c e we spot a ridge of b r y o z o a n s s t a n d i n g a l m o s t a m e t e r
a b o v e the s u r r o u n d i n g s , a n d w e h e a d toward it. C o u l d w e h e a p p r o a c h i n g a
s h a r p d r o p - o f f l e a d i n g to d e e p e r water? A p p r o a c h i n g closer, we c a n see a
r i c h n e s s of life a r o u n d this ridge, a n d we sense a g e n t l e c u r r e n t flowing a l o n g
the b o t t o m , parallel to the g r a d u a l slope, w h i c h intensifies as we reach the
ridge. T h e d e p t h g a u g e reads 15 meters (50 feet), a n d we h a v e d e s c e n d e d
b e l o w the d e p t h w h e r e t h e s m a l l s u r f a c e w a v e s stirred the b o t t o m . H e r e ,
c u r r e n t flow takes o v e r a n d follows t h e c o n t o u r s o f the slope. T h e ridge i s
a c t u a l l y a m o u n d b u i l t entirely o f the b r a n c h i n g b r y o z o a n c o l o n i e s o f g e n u s
Parvohallopora; it projects o u t w a r d from the slope, The c u r r e n t is diverted
a n d g a i n s v e l o c i t y as it flows over t h e ridge. C r i n o i d s f o r m i n g a d e n s e c l u m p
are of g e n u s Glyptocrinus; their stalks are coiled around the bryozoan
b r a n c h e s , a n d t h e c r i n o i d s stand a b o v e t h e m like a forest c a n o p y . Their
c r o w n s are splayed o u t in feathery filtration fans that are all a l i g n e d p e r p e n -
d i c u l a r to t h e c u r r e n t . S t a n d i n g a b o v e t h e m are a few g i g a n t i c crinoids of
g e n u s Anomalocrinus, w i t h thick s t e m s a m e t e r or m o r e in l e n g t h a n d broad,
d e n s e filtration fans s o m e 30 c e n t i m e t e r s in d i a m e t e r , also oriented by the
c u r r e n t . M a n y o t h e r invertebrates m a k e their h o m e s i n this intricate thicket.
Gyclonema snails are a t t a c h e d to the c a l y c e s of practically every s p e c i m e n of
Glyptocrinus a n d c l u m p s of s m a l l Zygospira b r a c h i o p o d s attach to the cri-
n o i d stems. O t h e r b r a c h i o p o d s o f g e n u s Platystrophia and p e l e c y p o d s o f
Ambonychia are n e s t l e d in a m o n g the b r y o z o a n b r a n c h e s .
S u d d e n l y w e are s t a r t l e d b y streaky s h a d o w s p a s s i n g o v e r us, a n d look
u p t o see l a r g e , c o n i c a l n a u t i l o i d c e p h a l o p o d s g l i d i n g a b o v e u s , a l i g n e d
l i k e a i r p l a n e s i n f o r m a t i o n . T h e y b e h a v e m u c h like s c h o o l i n g s q u i d , b u t
e a c h has a n o u t e r s h e l l , w i t h a p a t t e r n o f c o l o r b a n d i n g . O n e i s h o l d i n g a
l a r g e , w r i g g l i n g trilobite i n its t e n t a c l e s . T h e s e c e p h a l o p o d s are the largest
creatures we have seen, and s o m e of these reach a meter or more in length,
a n d their l a r g e e y e s a n d n u m e r o u s t e n t a c l e s are m o r e t h a n a little m e n a c -
ing. We h u n k e r d o w n next to the ridge and point the camera upward as
t h e y s h o o t b y o v e r h e a d . I f this p i c t u r e d o e s not m a k e t h e c o v e r o f Science,
i t w i l l a t least b e t h e hit o f t h e n e x t m e e t i n g o f t h e P a l e o n t o l o g i c a l S o c i e t y !
In a flash t h e y are g o n e , a n d o u r b r e a t h i n g rate settles d o w n .
In a h o l l o w of t h e r i d g e we find a l a r g e Isotelus trilobite, its p y g i d i u m
(tail s h i e l d ) f o l d e d n e a t l y u n d e r its c e p h a l o n ( h e a d s h i e l d ) i t m u s t h a v e
s e e n us c o m i n g a n d r o l l e d up for p r o t e c t i o n . It is a g i a n t , a b o u t 25 c e n t i -
m e t e r s b e t w e e n t h e s h a r p tips o f t h e g e n a l s p i n e s . T h e t e m p t a t i o n t o grasp
a l i v i n g trilobite is t o o g r e a t , a n d as we r e a c h o u t for this p r i z e , t h e a n i m a l
s u d d e n l y u n f o l d s w i t h a s n a p a n d g l i d e s a w a y , v e n t r a l side u p , o u t o f r e a c h .
I t t h e n f l i p s b a c k o v e r a n d c l i n g s tightly t o t h e b o t t o m , p r e s s i n g t h e dorsal
c a r a p a c e d o w n into t h e s e d i m e n t . H a d this o n e n o t g o t t e n away, i t m i g h t
h a v e s u r p a s s e d t h e w o r l d r e c o r d s p e c i m e n o f Isotelus f r o m t h e O r d o v i c i a n

252 A Sea without Fish

o f n o r t h e r n C a n a d a ! N e v e r t h e l e s s , w e h a v e m a n a g e d t o g a t h e r u p several
s m a l l e r trilobites, a n d m a y b e their D N A f i n a l l y will resolve the q u e s t i o n o f
h o w trilobites are related to o t h e r a r t h r o p o d s .
We v e n t u r e out beyond the bryozoan ridge o n t o a seemingly level plain
with patches o f b r a c h i o p o d p a v e m e n t and b r y o z o a n s . The b r a c h i o p o d s are
n o t i c e a b l y s m a l l e r forms like Dalmanella a n d Sowerhyella. B r y o z o a n s are
delicate, twig-like c o l o n i e s with fewer sheet-like or m a s s i v e forms. Flexicaly-
mene trilobites are here, hut are s m a l l e r t h a n t h o s e we f o u n d in s h a l l o w e r
water; s o m e n e w trilobites of g e n e r a like Cryptolithus and Triarthrus c r u i s e
o n the m u d d y p a t c h e s , l e a v i n g g r o o v e d trails. E x t e n d i n g u p w a r d from s o m e
b r y o z o a n thickets are very slender yet l o n g - s t e m m e d c r i n o i d s of the taxa
Ectenocrinus and Cincinnaticrinus. A l t h o u g h they h a v e fewer a r m s t h a n the
other crinoids we have seen today, they splay t h e m into c o n i c a l filtration fans
in a l i g n m e n t with the g e n t l e c u r r e n t (not o p e n into the flow b u t w i t h the
c o n c a v e side b e a r i n g the food g r o o v e s d o w n c u r r e n t ) . S o m e s e d i m e n t p a t c h e s
are sands entirely c o m p o s e d o f f r a g m e n t s o f disarticulated c r i n o i d s , w i t h o d d
b u n d l e s o f l o n g , still-articulated steins. T h e sands h a v e b r o a d , s i n u o u s ripple
marks like those a l o n g a b e a c h . H o w c o u l d s u c h ripples form at o u r present
d e p t h of 30 meters (100 feet)? We sense no w a v e m o t i o n a n d o n l y slight cur-
rent; however, as e x p e r i e n c e d divers, we k n o w that severe storm c o n d i t i o n s
a t the surface, i n c l u d i n g h u r r i c a n e s , c a n p r o d u c e o c c a s i o n a l , s t r o n g w a v e
oscillation or currents a l o n g the d e e p e r sea floor w h e r e n o r m a l , fair w e a t h e r
wave m o t i o n s d o not penetrate. W e have a n u n e a s y f e e l i n g that w e h a v e
v e n t u r e d into a n e n v i r o n m e n t that c a n t u r n v i o l e n t v e r y q u i c k l y , a s w e see
a r o u n d u s the r e m n a n t s o f shattered b r y o z o a n s a n d c u r r e n t - w i n n o w e d shells.
Areas o f p u r e m u d h a v e now b e c o m e w i d e r t h a n areas o f shelly s e d i m e n t ,
and w e r e c o g n i z e the characteristics o f what will c o n i c t o b e k n o w n a s the
Kope Formation.
A g l a n c e at the dive c o m p u t e r s h o w s we h a v e a l m o s t e x c e e d e d o u r al-
lowable b o t t o m t i m e at t h e m a x i m u m d e p t h of 30 m e t e r s , so it is t i m e to
return to the shallows. We turn b a c k on a reverse c o m p a s s c o u r s e , h o p i n g
we we will surface close to the boat. As we a g a i n a p p r o a c h t h e b r y o z o a n ridge,
w e n o t i c e a n o v e r h a n g w e h a d o v e r l o o k e d . S o m e k i n d o f m o v e m e n t i s appar-
e n t i n t h e s h a d o w s b e n e a t h this l e d g e . W h i p - l i k e , spiny a p p e n d a g e s are
w a v i n g at us; this is s o m e t h i n g n e w w e m u s t investigate! U s i n g a l o n g ,
slender pole with a hook at o n e e n d , we reach b a c k u n d e r the l e d g e . After a
few u n s u c c e s s f u l tries, we h a v e s n a g g e d a large a n d b i z a r r e c r e a t u r e . We
r e c o g n i z e the flailing a p p e n d a g e s of a e u r y p t e r i d of g e n u s Megalograptus. It
is the size of a M a i n e lobster but has very strange a n d spiny, non-lobster-like
a p p e n d a g e s . The e l o n g a t e , s e g m e n t e d tail s e c t i o n is flexible like a lobster tail.
We do not h a v e t i m e to o b s e r v e t h e e u r y p t e r i d s n o w , so we p o p a few into a
m e s h h a g for later study. T h e m y s t e r y o f h o w t h e y u s e their a p p e n d a g e s will
r e m a i n until t h e n . Perhaps we will see w h e t h e r o n e tastes like lobster!
A f t e r m o r e f i n n i n g toward t h e s h a l l o w s , o u r d i v e c o m p u t e r b e e p s , a n d
w e m u s t a s c e n d . W e b r e a k t h e s u r f a c e a n d s w i t c h t o snorkel b e c a u s e w e
are l o w o n air. W e spot the b o a t i n t h e d i s t a n c e , p e r h a p s 200 m e t e r s away.
It is a l o n g s w i m on t h e s u r f a c e . We are l a d e n w i t h s p e c i m e n b a g s a n d g e a r ,
a n d , b y t h e t i m e w e reach the b o a t , w e are e x h a u s t e d f r o m g u l p i n g t h e

Epilogue 253
 o x y g e n - p o o r air. W e h a u l o u r s e l v e s o v e r t h e g u n w a l e , peel off o u r w e t s u i t s ,
a n d just lie i n t h e b o a t , c a t c h i n g o u r b r e a t h , o u r m i n d s r a c i n g w i t h the
sights w e h a v e s e e n . W h a t a d i v e !
S u d d e n l y , w e s e n s e t h e w a r m t h o f t h e a f t e r n o o n s u n , a m i d the c o o l -
ness o f a c r i s p a u t u m n day. W e are s t a r i n g a t t h e fossil-covered s u r f a c e o f
a b e d o f O r d o v i c i a n l i m e s t o n e littered w i t h rusty a u t u m n leaves. W e are
sitting o n t h e b a n k s o f S t o n e l i c k C r e e k , i n C l e r m o n t C o u n t y , O h i o , h a v i n g
l u n c h o n a field trip w i t h o u r s t u d e n t s a n d m a y b e y o u !

254 A Sea without Fish

APPENDIX 1. RESOURCES: WHERE
TO GO FOR MORE INFORMATION

T h e r e are m a n y t e x t b o o k s in p a l e o n t o l o g y , but we restrict t h e f o l l o w i n g list Paleontology

to some of the most r e c e n t as w e l l as o n e older, classic w o r k . textbooks
Fossil Invertebrates ( B o a r d m a n et al. 1987)
Principles of Paleontology, 3rd e d . ( F o o t e a n d M i l l e r 2007)
Invertebrate Fossils ( M o o r e et al. 1952)
Invertebrate Palaeontology and Evolution, 4th ed. (Clarkson 1998)

Ohio Fossils ( L a R o c q u e a n d M a r p l e 1955) Publications of

Fossils of Ohio (Feldmann a n d H a c k a t h o r n 1996) Geological Surveys
Exploring the Geology of the Cincinnati/Northern Kentucky Region, 2nd
e d . (Potter 2007)

Cincinnati Fossils ( D a v i s 1985,1992) and its p r e d e c e s s o r s ( C a s t e r et al. 1955, Locally published

1961) books

Index Fossils of North America ( S h i n i e r a n d S h r o c k 1944) Encyclopedic w o r k s

Treatise on Invertebrate Paleontology

R. A. Davis's " T h e T y p e - C i n c i n n a t i a n , " http://inside.nisj.edu/acadeinics/ Internet w e b s i t e s

f a c u l t v / d a v i s r / c i n t i a n / i n d e x . h t m . T h e C o l l e g e o f M o u n t St. J o s e p h , C i n -
c i n n a t i , O h i o (accessed F e b r u a r y 18, 2008).
The D r y D r e d g e r s , http://drydredgers.org/, D r y D r e d g e r s , Inc., C i n c i n n a t i ,
O h i o (accessed F e b r u a r y 18, 2008).
E a r t h T i m e , h t t p : / / w w w . e a r t h - t i i u e . o r g / a b o u t . h t m l ( a c c e s s e d F e b r u a r y 18,
2008).
"History of Life through T i m e , " http://www.ucmp.berkeley.edu/exhibits/
historyoflite.php. University of C a l i f o r n i a M u s e u m of Paleontology (ac-
c e s s e d February 18, 2008).
Steven H o l l a n d ' s " T h e Stratigraphy and fossils o f the U p p e r O r d o v i c i a n near
C i n c i n n a t i , O h i o , " http://www.uga.edu/~strata/cincy/index.html,The Uni-
versity of G e o r g i a Stratigraphy L a b (accessed F e b r u a r y 18,2008).
I n d i a n a G e o l o g i c a l S u r v e y ; http://igs.indiana.edu/, I n d i a n a U n i v e r s i t y (ac-
c e s s e d F e b r u a r y 18, 2008).
International G e o l o g i c a l Correlation P r o g r a m m e , I G C P 4 1 0 , " T h e Great
O r d o v i c i a n B i o d i v e r s i f i c a t i o n E v e n t . I m p l i c a t i o n s for G l o b a l C o r -

255
 relation a n d R e s o u r c e s , " h t t p : / / w w w . e s . m q . e d u . a u / M U C E P / i g c p 4 1 0 / ,
Macquarie University, Sydney, Australia (accessed F e b r u a r y 18,
2008).
K e n t u c k y G e o l o g i c a l S u r v e y , http://wwvv.uky.edu/KGS/, University o f K e n -
t u c k y ( a c c e s s e d F e b r u a r y 18, 2008).
K e n t u c k y P a l e o n t o l o g y S o c i e t y , h t t p : / / w w w . u k y . e d u / O t h e r o r g s / K P S / (ac-
c e s s e d F e b r u a r y 18, 2008).
O h i o G e o l o g i c a l Survey, http://www.ohiodnr.com/geosurvey Division of
G e o l o g i c a l S u r v e y , O h i o D e p a r t m e n t o f N a t u r a l R e s o u r c e s (ac-
c e s s e d F e b r u a r y 18, 2008).
University of Cincinnati Department of Geology, http://www.uc.edu/
g e o l o g y / ( a c c e s s e d F e b r u a r y 18, 2008).

Field g u i d e s T h e s e are g u i d e b o o k s t o f i e l d trips p e r t a i n i n g t o the O r d o v i c i a n g e o l o g y

o f t h e O h i o , I n d i a n a , a n d K e n t u c k y r e g i o n s . Most c o n t a i n d e t a i l e d road-
logs a n d d i r e c t i o n s to g e o l o g i c a l l o c a l i t i e s , as w e l l as d e t a i l e d d e s c r i p t i o n s
o f e x p o s e d s t r a t i g r a p h i c s e c t i o n s . L o c a l i t i e s listed i n o l d e r g u i d e b o o k s m a y
no longer be accessible.

C a s t e r 1961b; Hattin et al. 1961; P o p e a n d M a r t i n 1977; Hay et al. 1981;

M e y e r et al. 1981; M e y e r et a l . 1985; D a v i s 1986; H a n e b e r g et al. 1992;
S h r a k e 1992; D a v i s a n d C u f f e y 1998; A l g e o a n d Brett 2001; M c L a u g h l i n
e t al. f o r t h c o m i n g .

Museums Behringer-Crawford Museum, Covington, Kentucky http://www

. b c m u s e u m . o r g / b c m u s e u m / d e f a u l t . a s p x (accessed February 18, 2008)
C i n c i n n a t i M u s e u m C e n t e r h t t p : / / w w w . c i n c y m u s e u m . o r g / (accessed F e b -
r u a r y 18, 2008)
L i m p e r G e o l o g i c a l M u s e u m , M i a m i University, O x f o r d , O h i o http://www
. c a s . m u o h i o . e d u / l i m p e r m u s e u m / (accessed F e b r u a r y 18, 2008)
O r t o n G e o l o g i c a l M u s e u m , O h i o State University, C o l u m b u s , O h i o http://
w w w . g e o l o g y . o h i o - s t a t e . e d u / f a c i l i t i e s . p h p ( a c c e s s e d F e b r u a r y 18, 2008)

Outdoor C a e s a r C r e e k S t a t e Park, n e a r W a y n e s v i l l e , O h i o http://www.dnr.state

e d u c a t i o n areas .oh.us/tabid/72o/default.aspx/ ( a c c e s s e d F e b r u a r y 18, 2008)
A t t h e V i s i t o r s ' C e n t e r , t h e r e i s a n e x h i b i t a b o u t t h e g e o l o g y a n d fossils
to be f o u n d at t h e o v e r f l o w spillway, w h e r e fossil c o l l e c t i n g is p e r m i t t e d
i n a c c o r d a n c e w i t h c e r t a i n r e g u l a t i o n s a v a i l a b l e a t the Visitors' C e n t e r .
Hueston Woods State Park, near Oxford, Ohio http://www.dnr.state
.oh.us/parks/tabid/745/Default.aspx ( a c c e s s e d F e b r u a r y 18, 2008)
Fossil c o l l e c t i n g is p e r m i t t e d in c e r t a i n areas of t h e park.
C i n c i n n a t i Nature C e n t e r , near Milford, O h i o http://www.cincynature
.org/index2.asp/ ( a c c e s s e d F e b r u a r y 18, 2008)
A l t h o u g h fossil c o l l e c t i n g i s not p e r m i t t e d , t h e r e are g o o d e x p o s u r e s o f
C i n c i n n a t i a n strata a t several sites o n t h e N a t u r e C e n t e r p r o p e r t i e s .

256 Appendix 1
Sawyer Point Geological Timeline, Cincinnati, Ohio http://www
.cincinnati-oh.gov/crc/pages/-5708-/ ( a c c e s s e d F e b r u a r y 18, 2008)
T h i s timeline begins with the Late Ordovician and continues through
the f o u n d i n g o f C i n c i n n a t i w i t h e a c h p a v e m e n t b l o c k r e p r e s e n t i n g o n e
m i l l i o n years. I m p o r t a n t g e o l o g i c a l e v e n t s are e n g r a v e d o n b l o c k s a t a p -
propriate intervals.
Trammel Fossil Park, Sharonville, Ohio http://www.sharonville.org/
fossilpark.aspx ( a c c e s s e d F e b r u a r y 18, 2008)
T h i s park i s d e d i c a t e d t o e d u c a t i o n a b o u t O r d o v i c i a n g e o l o g y a n d p a l e -
o n t o l o g y (see F i g u r e 1.8).

P a l e o n t o l o g i c a l S o c i e t y . T h e P a l e o n t o l o g i c a l S o c i e t y i s t h e largest p a l e o n - Scientific societies

t o l o g i c a l o r g a n i z a t i o n in t h e U n i t e d States. It p u b l i s h e s b o t h t h e J o u r n a l and institutions
of Paleontology a n d Paleobiology. A series of e d u c a t i o n a l b r o c h u r e s a b o u t
fossils c a n b e d o w n l o a d e d f r o m their w e b s i t e http://paleosoc.org/ (ac-
c e s s e d F e b r u a r y 18, 2008).
P a l e o n t o l o g i c a l R e s e a r c h Institution, Ithaca, N e w York. P R I p u b l i s h e s b o t h
the Bulletins of American Paleontology and Palaeontographica Americana,
as well as a p o p u l a r m a g a z i n e , American Paleontologist. T h e i r website is
http://www.priweb.org/ (accessed F e b r u a r y 18, 2008).
G e o l o g i c a l Society of A m e r i c a . T h e G S A is the l e a d i n g geological organi-
z a t i o n in N o r t h A m e r i c a a n d it p u b l i s h e s b o t h t h e Geological Society of
America Bulletin a n d Geology. It sponsors m a n y r e g i o n a l a n d national
scientific m e e t i n g s and field trips. T h e i r website is http://www.geosociety
.org/ ( a c c e s s e d F e b r u a r y 18, 2008).

Appendix 1 257
APPENDIX 2. INDIVIDUALS AND
INSTITUTIONS ASSOCIATED WITH
THE TYPE-CINCINNATIAN

T h e f o l l o w i n g i s a list o f t h e n a m e s o f i n d i v i d u a l s a n d i n s t i t u t i o n s a s s o c i -
ated w i t h t h e C i n c i n n a t i r e g i o n , a n d , e s p e c i a l l y , its g e o l o g y a n d p a l e o n t o l -
ogy. S o m e o f the i n d i v i d u a l s listed w e r e m e m b e r s o f t h e C i n c i n n a t i S c h o o l ;
most were not.
T h e r e are s o m e potential p r o b l e m s w i t h this list. I n s o m e i n s t a n c e s ,
there are t w o p e o p l e w i t h similar, b u t different n a m e s , b u t w h o m a y n o t b e
different p e o p l e . For e x a m p l e , different s o u r c e s refer to a J. H. H a l l a n d a
John W . H a l l associated w i t h t h e C i n c i n n a t i S o c i e t y o f N a t u r a l History, a n d
there is I. Harris, I. H. Harris, a n d I. M. H a r r i s , all of W a y n e s v i l l e , O h i o .
G e o r g e V a l l a n d i n g h a m a n d G e o r g e V a l l a n d i g h a m are a l m o s t c e r t a i n l y t h e
s a m e p e r s o n , a n d the latter p r o b a b l y i s t h e c o r r e c t s p e l l i n g , b u t m a y b e not.
I n this v o l u m e , w e present p h o t o g r a p h s o f s o m e o f t h e p e o p l e d i s c u s s e d .
M a n y o f the i n d i v i d u a l s p o r t r a y e d are sufficiently w e l l k n o w n that t h e r e i s
little q u e s t i o n of identification. In s o m e i n s t a n c e s , h o w e v e r , a p h o t o g r a p h is
the o n l y o n e of w h i c h we are a w a r e that is s u p p o s e d to r e p r e s e n t t h e p e r s o n
i n q u e s t i o n . T h e identification m a y b e b a s e d o n a h a n d w r i t t e n n o t a t i o n o n
t h e p h o t o g r a p h o r o n t h e a l b u m p a g e that b e a r s t h e p h o t o g r a p h , w i t h n o
i n d e p e n d e n t verification. W e h o p e that s u c h identifications are c o r r e c t .

L i s t e d as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936). Albers, H. E.

Publications by T r u m a n H e m i n w a y Aldrich on Tertiary a n d present-day Aldrich, T. H.

m o l l u s c s a p p e a r e d in early n u m b e r s of the Journal of the Cincinnati Society (1848-1932)
of Natural History ( C o a n , K a b a t , a n d Petit 2007; J o h n s o n 2002; t h e g i v e n
n a m e " T h o m a s " in C a s t e r [1982] a p p a r e n t l y w a s a t y p o g r a p h i c a l error).

D . R . A n d e r s o n c o l l e c t e d o n e o f t h e o r i g i n a l s p e c i m e n s o f t h e starfish, A n d e r s o n , D. R.
Palaeasterina approximata S. A. M i l l e r , 1878.

A n t h o n y was elected recording secretary of the Western A c a d e m y of N a t u - A n t h o n y , John

ral S c i e n c e s o n M a y 1,1838. I n 1840, h e w a s e l e c t e d s e c r e t a r y o f t h e N a t u r a l Gould (1804-1877)
S c i e n c e S e c t i o n o f t h e C i n c i n n a t i S o c i e t y for t h e P r o m o t i o n o f U s e f u l
K n o w l e d g e , w h i c h m e r g e d w i t h t h e a c a d e m y later that year. H e i s r e c o r d e d
a s h a v i n g g i v e n s o m e fossils t o t h e a c a d e m y , a n d h e p u b l i s h e d o n t y p e -

259
 C i n c i n n a t i a n a n d o t h e r t r i l o b i t e s . A n t h o n y w a s o n e o f t h e hosts w h e n
C h a r l e s L y e l l , p r o b a b l y t h e f o r e m o s t g e o l o g i s t o n E a r t h , visited t h e C i n c i n -
nati a r e a in t h e 1840s. A n t h o n y w a s a u t h o r , a l o n g w i t h U. P. J a m e s , of a
paper on e c h i n o d e r m s in the local rocks, and he described an unusual
s p e c i m e n o f a fossil c e p h a l o p o d f r o m t h e t y p e - C i n c i n n a t i a n . I n 1854, h e
r e s i g n e d f r o m t h e a c a d e m y , a n d , i n 1863, h e b e c a m e c u r a t o r o f c o n c h o l o g y
a t H a r v a r d University, u n d e r L o u i s A g a s s i z . A c c o r d i n g t o C l e n c h (1936,69),
A n t h o n y ' s ". . . c h i e f interest w a s in f r e s h w a t e r m o l l u s k s , a n d he b u i l t up a
series o f A m e r i c a n f r e s h w a t e r f o r m s t h a t for its t i m e w a s s u p e r i o r t o a n y
c o l l e c t i o n i n t h i s c o u n t r y " ( A n t h o n y 1838,1839a, 1839b, 1847,1848; A n t h o n y
a n d J a m e s 1846; B r a n d t a n d D a v i s 2007; C o a n , K a b a t , a n d Petit 2007;
H e n d r i c k s o n 1947; J o h n s o n 2002; L y e l l 1845).

Austin, G e o r g e M. D r . A u s t i n , a p h y s i c i a n f r o m W i l m i n g t o n , O h i o , w a s listed a s a l o c a l c o l -
l e c t o r b y N i c k l e s (1936). H i s c o l l e c t i o n w e n t t o t h e U n i t e d States N a t i o n a l
M u s e u m , w h i c h p u b l i s h e d a p a p e r b y h i m o n fossil z o n e s i n t h e R i c h m o n -
d i a n r o c k s o f t h e t y p e - C i n c i n n a t i a n a r e a ( A u s t i n 1927; B e c k e r 1938; N i c k l e s
1936; S h i d e l e r [1952] 2002).

Barnhart, W. C. W . C . B a r n h a r t c o l l e c t e d t h e t y p e - s p e c i m e n o f t h e starfish, Palaeasterina

speciosa S. A. M i l l e r a n d D y e r , 1878. He sold it to J. W. H a r v e y , w h o , in
t u r n , sold i t t o C . B . D y e r ( M i l l e r a n d D y e r 1878a).

Bassler, Ray S. ( S e e c h a p t e r 2)
(1878-1961)

Bean, W. H. T h e b r y o z o a n s p e c i e s , Ceramopora ? beam, w a s n a m e d after W . H . B e a n ,

o f L e b a n o n , W a r r e n C o u n t y , O h i o ( U . R J a m e s 1884a).

Best, Robert Dr. Best was the principal curator of the Western M u s e u m at the time of
its o p e n i n g in 1820 ( D r a k e a n d M a n s f i e l d 1827).

Braun, E. Lucy E m m a L u c y B r a u n g a i n e d world f a m e a s a b o t a n i s t a n d plant e c o l o g i s t , b u t

(1889-1971) prior to that s h e a u t h o r e d t h e first p a l e o n t o l o g y thesis at t h e University of
C i n c i n n a t i , a master's thesis on C i n c i n n a t i a n b r a c h i o p o d s that later was p u b -
lished by t h e C i n c i n n a t i S o c i e t y of N a t u r a l History ( B r a u n 1916; C a s t e r 1981}.

Braun, Fred F r e d e r i c k B r a u n w a s listed as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936), a n d a

c o l l e c t i o n of his fossils w e n t to Y a l e ( S h i d e l e r [1952] 2002). B r a u n sold fossils
a s " F r e d B r a u n a n d C o . " a n d " t h e W e s t e r n Naturalist's A g e n c y " ( C a s t e r 1982,

260 Appendix 2
25), a n d , i n c o n j u n c t i o n w i t h Paul M o h r , a n o t h e r C i n c i n n a t i c o l l e c t o r , u n -
dertook l o n g - t e r m a n d e x t e n s i v e e x c a v a t i o n s for c r i n o i d s f r o m C a r b o n i f e r o u s
rocks a t C r a w f o r d s v i l l e , I n d i a n a ( V a n S a n t a n d L a n e 1964).

Dr. Bridge was born in N o r w o o d , O h i o , adjacent to C i n c i n n a t i , and he Bridge, Josiah

r e c e i v e d his b a c h e l o r ' s d e g r e e f r o m t h e U n i v e r s i t y o f C i n c i n n a t i i n 1913. (1890-1953)
His h i g h e r e d u c a t i o n w a s e l s e w h e r e , a n d p r o f e s s i o n a l l y h e w a s a s s o c i a t e d
with the United States ( G e o l o g i c a l Survey a n d the United States N a t i o n a l
M u s e u m in W a s h i n g t o n , D . C . He was a long-time associate of Ulrich
( B e c k e r 1938; C r o n e i s 1963).

Dr. Richard M a h a n Byrnes was a f o u n d i n g m e m b e r <>l the C i n c i n n a t i S o c i - Byrnes, Richard M.

ety of N a t u r a l History, a n d he served as its c u r a t o r of m i n e r a l o g y f r o m 1871 (1835-1892)
until 1880, w h e n h e w a s t h r i c e e l e c t e d president a n d , t h u s , served t h r e e years
i n that position. H e w a s a m e m b e r o f the society's C o m m i t t e e o n G e o l o g i c a l
N o m e n c l a t u r e (S. A. M i l l e r et al. 1879) a n d w a s a c o - a u t h o r of t h e e u l o g y of
C. B. D y e r . He w a s listed as a local fossil c o l l e c t o r by N i c k l e s (1936), a n d he
h a d a large c o l l e c t i o n of fossils, b u t w a s interested in m i n e r a l s , in terrestrial
and freshwater m o l l u s c s , i n plants, a n d m u c h m o r e ( A n o n . 1 8 7 6 , 1 8 7 8 , 1 8 9 2 ;
B y r n e s , C o t t o n , a n d L a n g d o n 1883; J o h n s o n 2002).

S. T. Carley, of B a n t a m , C l e r m o n t C o u n t y , O h i o , was an active m e m b e r Carley, S. T.

o f t h e W e s t e r n A c a d e m y o f N a t u r a l S c i e n c e s , b e g i n n i n g i n t h e 1830s, a n d
h e w a s o n e o f t h e s e v e n s u r v i v i n g m e m b e r s - o f t h e a c a d e m y w h e n its assets
w e r e d o n a t e d t o t h e C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y i n 1871 a n d t h e
a c a d e m y c e a s e d t o exist. I n a b o u t 1846 D a v i d D a l e O w e n e n l i s t e d t h e h e l p
o f t h e a c a d e m y t o i m p r o v e t h e t a x o n o m i c n o m e n c l a t u r e o f t h e region's
fossils; C a r l e y a n d U. P. J a m e s " w e r e a p p o i n t e d a c o m m i t t e e to p r e p a r e a
report o n t h e n a m i n g o f t h e Strophomena o f t h e C i n c i n n a t i b l u e l i m e -
stone. I n this t h e y said that t h e y h a d ' c a r e f u l l y e x a m i n e d a n d c o m p a r e d a
great n u m b e r o f s p e c i m e n s ' a n d w e r e 'satisfied t h a t , b y far, t o o m a n y s p e -
cies h a v e b e e n m a d e , ' t h u s a f f i r m i n g O w e n ' s c o n t e n t i o n . " " T h i s r e p o r t w a s
p u b l i s h e d in t h e Cincinnati Gazette, S e p t e m b e r 5, 1846" ( H e n d r i c k s o n
1947, 143). In 1849 t h e r e w a s p u b l i s h e d a c a t a l o g u e of f r e s h w a t e r m u s s e l s ,
as recognized by the a c a d e m y ; the p a m p h l e t was published anonymously,
b u t a c c o r d i n g t o H e n d r i c k s o n (1947), i t w a s l a r g e l y t h e w o r k o f C a r l e y , John
G o u l d A n t h o n y , U . P . J a m e s , a n d Joseph C l a r k ( b u t see J o h n s o n 2002).
Carley was one of the collectors whose s p e c i m e n s were used in the
o r i g i n a l d e s c r i p t i o n of t h e s p e c i e s of t r a c e fossils, Licrophycus flabellum S.
A . M i l l e r a n d D y e r , 1878a. T h e w e l l - k n o w n s p e c i e s o f a r t i c u l a t e b r a c h i o -
p o d s , Retrorsirostra carleyi, w a s n a m e d after S. T. C a r l e y ( A n o n . 1878; H e n -
d r i c k s o n 1947).

Appendix 2 261
Christy, David D a v i d C h r i s t y w a s a C i n c i n n a t i - a r e a g e o l o g i s t , anti-slavery writer, printer,
a n d n e w s p a p e r m a n . H i s g e o l o g i c a l letters w e r e p u b l i s h e d i n 1848, origi-
n a l l y in a n e w s p a p e r , t h e Cincinnati Gazette, b u t t h e n separately. In a r o u n d
1858 h i s c o l l e c t i o n w a s sold t o M i a m i University, O x f o r d , O h i o , for $2,200
( B e c k e r 1938) or, a c c o r d i n g to S h i d e l e r ([1952] 2002, 2), for $5000, " a n al-
m o s t u n h e a r d o f s u m for s u c h a p u r p o s e , i n t h o s e d a y s . " T h e s a m e c o l l e c -
tion m u s t h a v e b e e n a financial b e n e f i t to at least o n e o t h e r party, t o o ;
a g a i n a c c o r d i n g t o S h i d e l e r ([1952] 2 0 0 2 , 4 ) : " I h a v e m e n t i o n e d t h e C h r i s t y
c o l l e c t i o n , a c q u i r e d b y M i a m i University. S o m e years a g o w h e n t h e U n i -
versity o f C h i c a g o w a s c l e a n i n g h o u s e w e w e r e g i v e n s o m e o f their u n -
w a n t e d m a t e r i a l . I n c l u d e d w e r e fossils d i s t i n c t l y m a r k e d ' C h r i s t y C o l l e c -
t i o n . ' T h e C h r i s t y c o l l e c t i o n h a d d i s a p p e a r e d f r o m M i a m i w h e n that
i n s t i t u t i o n w a s c l o s e d b e t w e e n 1873 a n d 1885. C h i c a g o h a d b o u g h t the
c o l l e c t i o n f r o m J a m e s H a l l w h o o p e r a t e d o u t o f A l b a n y , N.Y. Just w h o
s w i p e d t h e c o l l e c t i o n a n d sold i t t o H a l l h a s n ' t b e e n d e t e r m i n e d . O t h e r
material from the C h r i s t y collection has c o m e back in similar ways"
( B e c k e r 1938; C h r i s t y 1848; M e r r i l l [1924] 1964; S h i d e l e r [1952] 2002).

The Cincinnati ( S e e T h e C i n c i n n a t i S o c i e t y o f N a t u r a l History)

M u s e u m of
Natural History

T h e Cincinnati " N o r m a l s c h o o l s " were institutions that trained teachers and would-be
Normal School teachers. T h e C i n c i n n a t i N o r m a l S c h o o l was established by the C i n c i n -
nati B o a r d o f E d u c a t i o n i n 1868 a n d h o u s e d i n o n e o f its s c h o o l b u i l d i n g s ;
i t w a s s u s p e n d e d shortly after t h e start o f t h e t w e n t i e t h c e n t u r y . John M i c k -
l e b o r o u g h (q.v.) w a s p r i n c i p a l o f t h e C i n c i n n a t i N o r m a l S c h o o l for s e v e n
y e a r s , f r o m 1878 u n t i l 1885 ( L a t h r o p 1900,1902).

The Cincinnati (See A n t h o n y , John G o u l d ; T h e Western A c a d e m y o f Natural Sciences)

Society for t h e
Promotion of
Useful K n o w l e d g e

The Cincinnati T h e C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y w a s f o u n d e d i n 1870 ( A n o n .

Society of 1878, 1902). I n 1871 t h e r e m a i n i n g m e m b e r s o f t h e W e s t e r n A c a d e m y o f
Natural History N a t u r a l S c i e n c e s d o n a t e d all t h e assets o f t h e a c a d e m y , i n c l u d i n g m o n e y ,
b o o k s , a n d t h e i r c o l l e c t i o n , t o t h e C i n c i n n a t i S o c i e t y o f N a t u r a l History.
Initially, t h e s o c i e t y r e n t e d r o o m s i n t h e b u i l d i n g o f t h e C i n c i n n a t i C o l -
l e g e . I n 1877 t h e s o c i e t y , u s i n g p a r t o f a b e q u e s t r e c e i v e d f r o m m e m b e r
C h a r l e s B o d m a n , a c q u i r e d its o w n b u i l d i n g a t t h e s o u t h e a s t c o r n e r o f A r c h
a n d B r o a d w a y . I n 1934 t h e s o c i e t y m o v e d t o n e w p r e m i s e s i n t h e b u i l d i n g
o f t h e O h i o M e c h a n i c s Institute o n C e n t r a l P a r k w a y , a n d i n 1957 r e l o c a t e d

262 Appendix 2
t o its o w n b u i l d i n g o n G i l b e r t A v e n u e , i n t h e s o u t h w e s t c o r n e r o f E d e n
Park ( A n o n . 1978). A t t h e s a m e t i m e , t h e n a m e o f t h e o r g a n i z a t i o n w a s
c h a n g e d t o t h e C i n c i n n a t i M u s e u m o f N a t u r a l H i s t o r y a n d , w i t h t h e ad-
dition of a p l a n e t a r i u m , the C i n c i n n a t i M u s e u m of N a t u r a l History a n d
P l a n e t a r i u m . W i t h t h e a b a n d o n m e n t o f t h e G i l b e r t A v e n u e facilities a n d
a b s o r p t i o n into t h e C i n c i n n a t i M u s e u m C e n t e r a t U n i o n T e r m i n a l i n t h e
1990s, w h a t w a s o r i g i n a l l y t h e C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y c e a s e d
to exist as a s e p a r a t e entity.

L i s t e d as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936). C o o k , W. E.

E l e c t e d a m e m b e r o f t h e C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y i n 1878 a n d Cooper, Edward M.

served a s c u r a t o r o f p a l e o n t o l o g y o f t h e s o c i e t y ( A n o n . 1 8 7 9 , 1 8 8 5 a ) .

E l i z a b e t h A . D a l v e , k n o w n t o h e r friends a s B e t t i n a , w a s a n illustrator a n d Dalve, Elizabeth A.

part-time m u s e u m assistant i n t h e D e p a r t m e n t o f G e o l o g y a t t h e U n i v e r s i t y
of C i n c i n n a t i . S h e p r e p a r e d a list of t h e stratigraphic o c c u r r e n c e s of t h e fos-
sils i n t h e t y p e - C i n c i n n a t i a n ( B r a n d t a n d D a v i s 2007; D a l v e 1948,1951).

Deiss was born in C o v i n g t o n , Kentucky, and graduated from M i a m i Uni- Deiss, Charles
versity, O x f o r d , O h i o , w h e r e h e w a s a s t u d e n t o f S h i d e l e r . H e w a s t h e h e a d Frederick
o f t h e I n d i a n a G e o l o g i c a l S u r v e y f r o m 1945 u n t i l t h e t i m e o f his d e a t h (1903-1959)
( B e c k e r 1938; C r o n e i s 1963).

L i s t e d as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936); he is p r e s u m a b l y t h e s a m e Dickhaut, Henry E.

p e r s o n a s H . E . D i c k h a u t , listed b y U l r i c h (1879b, 30).

T h e p r o p r i e t o r o f t h e W e s t e r n M u s e u m f r o m 1823 u n t i l s o m e t i m e i n t h e Dorfeuille, Joseph

late 1830s ( D r a k e a n d M a n s f i e l d 1827; T r o l l o p e 1832; K e l l o g g 1945).

Dr. D r a k e w a s t h e o n e o f t h e m o s t s i g n i f i c a n t f o r c e s i n t h e i n t e l l e c t u a l life Drake, Daniel

o f C i n c i n n a t i i n t h e first h a l f o f t h e n i n e t e e n t h c e n t u r y . H e w a s t h e m o t i - (1785-1852)
v a t i n g force b e h i n d b o t h t h e W e s t e r n M u s e u m a n d t h e W e s t e r n A c a d e m y
of Natural Sciences, and he was one of the "managers" of the Western
M u s e u m a t t h e t i m e o f its f o u n d i n g ( C a s t e r 1982; H e n d r i c k s o n 1947; J o h n -
son 2002; S. A. M i l l e r 1882a; [ S i l l i m a n ] 1819).

T h e D r y D r e d g e r s are a g r o u p o f a m a t e u r fossil c o l l e c t o r s f o u n d e d i n 1942 Dry D r e d g e r s

in t h e w a k e of a d i s c u s s i o n a n d field trip series s p o n s o r e d by t h e D e p a r t -

Appendix 2 263
 m e n t o f G e o l o g y a t t h e U n i v e r s i t y o f C i n c i n n a t i . T h e y are still g o i n g s t r o n g
(Brandt and Davis 2007; Dalve 1951).

Dyche, D. T. D. ( S e e c h a p t e r 2)

Dyer, Charles Brian ( S e e c h a p t e r 2)

(1806-1883)

Embree, Jesse O n e o f t h e " m a n a g e r s " o f t h e W e s t e r n M u s e u m a t t h e t i m e o f its f o u n d i n g ,

about 1820 ([Silliman] 1819).

Faber, Charles ( S e e c h a p t e r 2)
(died 1930)

Fales, J. C. Professor J. C. Kales, of C e n t r e C o l l e g e . D a n v i l l e , K e n t u c k y , was the person

w h o p r o v i d e d t h e s p e c i m e n s on w h i c h Monticulipora falesi U. P. James, 1884
w a s b a s e d a n d after w h o m t h e s p e c i e s w a s n a m e d (U. P. James 1884b).

Fenton, Carroll Lane W a s t h e first F a b e r C u r a t o r i n t h e D e p a r t m e n t o f G e o l o g y a t t h e University

o f C i n c i n n a t i ( C a s t e r 1982).

Findlay, James O n e o f t h e " m a n a g e r s " o f t h e W e s t e r n M u s e u m a t the t i m e o f its f o u n d i n g

( [ S i l l i m a n ] 1819).

Foerste, A u g u s t A u g u s t Frederick Foerste was a long-time h i g h school teacher in D a y t o n ,

F. (1862-1936) O h i o . H e w a s o n e o f f o r e m o s t w o r k e r s o n fossil n a u t i l o i d c e p h a l o p o d s . I n
a d d i t i o n , h e w o r k e d e x t e n s i v e l y o n t h e fossils a n d stratigraphy o f t h e t y p e -
C i n c i n n a t i a n . His collection w e n t to the U.S. National M u s e u m in Wash-
i n g t o n , D . C . I t w a s F o e r s t e w h o n a m e d t h e s p e c i e s o f trilobite n o w k n o w n
as Flexicalymene meeki. T h e r e is e x t e n s i v e b i o g r a p h i c a l m a t e r i a l on Foerste
a v a i l a b l e (Bassler 1937, 1947; B e c k e r 1938; B r a n d t a n d D a v i s 2007; C a s t e r
1 9 5 1 , 1 9 8 1 , 1 9 8 2 ; H o l l a n d 2000; S a n d y 1994; S h i d e l e r [1952] 2002).

Foster, W. B. D o n a t e d a " v a l u a b l e c o l l e c t i o n o f fossils a n d m i n e r a l s " t o t h e C i n c i n n a t i

S o c i e t y o f N a t u r a l H i s t o r y i n S e p t e m b e r o f 1878. I n a b o u t 1869 S c h u c h e r t ' s
father t o o k t h e y o u n g lad t o v i s i t " . . . W i l l i a m Foster's g e o l o g i c a l m u s e u m
which opened to me an u n k n o w n world" (Schuchert, quoted by Becker
1938,193); this c o u l d h a v e b e e n W . B . Foster ( A n o n . 1879; B e c k e r 1938).

264 Appendix 2
T o w a r d t h e e n d o f its e x i s t e n c e , t h e W e s t e r n M u s e u m i n C i n c i n n a t i w a s Franks' M u s e u m
best k n o w n as a c h a m b e r of horrors; in that b u s i n e s s it h a d a rival in t h e
form o f F r a n k s ' M u s e u m ( T u c k e r 1965, 32).

L a w y e r ; listed as a l o c a l c o l l e c t o r by S h i d e l e r ([1952] 2002). Fulton, Robert

L i s t e d as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936). Gault, W m .

G e o r g e G r a h a m was one of the original m e m b e r s of the Western A c a d e m y Graham, G e o r g e

o f N a t u r a l S c i e n c e s i n 1835, a n d h e w a s o n e o f t h e s e v e n s u r v i v i n g m e m b e r s
o f the a c a d e m y w h e n its assets w e r e d o n a t e d t o t h e C i n c i n n a t i S o c i e t y o f (1798-1881)
N a t u r a l History i n 1871. H i s c o l l e c t i o n o f fossils, s h e l l s , a n d p l a n t s , u n f o r -
tunately, w a s d e s t r o y e d in a fire ( A n o n . 1878; H e n d r i c k s o n 1947; J a m e s ,
H o w e , a n d N o r t o n 1881).

A u t h o r of the s p e c i e s Nereidavus varians G r i n n e l l , 1877, w h i c h w a s b a s e d on Grinnell,

s c o l e c o d o n t s ; A l b e r t G a l l a t i n W e t h e r b y h a d sent G r i n n e l l s p e c i m e n s o f t h e s e G e o r g e Bird
jaws o f a n n e l i d w o r m s from t h e t y p e - C i n c i n n a t i a n ( K n e l l , pers. c o m m . ) .

H e n r i M i l n e - E d w a r d s a n d Jules H a i m e d e s c r i b e d t h e t y p e - C i n c i n n a t i a n Haime, Jules

bryozoans now known as Parvohallopora rugosa and Dekayia aspera
( C u f f e y , D a v i s , a n d U t g a a r d 2002).

Mrs. M. P. Haines, of R i c h m o n d , Indiana, collected one of the specimens Haines, M. P.

of t h e s p e c i e s of starfish, Palaeasterina approximate S. A. M i l l e r a n d D y e r ,
1878a.

Listed a s curator o f p a l e o n t o l o g y o f t h e C i n c i n n a t i S o c i e t y o f N a t u r a l History Hall, J. H.

( A n o n . 1876); a l m o s t c e r t a i n l y t h e s a m e p e r s o n a s John W . H a l l , Jr., listed
below.

James H a l l p r o p o s e d t h e r e s o l u t i o n for t h e e s t a b l i s h m e n t o f t h e W e s t e r n Hall, James

A c a d e m y o f N a t u r a l S c i e n c e s o n A p r i l 25, 1835 ( H e n d r i c k s o n 1947, 140).
T h i s i s a l m o s t c e r t a i n l y n o t t h e s a m e p e r s o n a s t h e J a m e s H a l l (1811-1898)
w h o w a s o n e o f t h e m o s t e m i n e n t p a l e o n t o l o g i s t s o f t h e U n i t e d States i n
t h e n i n e t e e n t h c e n t u r y . T h e latter h a i l e d f r o m N e w Y o r k , b u t d i d p a l e o n -
t o l o g i c a l w o r k all o v e r t h e c o u n t r y , i n c l u d i n g t h e C i n c i n n a t i r e g i o n , a n d
he had c o n n e c t i o n s with various m e m b e r s of the C i n c i n n a t i S c h o o l (Hall
1845,1861,1872a, 1872b, 1883a, 1883b; H a l l a n d W h i t f i e l d 1875). B i o g r a p h i c a l

Appendix 2 265
 i n f o r m a t i o n a b o u t J a m e s H a l l o f N e w Y o r k i s v o l u m i n o u s a n d far f l u n g
( B e c k e r 1938; C a s t e r 1951, 1982, 1985; C r o n e i s 1963; M e r r i l l [1924] 1964;
S h e r b o r n 1940; S h i d e l e r [1952] 2002).

Hall, John W., Jr. J. W. H a l l , Jr., is listed as c u r a t o r of p a l e o n t o l o g y of the C i n c i n n a t i S o c i e t y of

N a t u r a l History from 1874 to 1877, a n d as c o r r e s p o n d i n g secretary of t h e soci-
ety in 1877 a n d 1878 ( A n o n . 1878). L a t e r that year, he d o n a t e d s o m e fossils to
t h e s o c i e t y ( A n o n . 1879). H e also served t h e s o c i e t y a s curator o f m i n e r a l o g y
( A n o n . 1885b). H e c o - a u t h o r e d a p a p e r w i t h a n u m b e r o f m e m b e r s o f t h e
C i n c i n n a t i S c h o o l a n d o t h e r l o c a l fossil c o l l e c t o r s (S. A . M i l l e r e t al. 1879).
J. W. H a l l , of C o v i n g t o n , is listed as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936).

H a m m o n d , W. E. O f M a d i s o n , s o listed a s a l o c a l fossil c o l l e c t o r b y N i c k l e s (1936).

Harper, G e o r g e ( S e e c h a p t e r 2)
W. (1832-1918)

Harris, I. H. The collection of I. H. Harris, of Waynesville, O h i o , provided specimens

u p o n w h i c h a n u m b e r of species originally were based, including some
n a m e d after h i m (S. A . M i l l e r 1 8 7 8 , 1 8 8 0 , 1 8 8 1 , 1 8 8 2 b , 1882d, 1883b, 1884. A t
least p a r t o f his c o l l e c t i o n e n d e d u p i n t h e N a t i o n a l M u s e u m o f N a t u r a l
History, i n W a s h i n g t o n , D . C . ( S a n d y 1996; S h i d e l e r [1952] 2002). T h e " I . M .
H a r r i s " listed as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936) is probably the s a m e
p e r s o n a s " I . H . H a r r i s , " a s i s t h e " I . H a r r i s " o f B e c k e r (1938).

Harvey, J. W. J. W. H a r v e y b o u g h t t h e t y p e - s p e c i m e n of t h e starfish, Palaeasterina spe-

ciosa S . A . M i l l e r a n d D y e r , 1878 f r o m W . C . B a r n h a r t a n d , i n t u r n , sold i t
t o C . B . D y e r ( M i l l e r a n d D y e r 1878a).

Hayden, F. V. F e r d i n a n d V a n d e v e e r H a y d e n i s m o s t f a m o u s for his h a v i n g b e e n t h e leader

o f t h e U n i t e d States G e o l o g i c a l a n d G e o g r a p h i c a l S u r v e y o f the Territories,
one o f t h e g r e a t surveys o f t h e A m e r i c a n W e s t , w h i c h e m p l o y e d L e s q u e r e u x
(q.v.), a m o n g m a n y o t h e r n o t a b l e g e o l o g i s t s o f t h e day. I n 1879 t h e H a y d e n
S u r v e y w a s o n e o f t h r e e federal surveys c o n s o l i d a t e d t o b e c o m e the t h e n - n e w
U n i t e d States G e o l o g i c a l Survey. M e e k a n d W o r t h e n , a l o n g w i t h F . B . M e e k
(q.v.), p r o p o s e d t h e u s e o f t h e t e r m " C i n c i n n a t i G r o u p " for t h e b o d y o f rocks
in the C i n c i n n a t i r e g i o n that h i t h e r t o h a d b e e n referred to as the " H u d s o n
River G r o u p " ; t h e y thus p r o p o s e d t h e c o n c e p t o f t h e C i n c i n n a t i a n (Bartlett
1962; M e r r i l l [1924] 1964; M e e k a n d W o r t h e n 1865a; W i l m a r t h 1925).

266 Appendix 2
" A n d r e w H e r r l i n g e r , later an a t t o r n e y in C i n c i n n a t i " is listed as a l o c a l Herrlinger, A n d r e w
fossil c o l l e c t o r by N i c k l e s (1936).

It was Reverend Herzer w h o introduced E. O. U l r i c h , then seven years old, Herzer, Henry
t o t h e w o n d e r s o f fossil c o l l e c t i n g ( C r o n e i s 1963; U l r i c h 1891).

Dr. Hill was associated with the C i n c i n n a t i Society of Natural History in Hill, H. H.
t h e 1870s, i n c l u d i n g v a r i o u s t e r m s a s c u r a t o r o f a r c h a e o l o g y , c u r a t o r o f
c o n c h o l o g y , a n d l i b r a r i a n ( A n o n . 1876, 1878). He a l s o is listed as a l o c a l
fossil c o l l e c t o r by N i c k l e s (1936).

Professor R . H . H o l b r o o k , o f t h e N a t i o n a l N o r m a l U n i v e r s i t y , L e b a n o n , Holbrook, R. H.
W a r r e n C o u n t y , O h i o , l e n t t h e m a t e r i a l u p o n w h i c h Stromatopora sub-
cylindrica o r i g i n a l l y w a s b a s e d (U. P. J a m e s 1884a).

L . M . H o s e a w a s c o - p r o p r i e t o r a n d c o - e d i t o r o f t h e Cincinnati Quarterly Hosea, L. M.

Journal of Science, a l o n g w i t h S. A. M i l l e r , d u r i n g its s e c o n d y e a r of p u b l i -
c a t i o n , 1875, a c c o r d i n g t o t h e c o v e r s o f t h e j o u r n a l .

Curator of comparative a n a t o m y in the C i n c i n n a t i S o c i e t y of Natural His- Howe, A. J.

tory ( A n o n . 1876).

D r . H u n t e r c o l l e c t e d o n e o f t h e s p e c i m e n s o n w h i c h t h e s p e c i e s Conularia Hunter, W. H. H.
formosa w a s b a s e d ( M i l l e r a n d D y e r 1878a).

T h o m a s H e n r y H u x l e y was o n e of the foremost naturalists in G r e a t Britain Huxley, T. H.

during the second half of the nineteenth century, and he was a vigorous
p r o p o n e n t o f t h e idea o f o r g a n i c e v o l u t i o n . H u x l e y visited C i n c i n n a t i o n
a t least t w o o c c a s i o n s , b u t h e o n l y s t a y e d o v e r n i g h t e a c h t i m e , a n d h e d i d
n o t give a n y s p e e c h e s n o r m a k e a n y f i e l d trips (Kritsky, p e r s . c o m m . ) .

James, Joseph
( S e e c h a p t e r 2)
Francis (1857-1897)

James, Uriah
( S e e c h a p t e r 2)
Pierson (1811-1889)

Appendix 2 267
Kemper, Willis L a w y e r ; listed as a l o c a l c o l l e c t o r by S h i d e l e r ([1952] 2002).

Lathrop, J. P. D o n a t e d a c o l l e c t i o n o f fossils t o t h e C i n c i n n a t i S o c i e t y o f N a t u r a l History

( A n o n . 1882).

Lesquereux, Leo ( C h a r l e s ) L e o L e s q u e r e u x w a s b o r n i n S w i t z e r l a n d i n 1806 b u t m o v e d t o

(1806-1889) t h e U n i t e d States i n 1848. H e b e c a m e , p e r h a p s , t h e f o r e m o s t p a l e o b o t a n i s t
o f his d a y i n t h e U n i t e d States. H e w o r k e d for a n u m b e r o f t h e state g e o l o g i -
c a l s u r v e y s , i n c l u d i n g t h e Illinois s u r v e y o f A . H . W o r t h e n (q.v.), a n d also
w o r k e d for t h e g r e a t s u r v e y o f t h 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 y d e n
(q.v.). L e s q u e r e u x a u t h o r e d t w o p a p e r s o n w h a t w e r e t h e n c o n s i d e r e d t o b e
l a n d p l a n t s i n t h e r o c k s o f t h e t y p e - C i n c i n n a t i a n . H e d i e d i n 1889, a n d
E d w a r d O r t o n (q.v.) w a s o n e o f h i s p a l l b e a r e r s ( B a r t l e t t 1962; M e r r i l l [1924]
1964; L e s q u e r e u x 1 8 7 4 , 1 8 7 8 , 2006; T r i t t 2006).

Letton's M u s e u m M e s s r s . L e t t o n a n d W i l l e t f o u n d e d a m u s e u m i n C i n c i n n a t i i n 1818just
a b o u t t h e s a m e t i m e t h a t t h e W e s t e r n M u s e u m w a s initiated b y D r . D a n i e l
D r a k e a n d his c o l l e a g u e s . I n 1826 R a l p h L e t t o n w a s t h e proprietor, a n d the
m u s e u m o c c u p i e d t w o storeys o f a b u i l d i n g a t t h e c o r n e r o f M a i n a n d
F o u r t h Streets. E x h i b i t e d there w e r e birds, m a m m a l s , minerals, shells,
fossils ( i n c l u d i n g f i f t y m a m m o t h b o n e s ) , a n t i q u i t i e s , a n d , a s w a s c o m m o n
i n t h o s e d a y s , w a x f i g u r e s . ( D r a k e a n d M a n s f i e l d 1827).

Lindahl, Josua D r . ( J o h a n H a r a l d ) Josua L i n d a h l w a s b o r n a n d e d u c a t e d i n S w e d e n . H e

(1844-1912) w a s c u r a t o r o f t h e Illinois State M u s e u m o f N a t u r a l History a n d state g e o l o -
gist o f I l l i n o i s , a n d t h e n w a s d i r e c t o r o f t h e C i n c i n n a t i S o c i e t y o f N a t u r a l
History, 1 8 9 5 - 1 9 0 6 ( A n o n . 1912; L i n d a h l 1906; U l r i c h 1891).

Locke, John Dr. L o c k e was a c h e m i s t , educator, geologist, paleontologist, inventor of

(1792-1856) s c i e n t i f i c i n s t r u m e n t s , a n d m o r e ; for e x a m p l e , h e w a s o n e o f t h e earliest
p h o t o g r a p h e r s i n C i n c i n n a t i ( G a g e l 1998). A s a n assistant g e o l o g i s t i n t h e
M a t h e r Survey, the first geological survey of O h i o , he was the author of the
1838 r e p o r t o n t h e g e o l o g y o f s o u t h w e s t e r n O h i o ( L o c k e 1838). H e has b e e n
credited as the first local a m a t e u r to b e c o m e a professional geologist, and
he was the first person to r e c o g n i z e what is n o w called the C i n c i n n a t i
A r c h . He was o n e of the hosts w h e n C h a r l e s Lyell, probably the foremost
g e o l o g i s t o n E a r t h , visited t h e C i n c i n n a t i a r e a ( L y e l l 1845). T h e majority
o f D r . L o c k e ' s p a l e o n t o l o g i c a l c o n t r i b u t i o n s d e a l t w i t h l o c a l trilobites. B i o -
g r a p h i c a l m a t e r i a l s o n D r . L o c k e are v o l u m i n o u s a n d far f l u n g ( B a r t k y
2000; B e c k e r 1938; B r a n d t a n d D a v i s 2007; D e x t e r 1979; D r a k e a n d M a n s -
f i e l d 1827; H a n s e n a n d C o l l i n s 1 9 7 9 ; H e n d r i c k s o n 1947; M e r r i l l [1924] 1964;
S . A . M i l l e r 1882a; S h o t w e l l 1902; T r o l l o p e 1832; W i n c h e l l 1894).

268 Appendix 2
T h e E n g l i s h m a n , C h a r l e s Lyell, was o n e of the founders of the science of Lyell, Charles
g e o l o g y as we k n o w it today. In 1842 he a c c e p t e d an invitation f r o m t h e W e s t -
ern A c a d e m y o f N a t u r a l S c i e n c e s t o c o m e t o C i n c i n n a t i . His hosts w e r e John
L o c k e , R o b e r t B u c h a n a n , J . G . A n t h o n y , a n d Joseph C l a r k , a n d t h e y visited
localities i n t h e area, i n c l u d i n g B i g B o n e L i c k , K e n t u c k y , w o r l d f a m o u s for
its r e m a i n s of Ice A g e m a m m a l s ( H e n d r i c k s o n 1947; L y e l l 1845).

S i d n e y S m i t h L y o n w a s b o r n i n C i n c i n n a t i . H e w a s o n e o f t h e assistant Lyon, Sidney S.

g e o l o g i s t s i n t h e g e o l o g i c a l s u r v e y o f K e n t u c k y b y D a v i d D a l e O w e n (q.v.), (1808-1872)
a n d h e a u t h o r e d o r c o - a u t h o r e d a n u m b e r o f s c i e n t i f i c p a p e r s o f fossil
e c h i n o d e r m s o f I n d i a n a , K e n t u c k y , a n d O h i o (S. A . M i l l e r 1882a).

W. W. M a t h e r was c h i e f geologist of the first geological survey of O h i o Mather, William

(1836-1838). Assistant g e o l o g i s t s i n c l u d e d John L o c k e a n d C h a r l e s W h i t - W. (1804-1859)
tlesey ( M e r r i l l [1924] 1964; S. A. M i l l e r 1882a).

Of O x f o r d , so listed as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936). McCord, D. A.

Fielding Bradford M e e k , a l t h o u g h born in M a d i s o n , Indiana, was mostly M e e k , F. B.

associated with the S m i t h s o n i a n Institution in W a s h i n g t o n , D . C . He is (1817-1876)
w e l l k n o w n for his i n v o l v e m e n t w i t h t h e g r e a t g e o l o g i c a l s u r v e y s o f t h e
A m e r i c a n W e s t ( B a r t l e t t 1962), a n d h e a u t h o r e d m a n y p a l e o n t o l o g i c a l c o n -
t r i b u t i o n s . O f m o s t interest t o r e a d e r s o f this v o l u m e i s t h e fact t h a t h e w a s
t h e a u t h o r o f t h e 1873 O h i o G e o l o g i c a l S u r v e y r e p o r t o n fossils o f t h e t y p e -
C i n c i n n a t i a n ( M e e k 1873), a n d a c o l l e c t i o n o f h i s O h i o s p e c i m e n s w e n t t o
C o l u m b i a University (Sherborn 1940). Moreover, M e e k and Worthen
(1865a) p r o p o s e d t h e u s e o f t h e t e r m " C i n c i n n a t i G r o u p " for t h e b o d y o f
rocks i n t h e C i n c i n n a t i r e g i o n t h a t h i t h e r t o h a d b e e n referred t o t h e " H u d -
son River G r o u p " ; t h e y t h u s p r o p o s e d t h e c o n c e p t o f t h e C i n c i n n a t i a n . B i o -
g r a p h i c a l m a t e r i a l s o n F . B . M e e k are e x t e n s i v e ( B e c k e r i 9 3 8 ; B r a n d t a n d
D a v i s 2007; C a s t e r 1981,1982; C r o n e i s 1963; M e r r i l l [1924] 1964; S . A . M i l l e r
1882a; W h i t e 1 8 7 7 , 1 8 9 6 , 1 9 0 2 ) .

( S e e c h a p t e r 2) Mickleborough,
John

D r . M i l l e r c o l l e c t e d o n e o f t h e s p e c i m e n s o n w h i c h t h e s p e c i e s Conularia Miller, C. A.
formosa w a s b a s e d ( M i l l e r a n d D y e r 1878a). T h e l o c a l t r i l o b i t e , Ceraurus
milleranus, w a s n a m e d after h i m (S. A . M i l l e r a n d G u r l e y 1893). D r . M i l l e r
also w a s t h e a u t h o r o f a s h o r t p a p e r o n p r e s e n t - d a y u n i o n i d c l a m s ( C . A .
M i l l e r 1874; J o h n s o n 2002).

Appendix 2 269
Miller, Samuel ( S e e c h a p t e r 2)
A l m o n d (1837-1897)

Milne-Edwards, H e n r i M i l n e - E d w a r d s a n d Jules H a i m e d e s c r i b e d t h e C i n c i n n a t i a n b r y o -
Henri z o a n s n o w k n o w n as Parvohallopora rugosa a n d Dekayia aspera ( C u f f e y ,
D a v i s , a n d U t g a a r d 2002).

Misener, John L i s t e d as a c o l l e c t o r of fossils at R i c h m o n d , I n d i a n a , by Foerste (1917), w h o

n a m e d Conchopeltis miseneri a n d Conularia miseneri after h i m .

Misener, S. R. Of R i c h m o n d , I n d i a n a , so listed as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936).

Mohr, Paul M o h r w a s associated w i t h t h e C i n c i n n a t i S o c i e t y o f N a t u r a l History i n the

1870s (S. A. M i l l e r et al. 1879), a n d t h e c e p h a l o p o d Orthoceras mohri w a s
n a m e d after h i m (S. A . M i l l e r 1875b). M o h r also w a s involved w i t h F r e d e r i c k
B r a u n , a n o t h e r C i n c i n n a t i c o l l e c t o r , i n e x c a v a t i o n s for c r i n o i d s f r o m C a r -
b o n i f e r o u s rocks a t C r a w f o r d s v i l l e , I n d i a n a ( V a n S a n t a n d L a n e 1964).

M o o r e , Richard L i s t e d a s a l o c a l fossil c o l l e c t o r b y N i c k l e s (1936). H e w a s a n o r i g i n a l m e m -

B. (1815-1885) b e r o f t h e C i n c i n n a t i S o c i e t y o f N a t u r a l History, c u s t o d i a n 1873-1877,
t r u s t e e 1 8 7 6 - 1 8 7 8 , a n d p r e s i d e n t 1 8 7 7 - 1 8 7 8 ( A n o n . 1876, 1878; C o t t o n ,
B y r n e s , a n d H e i g h w a y 1885).

Newberry, J. S. John S t r o n g N e w b e r r y w a s b o r n i n C o n n e c t i c u t , b u t m o v e d t o O h i o a t the

(1822-1892) a g e o f t w o ; h e r e c e i v e d h i s m e d i c a l d e g r e e i n t h e C l e v e l a n d area i n 1848.
H i s c o n n e c t i o n t o t h e t y p e - C i n c i n n a t i a n lies i n t h e fact that h e h e a d e d t h e
S e c o n d G e o l o g i c a l S u r v e y o f O h i o (1869-1878); h e e m p l o y e d F . B . M e e k
(q.v.), a n d t h e O h i o S u r v e y p u b l i s h e d o n e o f t h e i m p o r t a n t c o n t r i b u t i o n s
t o t h e p a l e o n t o l o g y o f t h e t y p e - C i n c i n n a t i a n ( M e e k 1873). N e w b e r r y ' s 1874
paper, " O n the So-called L a n d Plants from the L o w e r Silurian of O h i o , "
w a s r e p u b l i s h e d t h e s a m e year, b u t w i t h o u t t h e f i g u r e s , in t h e Cincinnati
Quarterly Journal of Science, w h i c h w a s , of c o u r s e , e d i t e d by S. A. M i l l e r
( M e e k 1873; M e r r i l l [1924] 1964; N e w b e r r y 1874).

N e w t o n , A. J. Dr. N e w t o n , of R i c h m o n d , Indiana, collected one of the specimens of

Enoploura d e s c r i b e d by W e t h e r b y (1879a).

Nicholson, H. Alleyn A l t h o u g h h e did study C i n c i n n a t i a n b r y o z o a n s i n t h e 1870s a n d 1880s, N i c h -

(1844-1899) o l s o n , a professor at t h e U n i v e r s i t y of A b e r d e e n , S c o t l a n d , was n o t a m e m b e r

270 Appendix 2
o f the C i n c i n n a t i S c h o o l . I t w a s N i c h o l s o n w h o i n t r o d u c e d the study o f t h i n -
sections t o t h e study o f b r y o z o a n s ( C u f f e y , D a v i s , a n d U t g a a r d 2002). T h e
n a m e Ceramopora nicholsoni w a s p r o p o s e d for a s p e c i e s of b r y o z o a n by U. P.
James (1875).

( S e e c h a p t e r 2) Nickles, John M.
(1859-1945)

(See C i n c i n n a t i N o r m a l S c h o o l ) Normal School

G e o r g e O e h h a d a c o l l e c t i o n o f fossils that i n c l u d e d m a n y fine C i n c i n n a t i Oeh, George

crinoids, a n d Heterocrinus (locrinus) oehanus w a s n a m e d after h i m ( U l r i c h
1882).

O ' N e a l l w a s a fossil c o l l e c t o r f r o m L e b a n o n , O h i o ; t h e trilobite, Acidaspis O'Neall, J. Kelly

onealli, w a s n a m e d after h i m (S. A . M i l l e r 1875a). T h e s p e c i e s o f b r y o z o a n
n o w c a l l e d Parvohallopora onealli w a s n a m e d by U. P. J a m e s , w h o spelt
O ' N e a l l ' s m i d d l e n a m e " K e l l e y " ( U . P . J a m e s 1875).

T h e French paleontologist, Alcide d'Orbigny, described the C i n c i n n a t i a n d'Orbigny, Alcide

bryozoans Monticulipora mammulata, Parvohallopora ramosa, and Het-
erotrypa frondosa ( C u f f e y , D a v i s , a n d U t g a a r d 2002; d ' O r b i g n y 1850).

E d w a r d O r t o n was b o r n i n D e l a w a r e C o u n t y , N e w York, a n d his initial e d u c a - O r t o n , Edward

tion was in that state. He also w e n t to seminary, i n c l u d i n g a stint at t h e L a n e (1829-1899)
T h e o l o g i c a l S e m i n a r y in C i n c i n n a t i , a n d he was o r d a i n e d as a Presbyterian
minister. S o m e w h a t later, he b e c a m e professor of natural s c i e n c e at A n t i o c h
C o l l e g e , and then its president. In 1873, he was a p p o i n t e d president of a n e w
institution, the O h i o A g r i c u l t u r a l a n d M e c h a n i c a l C o l l e g e , and also professor
of g e o l o g y there; u n d e r his leadership, the c o l l e g e prospered and w a s r e n a m e d
t o b e c o m e T h e O h i o State University. I n 1869, O r t o n w a s o n e o f t w o principal
assistants t o John Strong N e w b e r r y a n d t h e S e c o n d G e o l o g i c a l S u r v e y o f O h i o .
In 1882, he s u c c e e d e d N e w b e r r y as state geologist of O h i o , a n d , as s u c h , he gave
e m p l o y m e n t to U l r i c h a n d to Foerste. O r t o n b e c a m e a nationally r e c o g n i z e d
scientific figure, a n d , in 1897, he was e l e c t e d president of the G e o l o g i c a l S o c i -
ety of A m e r i c a ( C r o n e i s 1963; H a n s e n a n d C o l l i n s 1979; M e r r i l l [1924] 1964).

E d w a r d O r t o n , Jr., w a s a p p o i n t e d state g e o l o g i s t o f O h i o u p o n t h e d e a t h O r t o n , Edward, Jr.

of his father in 1899 a n d s e r v e d in t h a t p o s i t i o n u n t i l h i s r e s i g n a t i o n in 1906
( H a n s e n a n d C o l l i n s 1979).

Appendix 2 271
O w e n , David Dale D a v i d D a l e O w e n , a l t h o u g h n o t a m e m b e r o f t h e C i n c i n n a t i S c h o o l , did
(1807-1860) m u c h work in the region, a n d , indeed, in m u c h of the A m e r i c a n Midwest.
He had c o n n e c t i o n s with the Western A c a d e m y of Natural S c i e n c e s , and
the C i n c i n n a t i a n bryozoan, Fistulipora oweni, was n a m e d after h i m
( B e c k e r 1938; C r o n e i s 1963; H e n d r i c k s o n 1947; U. P. J a m e s 1879c, 1884a;
M e r r i l l [1924] 1964; S. A. M i l l e r 1882a).

Patterson, W. J. W i l l i a m J. Patterson w a s listed as a l o c a l fossil c o l l e c t o r by W e t h e r b y (1879a)

a n d b y N i c k l e s (1936). T h e C i n c i n n a t i a n c r i n o i d , Glyptocrinus pattersoni,
w a s n a m e d after h i m (S. A . M i l l e r 1882b).

Plummer, John T. D r . P l u m m e r w a s t h e a u t h o r o f a n early p a p e r o n t h e g e o l o g y o f t h e area

a b o u t R i c h m o n d , I n d i a n a , i n w h i c h fossils are d i s c u s s e d a n d illustrated
( P l u m m e r 1843).

Probasco, Henry Henry Probasco was in the hardware business in C i n c i n n a t i . He was an

(1820-1902) o r i g i n a l m e m b e r o f t h e C i n c i n n a t i S o c i e t y o f N a t u r a l History, a n d his c o l -
l e c t i o n o f fossils c a m e t o t h e U n i v e r s i t y o f C i n c i n n a t i . I t w a s h e w h o c o m -
missioned the Tyler D a v i d s o n Fountain, w h i c h gives the n a m e to Fountain
S q u a r e i n d o w n t o w n C i n c i n n a t i . I n a r o u n d 1875 P r o b a s c o a p p a r e n t l y p r o -
posed the establishment of a " C i n c i n n a t i M u s e u m of Science and Art"
a l o n g t h e l i n e s o f t h e British M u s e u m ( A n o n . 1 8 7 5 , 1 8 7 8 ; C a s t e r 1951,1982;
C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y ; W r i t e r s ' P r o g r a m o f t h e W o r k Proj-
e c t s A d m i n i s t r a t i o n i n t h e S t a t e o f O h i o 1943).

Rafinesque, C. C o n s t a n t i n e S a m u e l R a f i n e s q u e - S c h m a l t z i s b e s t k n o w n t o fossil c o l l e c -
S. (1783-1840) tors b e c a u s e o f t h e c o m m o n C i n c i n n a t i a n b r a c h i o p o d , Rafinesquina H a l l
a n d C l a r k e , 1892. R a f i n e s q u e w a s b o r n i n T u r k e y , i n w h a t w a s t h e n C o n -
s t a n t i n o p l e , b u t h e t r a v e l e d w i d e l y , i n c l u d i n g t h e U n i t e d States, w h e r e h e
e v e n t u a l l y s e t t l e d . R a f i n e s q u e t a u g h t for a t i m e at T r a n s y l v a n i a C o l l e g e in
L e x i n g t o n , Kentucky, and he gave public lectures at the C i n c i n n a t i m u -
s e u m o f Joseph D o r f e u i l l e ((/.v.). H e s e e m s t o h a v e b e e n p r i m a r i l y a b o t a -
nist, b u t his interests w e r e v e r y far r e a c h i n g , a n d h e n a m e d m a n y g e n e r a
a n d s p e c i e s in a m u l t i t u d e of d i f f e r e n t t a x o n o m i c g r o u p s . He w a s a prolific
a u t h o r , b u t m a n y o f his w o r k s w e r e i n o b s c u r e p u b l i c a t i o n s o r e v e n self-
p u b l i s h e d , a n d m a n y are q u i t e rare. T h i s m a y b e p a r t o f t h e r e a s o n that his
s c i e n t i f i c w o r k i s n o t h i g h l y r e g a r d e d today. T h e r e i s e x t e n s i v e b i o g r a p h i c a l
m a t e r i a l on R a f i n e s q u e a v a i l a b l e ( C a s t e r 1982; J o h n s o n 2002; M e r r i l l [1924]
1964; S. A. M i l l e r 1882a; R a f i n e s q u e 1836).

Reinhardt, E. E . R e i n h a r d t c o l l e c t e d t h e t y p e - s p e c i m e n o f t h e c r i n o i d s p e c i e s Glyptocri-
nus angularis S. A. M i l l e r a n d D y e r , 1878a.

272 Appendix 2
J. M. Richardson, of 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 , found the original Richardson, J. M.
s p e c i m e n s of the c r i n o i d s p e c i e s Glyptocrinus richardsoni W e t h e r b y , 1880.

Dr. R i d d e l l , a f a m o u s botanist, w a s part of the f a c u l t y of the M e d i c a l D e p a r t - Riddell, John L.

m e n t o f C i n c i n n a t i C o l l e g e , a short-lived m e d i c a l c o l l e g e f o u n d e d b y D a n i e l
D r a k e (Writers' P r o g r a m o f t h e W o r k Projects A d m i n i s t r a t i o n i n t h e State o f
O h i o 1943). A b o u t t h e s a m e t i m e , Dr. R i d d e l l w a s a c u r a t o r o f t h e W e s t e r n
A c a d e m y of N a t u r a l S c i e n c e s , b e g i n n i n g in A p r i l of 1835 ( H e n d r i c k s o n 1947).

Carl L u d w i g R o e m i n g e r was born in G e r m a n y and earned a m e d i c a l degree Roeminger, Carl

(but w i t h a g e o l o g i c a l thesis!) at T u b i n g e n . At t h e t i m e of t h e 1848 r e v o l u -
tions, h e m o v e d t o C i n c i n n a t i , w h i c h , a t that t i m e , h a d a large G e r m a n -
s p e a k i n g p o p u l a t i o n , a n d h e p r a c t i c e d m e d i c i n e t h e r e for s o m e twenty-five
years. H e studied w h a t are n o w k n o w n t o b e fossil b r y o z o a n s a s spare t i m e
f r o m h i s m e d i c a l d u t i e s w o u l d allow. I t w a s h e w h o n a m e d Rhombotrypa
quadrata ( R o e m i n g e r , 1866). A b o u t that s a m e t i m e , h e m o v e d t o M i c h i g a n ,
w h e r e he s i m u l t a n e o u s l y c o n d u c t e d a m e d i c a l p r a c t i c e , w a s a professor of
g e o l o g y a t t h e University o f M i c h i g a n , a n d w a s M i c h i g a n ' s state g e o l o g i s t .
Roeminger's n a m e appears both with and without the u m l a u t over the " o " or
the " e " after t h e " o " ( C u f f e y , D a v i s , a n d U t g a a r d 2002; M e r r i l l [1924] 1964).

A p l u m b e r by trade, C h a r l e s S c h l e m m e r was an associate of E. O. Ulrich Schlemmer, Charh

a n d C h a r l e s S c h u c h e r t i n t h e i r e a r l y c o l l e c t i n g d a y s . S c h l e m m e r ' s interest (died 1933)
in fossils is said to h a v e b e e n a w a k e n e d l a r g e l y by C h a r l e s F a b e r , as w a s
that o f A s h e r m a n n , T w i t c h e l l , V a u p e l , a n d o t h e r s ( S h i d e l e r [1952] 2002).

( S e e c h a p t e r 2) Schuchert, Charle
(1858-1942)

Scott, w h o b e c a m e an e m i n e n t vertebrate paleontologist, was b o r n in C i n - Scott, William

c i n n a t i o n F e b . 12,1858. I n a r o u n d 1861 h i s f a m i l y d e p a r t e d C i n c i n n a t i for Berryman
g o o d a n d m o v e d t o C h i c a g o , a n d t h e n t o P r i n c e t o n , N e w Jersey. S c o t t ' s (1858-1947)
b i r t h p l a c e p r o b a b l y h a d little o r n o t h i n g t o d o w i t h his p a l e o n t o l o g i c c a r e e r
( C r o n e i s 1963; S i m p s o n 1984).

Dr. Scoville collected n e a r L e b a n o n , O h i o ; Orthis scovillei w a s n a m e d Scoville, S. S.

after h i m (S. A . M i l l e r 1882c).

S h a l e r was b o r n a n d reared i n n o r t h e r n K e n t u c k y . H e s t u d i e d u n d e r L o u i s Shaler, Nathaniel

A g a s s i z a n d spent t h e b u l k o f his c a r e e r a t H a r v a r d University. S h a l e r m u s t Southgate
(1841-1906)

Appendix 2 273
 h a v e b e e n a n e x t r a o r d i n a r y i n d i v i d u a l ; for e x a m p l e , f r o m 1874 t o 1880, h e
t a u g h t at H a r v a r d , w a s state g e o l o g i s t of K e n t u c k y , a n d w a s p r e s i d e n t of a
m i n i n g c o m p a n y i n M o n t a n a , all a t t h e s a m e t i m e . I n t h e u p p e r e c h e l o n o f
his profession, h e served a s p r e s i d e n t o f t h e G e o l o g i c a l S o c i e t y o f A m e r i c a i n
1895 a n d w a s a m e m b e r o f t h e N a t i o n a l A c a d e m y o f S c i e n c e s , b u t h e also was
a c o n t r i b u t o r to t h e Atlantic Monthly a n d to Scrihner's m a g a z i n e and w r o t e
five r o m a n t i c d r a m a s in b l a n k verse. A l t h o u g h S h a l e r did a u t h o r a p a p e r that
i n v o l v e d b r a c h i o p o d s o f t h e C i n c i n n a t i r e g i o n , h e w a s n o t really a m e m b e r
o f the C i n c i n n a t i S c h o o l ( B e c k e r 1938; C a s t e r 1951,1982; C r o n e i s 1963; H o b b s
1907; L i v i n g s t o n e 1987; S h a l e r 1876; S h i d e l e r [1952] 2002).

Shideler, W. H. W i l l i a m Henry Shideler was born in West M i d d l e t o w n , O h i o , and graduated

(1886-1958) i n 1907 f r o m M i a m i University, O x f o r d , O h i o . I m m e d i a t e l y u p o n receipt o f
his d o c t o r a t e a t C o r n e l l U n i v e r s i t y i n 1910, h e r e t u r n e d t o his a l m a mater,
w h e r e h e s p e n t t h e rest o f h i s life o n t h e faculty. H e w a s r e c o g n i z e d a s a n
e x c e p t i o n a l t e a c h e r , a n d , e v e n t h o u g h his p u b l i c a t i o n record w a s not e x t e n -
sive ( S h i d e l e r 1914,1916,1918,1939), his k n o w l e d g e of the l o c a l rocks a n d fos-
sils w a s extraordinary, a n d his i n f l u e n c e o n c o h o r t s o f students and c o l l e a g u e s
w a s p r o f o u n d . A l t h o u g h n o t really a m e m b e r o f t h e C i n c i n n a t i S c h o o l , h e
did k n o w s o m e o f t h e m p e r s o n a l l y a n d e v e n spent t i m e i n t h e f i e l d w i t h a t
least U l r i c h a n d F a b e r ( B e c k e r 1938; C a s t e r 1951,1961a, 1985; C r o n e i s 1963;
S h i d e l e r [1952] 2002).

Stevens, W. J. W . J . S t e v e n s , o f L e b a n o n , O h i o , c o l l e c t e d t h e t y p e - s p e c i m e n o f t h e star-
fish s p e c i e s Palaeaster simplex S. A. M i l l e r a n d D y e r , 1878a.

Trollope, Frances T h e m o t h e r o f British n o v e l i s t A n t h o n y T r o l l o p e . D u r i n g h e r stay i n C i n -

cinnati, she had s o m e association with the Western M u s e u m and with
Joseph D o r f e u i l l e . H e r r e f e r e n c e t o a " D r . L o c k " a p p a r e n t l y w a s t o John
L o c k e ( T r o l l o p e 1832).

T w e n h o f e l , William T w e n h o f e l w a s b o r n n e a r C o v i n g t o n , K e n t u c k y , just across t h e O h i o River

Henry (1875-1957)
from C i n c i n n a t i , a n d reared in the area. He taught in K e n t u c k y public
s c h o o l s for six years a n d e a r n e d a b a c h e l o r ' s d e g r e e from the N a t i o n a l Nor-
m a l University i n L e b a n o n , O h i o , i n 1904. H i s g e o l o g i c a l t r a i n i n g was a t Y a l e
University, w h e r e S c h u c h e r t t a u g h t . The b u l k o f T w e n h o f e l ' s c a r e e r w a s
s p e n t a t t h e U n i v e r s i t y o f W i s c o n s i n . H e served a s president o f t h e P a l e o n t o -
l o g i c a l S o c i e t y in 1931, b u t he is b e s t r e m e m b e r e d as o n e of the f o u n d e r s of
t h e f i e l d o f s e d i m e n t o l o g y ( B e c k e r 1938; C r o n e i s 1963; D o t t 2000).

Twitchell, G e o r g e G e o r g e T w i t c h e l l w a s a p h y s i c i a n a n d a m a t e u r fossil c o l l e c t o r w h o s e interest

B. (died 1933) in fossils is said to h a v e b e e n a w a k e n e d largely by C h a r l e s Faber, as w a s that

274 Appendix 2
of A s h e r m a n n , S c h l e m m e r , Vaupel, and others. He was elected to m e m b e r -
ship in the C i n c i n n a t i S o c i e t y of N a t u r a l History in 1885. H i s large c o l l e c t i o n
of type-Cincinnatian bryozoans, including thin-sections, was bequeathed to
the University o f C i n c i n n a t i . H e served a s c u r a t o r o f m i c r o s c o p y a t t h e s o c i e t y
and a u t h o r e d several scientific p a p e r s , a l t h o u g h n o t on fossils ( A n o n . 1885a;
C a s t e r 1982; S h i d e l e r [1952] 2002; T w i t c h e l l 1885,1887,1934).

( S e e c h a p t e r 2) Ulrich, E. O.
(1857-1944)

G e o r g e V a l l a n d i g h a m , o f C i n c i n n a t i , w a s a n a c t i v e c o l l e c t o r after w h o m Vallandigham,
Cyrtoceras vallandighami S. A. M i l l e r , 1874 w a s n a m e d . He a l s o c o l l e c t e d G e o r g e L.
o n e o f t h e s p e c i m e n s o n w h i c h t h e g e n u s o f c a r p o i d e c h i n o d e r m s , Enop-
loura, w a s b a s e d ( W e t h e r b y 1879a). T h i s i s c e r t a i n l y t h e G e o r g e L . V a l -
l a n d i n g h a m listed as a l o c a l fossil c o l l e c t o r by N i c k l e s (1936).

D e s c r i b e d t h e first-named t y p e - C i n c i n n a t i a n b r y o z o a n , Constellaria con- Van Cleve, J. W.

stellata, w h i c h w a s p u b l i s h e d f o r m a l l y b y D a n a (1846, 1849), V a n C l e v e
h a v i n g d i e d ( C u f f e y , D a v i s , a n d U t g a a r d 2002).

E r n s t V a u p e l w a s a c h i l d h o o d friend o f C h a r l e s S c h u c h e r t a n d o f John M . Vaupel, Ernst H.

N i c k l e s . I n 1885 V a u p e l , a l o n g w i t h S c h u c h e r t a n d C h a r l e s Faber, w a s p r o - (died 1942)
p o s e d for m e m b e r s h i p in t h e C i n c i n n a t i S o c i e t y of N a t u r a l History. A shirt
m a k e r by profession, V a u p e l a c c u m u l a t e d a large c o l l e c t i o n of l o c a l fossils
and d o n a t e d i t t o Y a l e University, w h e r e S c h u c h e r t t a u g h t ; h e t h e n a m a s s e d
a s e c o n d c o l l e c t i o n a n d g a v e it to t h e U n i v e r s i t y of C i n c i n n a t i ( A n o n . 1885a;
C a s t e r 1981,1982; N i c k l e s 1936; S h i d e l e r [1952] 2002; W e t h e r b y 1881).

John W a r d e r is r e c o r d e d as h a v i n g g i v e n s o m e fossils to the W e s t e r n A c a d e m y Warder, John

of N a t u r a l S c i e n c e s on M a y 1, 1838, a n d he a u t h o r e d a p a p e r on trilobites A s t o n (1812-1883)
(although not t y p e - C i n c i n n a t i a n ) p u b l i s h e d t h e s a m e year. H e w a s president
of the C i n c i n n a t i S o c i e t y of N a t u r a l History f r o m its f o u n d i n g in 1870 u n t i l
April 1875 ( M o o r e , H e i g h w a y , a n d H o w e 1883; H e n d r i c k s o n 1947).

O n e o f t h e o r i g i n a l m e m b e r s o f t h e D r y D r e d g e r s i n 1942 ( F e l t o n , pers. Weichold, Charles

c o m m . ) . K e n n e t h C a s t e r u s e d t o refer t o c e r t a i n r i n g - s h a p e d b r y o z o a n
colonies in the t y p e - C i n c i n n a t i a n as " W e i c h o l d R i n g s " or " W e i c h o l d
D o u g h n u t s , " after C h a r l e s W e i c h o l d .

Appendix 2 275
Welch, L. B. Dr. W e l c h practiced 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 . T h e
original specimens of Glyptocrinus richardsoni Wetherby, 1880, were in
W e l c h ' s c o l l e c t i o n ( W e t h e r b y 1880, 246; F o e r s t e 1893a, 1893b).

Wessels, Charles C h a r l e s W e s s e l s f o u n d t h e s p e c i m e n of Lepidocoleus jamesi d e s c r i b e d by

F a b e r , w h e n h e n a m e d t h e g e n u s Lepidocoleus ( F a b e r 1886,17).

The W e s t e r n T h e Western A c a d e m y of Natural Sciences was founded by Dr. Daniel

Academy of D r a k e i n 1835. I n 1840 t h e N a t u r a l S c i e n c e S e c t i o n o f t h e C i n c i n n a t i S o -
Natural Sciences c i e t y f o r the P r o m o t i o n o f U s e f u l K n o w l e d g e , o f w h i c h John G o u l d A n -
(1835-1871) t h o n y w a s s e c r e t a r y , m e r g e d w i t h t h e a c a d e m y . I n 1871 t h e r e m a i n i n g
m e m b e r s o f the W e s t e r n A c a d e m y o f N a t u r a l S c i e n c e s d o n a t e d all the
assets o f t h e a c a d e m y , i n c l u d i n g m o n e y , b o o k s , a n d their c o l l e c t i o n , t o the
C i n c i n n a t i S o c i e t y o f N a t u r a l History, w h i c h h a d b e e n f o u n d e d t h e previ-
o u s y e a r ( A n o n . 1 8 7 8 , 5 ; H e n d r i c k s o n 1947).

The W e s t e r n A s o c i e t y w a s f o r m e d in 1819 or shortly b e f o r e by a g r o u p , i n c l u d i n g D a n i e l

Museum D r a k e (q.v.), t o e s t a b l i s h a m u s e u m i n C i n c i n n a t i . T h e W e s t e r n M u s e u m
(1820-1867) o p e n e d o n J u n e 10,1820, w i t h D r . R o b e r t B e s t a s t h e p r i n c i p a l curator. John
J a m e s A u d u b o n w a s a n e a r l y e m p l o y e e . I n 1823 t h e W e s t e r n M u s e u m S o -
c i e t y t r a n s f e r r e d o w n e r s h i p to Joseph D o r f e u i l l e (q.v.). In 1839 he sold t h e
m u s e u m a n d m o s t o f its e x h i b i t s a n d c o l l e c t i o n s t o a l o c a l g r o u p a n d took
t h e rest w i t h h i m t o N e w Y o r k a n d o p e n e d a n e w e s t a b l i s h m e n t . I n 1867
t h e e x h i b i t s a n d c o l l e c t i o n s i n C i n c i n n a t i w e r e p u t u p for p u b l i c a u c t i o n ;
their w h e r e a b o u t s are u n k n o w n today. ( C o n t r a r y t o t h e t y p o g r a p h i c error
i n C a s t e r [1982, 23], i t w a s n o t t h e c o l l e c t i o n s o f t h e W e s t e r n M u s e u m , b u t ,
rather, t h o s e o f t h e W e s t e r n A c a d e m y o f N a t u r a l S c i e n c e s that w e r e t a k e n
o v e r b y t h e C i n c i n n a t i S o c i e t y o f N a t u r a l H i s t o r y i n 1871.) ( A n o n . 1878, 5 ;
D r a k e a n d M a n s f i e l d 1 8 2 7 , 4 4 - 4 6 ; H a r t - D a v i s 2004, 39; J o h n s o n 2002; K e l -
l o g g 1945; [ S i l l i m a n ] 1819; T u c k e r 1 9 6 5 , 1 9 6 7 ) .

Wetherby, Albert ( S e e c h a p t e r 2)
Gallatin (1833-1902)

Whitfield, R. P. R. P. W h i t f i e l d s e r v e d as a p a l e o n t o l o g i s t w i t h a n u m b e r of t h e federal a n d
state g e o l o g i c a l s u r v e y s i n t h e s e c o n d h a l f o f t h e n i n e t e e n t h c e n t u r y , c o m -
m o n l y associated with James Hall. W i t h respect to the t y p e - C i n c i n n a t i a n ,
h e w a s o n e o f t h e assistants t o t h e O h i o G e o l o g i c a l S u r v e y d u r i n g t h e
N e w b e r r y y e a r s a n d e x t e n d i n g o n into t h e E d w a r d O r t o n y e a r s , a n d s o m e
of his w o r k e v e n w a s p u b l i s h e d in t h e Journal of the Cincinnati Society of
Natural History, i n c l u d i n g d e s c r i p t i o n s of fossil p e l e c y p o d s o b t a i n e d f r o m
John M i c k l e b o r o u g h ( H a l l a n d W h i t f i e l d 1875; H a n s e n a n d C o l l i n s 1 9 7 9 ;

276 Appendix 2
M e r r i l l [1924] 1964; W h i t f i e l d 1878). T h e " R . P . W h i t e f i e l d " i n N i c k l e s
(1936) m u s t b e t h e s a m e p e r s o n .

A l t h o u g h p r i m a r i l y associated w i t h t h e C l e v e l a n d a r e a , C o l o n e l W h i t t l e s e y Whittlesey, Charles

m a d e a n a g r i c u l t u r a l survey o f H a m i l t o n C o u n t y , O h i o , i n 1844. Prior t o that,
h e h a d b e e n i n c h a r g e o f the t o p o g r a p h i c w o r k for W . W . M a t h e r ' s 1836-1838
g e o l o g i c a l survey o f O h i o . L a t e r h e w a s p a r t y t o a n a p p a r e n t l y v i c i o u s d i s p u t e
a s t o w h e t h e r N e w b e r r y o r W h i t t l e s e y s h o u l d h a v e b e c o m e state g e o l o g i s t o f
O h i o i n 1869; N e w b e r r y w a s s o a p p o i n t e d ( H a n s e n a n d C o l l i n s 1979; M e r r i l l
[1924] 1964).

(See Letton's M u s e u m ) Willet

Judge W i l l i a m W i l s o n , o f L e b a n o n , W a r r e n C o u n t y , O h i o , f o u n d t h e t y p e -
s p e c i m e n of Prioniodus dychei U. P. J a m e s , 1884, a n d t h e t h r e e s p e c i m e n s
on w h i c h Polygnathus wilsoni U. P. J a m e s , 1884 w a s b a s e d . H e , D y c h e , a n d
James w e r e f r i e n d s ( U . P. J a m e s 1884c, 1 4 7 - 1 4 8 ) .
Wilson, W m . W.

A m o s Henry W o r t h e n w a s b o r n in V e r m o n t in 1813, b u t , b e g i n n i n g in 1834,

t a u g h t s c h o o l i n C u m m i n s v i l l e , O h i o , n o w part o f C i n c i n n a t i . H e m o v e d t o
Warsaw, Illinois, in 1836, a n d d e v o t e d h i m s e l f to b u s i n e s s v e n t u r e s for a l i v i n g
a n d rocks and fossils in his spare t i m e . In 1855-1857 W o r t h e n w a s assistant in
W o r t h e n , A. H.
James Hall's g e o l o g i c a l survey of Iowa a n d , in 1858, w a s a p p o i n t e d t h e state
geologist o f Illinois. I n the latter post, h e hired M e e k a n d U l r i c h t o d o c u m e n t
the P a l e o z o i c fossils of that state. M e e k a n d W o r t h e n (1865a) p r o p o s e d the use
o f the term " C i n c i n n a t i G r o u p " for the b o d y o f rocks i n t h e C i n c i n n a t i r e g i o n
that h i t h e r t o h a d b e e n referred t o t h e " H u d s o n R i v e r G r o u p " ; t h e y t h u s p r o -
p o s e d t h e c o n c e p t o f t h e C i n c i n n a t i a n ( C r o n e i s 1963; M e e k a n d W o r t h e n
1865; M e r r i l l [1924] 1964; W i l m a r t h 1925).

Appendix 2 277
GLOSSARY

H o w t o p r o n o u n c e scientific t e r m s c a n b e a real b u g a b o o . I n t h e f o l l o w i n g
glossary, a n d e l s e w h e r e i n this v o l u m e , w e h a v e i n c l u d e d o c c a s i o n a l a d v i c e
on how to p r o n o u n c e terms. As you know, lexicographers have developed
a s c h e m e o f s y m b o l s t o i n d i c a t e h o w t h e y feel p a r t i c u l a r letters, s y l l a b l e s ,
and w o r d s s h o u l d b e p r o n o u n c e d . W e h a v e tried t o k e e p t h e u s e o f s u c h
s y m b o l s t o a m i n i m u m . W e h o p e t h a t , i n s o d o i n g , w e still h a v e m a n a g e d
to h e l p y o u p r o n o u n c e w o r d s in a w a y u s e f u l to y o u .
Unfortunately, n o t all scientific t e r m s are i n c l u d e d in d i c t i o n a r i e s , n o t
e v e n in the p r e m i e r d i c t i o n a r y of t h e E n g l i s h l a n g u a g e , The Oxford English
Dictionary ( S i m p s o n a n d W e i n e r 1989). T h e n a m e s of g e n e r a a n d s p e c i e s are
very rarely i n c l u d e d , e x c e p t i n s o m e s p e c i a l i z e d works. O n e that w e h a v e
found helpful is The Biologist's Handbook of Pronunciations (Jaeger 1960),
a n d there o t h e r similar works. A l a s ! S p a c e h e r e d o e s n o t p e r m i t us to list all
s u c h w o r k s , b u t t h e friendly professional librarians at t h e library of y o u r
c h o i c e c a n b e o f t r e m e n d o u s h e l p here.

symbols

absolute age (n.) g e o l o g i c a g e i n y e a r s , u s u a l l y d e t e r m i n e d b y r a d i o m e t r i c A

dating.
acritarch (n.) m i c r o f o s s i l o f o r g a n i c c o m p o s i t i o n r e p r e s e n t i n g u n k n o w n
b u t possibly a l g a l , s i n g l e - c e l l e d o r g a n i s m , p a r t i c u l a r l y a b u n d a n t i n
Paleozoic m a r i n e rocks.

279
 adapertural (adj.) i n c e p h a l o p o d s , t h e d i r e c t i o n t o w a r d t h e o p e n i n g o f the
shell (and, h e n c e , a w a y f r o m t h e a p e x o f the shell); s o m e w o r k e r s prefer
t o u s e t h e s y n o n y m o u s t e r m " a d o r a l " (cf.: a d a p i c a l ) .
adapical (adj.) i n c e p h a l o p o d s , t h e d i r e c t i o n t o w a r d t h e a p e x o f t h e shell
( a n d , h e n c e , a w a y f r o m t h e o p e n i n g o f t h e shell) (cf.: a d a p e r t u r a l ) .
adductor muscle (n.) o n e o f t h e m u s c l e s o f a b r a c h i o p o d o r a p e l e c y p o d
w h e r e b y t h e a n i m a l c l o s e s its s h e l l (cf: d i d u c t o r m u s c l e ; adjustor
muscle).
adjustor m u s c l e (n.) o n e o f t h e m u s c l e s o f a b r a c h i o p o d w h e r e b y t h e a n i -
m a l m o v e s its shell w i t h r e s p e c t t o t h e p e d i c l e (cf: a d d u c t o r m u s c l e ;
diductor muscle).
adoral (adj.) a d a p e r t u r a l (q.v.).
aegism (n.) "Associations for p r o t e c t i o n , either t h r o u g h c a m o u f l a g e of sur-
rounding vegetation, residence upon or within another animal, or even
transport w i t h i n the protective u m b r e l l a or aegis of a larger partner are very
c o m m o n i n the a n i m a l k i n g d o m . " " . . . this term collects u n d e r its b a n n e r
all those associations w h i c h are not principally c o n c e r n e d with food acqui-
sition. . . " ( M o r t o n 1989,10) (cf: c o m m e n s a l i s m ; m u t u a l i s m ; parasitism).
age (n.) g e o l o g i c t i m e u n i t , e q u i v a l e n t to stage in the time-stratigraphic clas-
sification; t h e h i e r a r c h y o f g e o l o g i c t i m e units, f r o m largest t o smallest
is: e o n , era, p e r i o d , e p o c h , a n d a g e (cf.: e o n , e p o c h , era, period).
aglaspids (n.) a r t h r o p o d s w i t h a h e a d s h i e l d a n d s e g m e n t e d b o d y , possibly
related t o h o r s e s h o e c r a b s a n d e u r y p t e r i d s ("sea s c o r p i o n s " ) , restricted
to C a m b r i a n and Ordovician.
anterior (adj., n.) o n o r t o w a r d t h e f r o n t o f t h e a n i m a l . ( N o t e that w h a t i s
functionally anterior in a given a n i m a l may not correspond to the ana-
t o m i c a l anterior. F o r e x a m p l e , i n h u m a n s , t h e h e a d i s a n a t o m i c a l l y
a n t e r i o r , b u t , a s y o u w a l k a r o u n d , y o u r h e a d i s a t t h e top o f 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 a l l y " d o r s a l " ; cf: distal; d o r s a l ; lateral; m e d i a l ;
p o s t e r i o r ; p r o x i m a l ; ventral.)
Aplacophora (n.) a class o f p h y l u m M o l l u s c a ; p r e s e n t - d a y a p l a c o p h o r a n s
that l a c k h a r d parts (from t h e G r e e k " a " + " p l a x , p l a k o s " + " p h o r e u s , "
m e a n i n g " n o plate b e a r e r " ) .
aragonitic (adj.) composed of the mineral aragonite (cf: calcareous,
calcific).
autoecology (n.) e c o l o g y of a s i n g l e s p e c i e s of o r g a n i s m s in a g i v e n t i m e
a n d p l a c e (cf.: s y n e c o l o g y ) .

B ball and pillow structure (n.) s e d i m e n t a r y s t r u c t u r e c h a r a c t e r i z e d b y b a i l -

o r p i l l o w - s h a p e d m a s s e s o f s a n d s t o n e o r l i m e s t o n e , often e m b e d d e d
in m a t r i x of a d i f f e r e n t l i t h o l o g y .
benthic, benthonic (adj.) l i v i n g i n , o n , o r a s s o c i a t e d w i t h t h e b o t t o m o f a
b o d y o f w a t e r (cf: n e k t o n i c ; p l a n k t o n i c ) .
benthos (n.) c o l l e c t i v e t e r m for b e n t h i c o r g a n i s m s .
bilaterally symmetrical (adj.) said o f a n a n i m a l i n w h i c h t h e left a n d right
sides o f t h e a n i m a l are m i r r o r i m a g e s o f o n e a n o t h e r ; h u m a n s are bi-
laterally s y m m e t r i c a l (cf: p l a n e o f s y m m e t r y , s y m m e t r y ) .

280 Glossary
binomen (see: s p e c i e s n a m e ) .
bioclaustration (n.) t h e p r o c e s s w h e r e b y a s o f t - b o d i e d o r g a n i s m t h a t in-
fests a n o t h e r o r g a n i s m o r c o l o n y i s e m b e d d e d b y t h e skeletal g r o w t h
o f that o t h e r o r g a n i s m o r c o l o n y . T h e w o r d w a s c o i n e d for i n s t a n c e s
in w h i c h a p a r t i c u l a r parasitic o r g a n i s m , s u g g e s t e d to be a h y d r o i d or
a colonial ascidiacian tunicate, was engulfed by the bryozoan colony
on w h i c h it w a s g r o w i n g ; t h e result is a p a t t e r n of h o l e s c a l l e d Catel-
locaula vallata P a l m e r a n d W i l s o n , 1988.
biofacies (n.) c h a r a c t e r i s t i c s of a s e d i m e n t a r y rock b a s e d on fossil c o n t e n t
(cf.: facies, lithofacies).
bioherm (n.) a f e a t u r e that is e l e v a t e d a b o v e t h e sea floor a n d w a s p r o -
d u c e d by o r g a n i s m s ( D a v i s 2004, 283); a fine, u p s t a n d i n g e x a m p l e is a
c o r a l reef, (cf.: b i o s t r o m e ) .
bioimmuration (n.) u s u a l l y a n e x t e r n a l m o l d o f a s o f t - b o d i e d o r g a n i s m
that was o v e r g r o w n b y a c a l c a r e o u s o r g a n i s m , f o l l o w e d b y d e c a y o f t h e
soft-bodied organism.
biostratigraphic unit (n.) a b o d y of r o c k c h a r a c t e r i z e d by fossils of a par-
t i c u l a r s p e c i e s or g r o u p of s p e c i e s ; at a g i v e n l o c a l i t y , t h i s is r e p r e -
sented b y t h e t h i c k n e s s o f strata s o c h a r a c t e r i z e d , ( c f : b i o z o n e , lith-
o s t r a t i g r a p h i c u n i t , t i m e - s t r a t i g r a p h i c unit).
biostratigraphy (n.) study of d i s t r i b u t i o n of fossils in s e d i m e n t a r y r o c k s .
biostrome (n.) s h e e t - l i k e , n o n - m o u n d e d , a c c u m u l a t i o n o f d e n s e l y p a c k e d
or intergrown calcareous organisms such as corals or bryozoans.
biotal succession (n.) t h e P r i n c i p l e o f B i o t a l S u c c e s s i o n w a s d e v e l o p e d b y
the E n g l i s h g e o l o g i s t W i l l i a m S m i t h in the years shortly prior to 1800.
H e r e a l i z e d t h a t , a s o n e g o e s u p w a r d i n t h e r o c k r e c o r d , o n e suite o f
fossils is s u c c e e d e d by a n o t h e r , w h i c h is, in t u r n , s u c c e e d e d by a t h i r d ,
a n d so o n , in a p a r t i c u l a r order. T h i s p r i n c i p l e is t h e basis for b i o -
stratigraphy a n d , h e n c e , for relative d a t i n g o f r o c k s b y t h e u s e o f fossils
(cf: e c o l o g i c s u c c e s s i o n ; fossil s u c c e s s i o n ) .
biozone (n.) a b i o s t r a t i g r a p h i c u n i t c h a r a c t e r i z e d by fossils of a p a r t i c u l a r
s p e c i e s o r g r o u p o f s p e c i e s ; s o m e t i m e s c a l l e d a "fossil z o n e " . C o m -
m o n l y t h e w o r d " z o n e " i s u s e d for this c o n c e p t , b u t , b e c a u s e that w o r d
is u s e d in m o r e t h a n o n e s e n s e in g e o l o g y , it is p r e f e r a b l e n o t to u s e
the term "zone", except with a modifier to indicate w h i c h kind of z o n e
is i n t e n d e d , (cf: e p i b o l e ) .
bivalved (adj.) said of a shell that c o n s i s t s of t w o s i m i l a r l y s i z e d p i e c e s ,
e a c h o f w h i c h i s c a l l e d a v a l v e ; b r a c h i o p o d s a n d p e l e c y p o d s are bi-
v a l v e d (cf: u n i v a l v e d ; p s e u d o b i v a l v e d ) .
bivalve (n.) a l t e r n a t i v e n a m e for a p e l e c y p o d .
b o d y fossil (n.) o n e that i n v o l v e s t h e r e m a i n s o f a n o r g a n i s m o r r e p l i c a o f
t h o s e r e m a i n s (cf: t r a c e fossil; i c h n o f o s s i l ; Lebensspur).

calcareous (adj.) c o m p o s e d o f c a l c i u m c a r b o n a t e , w h e t h e r t h e m i n e r a l c
c a l c i t e or t h e m i n e r a l a r a g o n i t e , or b o t h (cf: a r a g o n i t i c ; c a l c i f i c ) ,
calcite (n.) a c a l c i u m c a r b o n a t e m i n e r a l
calcific (adj.) c o m p o s e d of the m i n e r a l c a l c i t e (cf: a r a g o n i t i c , c a l c a r e o u s ) .

Glossary 281
 cast (n.) a r e p l i c a of an o r g a n i s m m a d e f r o m a m o l d of that o r g a n i s m (cf:
mold).
Chaetognatha (n.) a s m a l l p h y l u m o f m a r i n e invertebrates l i v i n g m o s t l y
a s p l a n k t o n , c o m m o n l y c a l l e d " a r r o w w o r m s " ; fossils are k n o w n f r o m
t h e P e n n s y l v a n i a n , a n d p o s s i b l y t h e C a m b r i a n ( V a l e n t i n e 2004).
chitin (n.) a c o m p l e x o r g a n i c m a t e r i a l p r o d u c e d by a r t h r o p o d s , for ex-
ample, by insects; the comparable material produced by molluscs is
termed c o n c h i o l i n , although some people use the term "chitin" to
refer n o t o n l y t o c h i t i n , i n t h e strict s e n s e , b u t t o c o n c h i o l i n a n d other
such organic materials.
chitinozoans (n.) a g r o u p o f m a r i n e p l a n k t o n i c microfossils w i t h g e n e r a l l y
f l a s k - s h a p e d o r g a n i c w a l l s ; o f u n c e r t a i n affinities, b u t p r e s u m a b l y a n i -
m a l s ( J a n s o n i u s a n d J e n k i n s 1978).
Chordata (n.) a n i m a l p h y l u m c h a r a c t e r i z e d b y t h e n o t o c h o r d , i n c l u d e s
the vertebrates.
Cincinnati Arch (n.) a b r o a d u p w a r p i n g o f strata e x t e n d i n g f r o m K e n -
tucky through O h i o .
Cincinnatian (n.) g e o l o g i c a g e t e r m for t h e L a t e O r d o v i c i a n E p o c h i n
North America.
C i n c i n n a t i a n Series (n.) t i m e - s t r a t i g r a p h i c t e r m for t h e U p p e r O r d o v i -
cian in North America.
class (n.) t h e l e v e l i n t h e L i n n a e a n h i e r a r c h y b e l o w p h y l u m a n d a b o v e
order.
classification (see: t a x o n o m y ) .
coelom (n.) b o d y c a v i t y i n c e r t a i n k i n d s o f a n i m a l s f o r m e d b y o r w i t h i n
m e s o d e r m a l tissue; a l s o c a l l e d a e u c o e l o m . A c o e l o m is l i n e d w i t h a
tissue c a l l e d t h e p e r i t o n e u m .
commensalism (n.) a r e l a t i o n s h i p in w h i c h an a n i m a l lives a t t a c h e d to or
a s a t e n a n t o f a n o t h e r , a n d o r d i n a r i l y s h a r e s its f o o d , b u t d o e s n o t
c o n s u m e its h o s t ( d i s t i n g u i s h e d f r o m a parasite, w h i c h f e e d s o n t h e
b o d y o f its host) (cf: a e g i s m ; m u t u a l i s m ; p a r a s i t i s m ; p r e d a t i o n ; host;
symbiosis).
competition, competitive interaction (n.) interaction(s) b e t w e e n t w o o r
m o r e s p e c i e s i n w h i c h all s p e c i e s are i n h i b i t e d o r h a r m e d , u s u a l l y
b e c a u s e they seek the s a m e resource, such as food, w h i c h is in limited
supply.
compression (n.) a k i n d of p r e s e r v a t i o n in w h i c h t h e fossil s p e c i m e n is
flattened; a fossil p r o d u c e d by s u c h a p r o c e s s .
conchiolin (n.) a c o m p l e x o r g a n i c m a t e r i a l p r o d u c e d b y m o l l u s c s ; t h e
o u t e r m o s t layer o f t h e m o l l u s c a n shell c o n s i s t s o f c o n c h i o l i n , a s d o t h e
o p e r c u l a of s o m e snails; the c o m p a r a b l e material p r o d u c e d by arthro-
pods is t e r m e d chitin.
consumer (n.) a n i m a l i n t h e f o o d c h a i n a b o v e t h e level o f p r i m a r y p r o d u c -
tion; it feeds on other organisms or organic matter.
convergent evolution (n.) o r g a n i c e v o l u t i o n i n w h i c h c h a n g e s i n t w o o r
m o r e e v o l u t i o n a r y l i n e a g e s result i n o r g a n i s m s that l o o k a l i k e . For
e x a m p l e , p o r p o i s e s , w h i c h are m a m m a l s , l o o k like f i s h .
coprophagous (adj.) f e e d i n g u p o n d u n g .

282 Glossary
coprolite (n.) a s p e c i m e n of fossil e x c r e m e n t (from t h e G r e e k " k o p r o s " +
"lithos," m e a n i n g " d u n g " + "rock").
correlation (n.) p r o c e s s o f d e t e r m i n i n g t h e c o r r e s p o n d e n c e o f strata i n
different s e c t i o n s o r l o c a t i o n s .
coquina (n.) a k i n d o f s e d i m e n t a r y r o c k t h a t c o n s i s t s a l m o s t e x c l u s i v e l y o f
s h e l l s , s h e l l - f r a g m e n t s , o r b o t h (from t h e S p a n i s h for " l i t t l e s h e l l " ;
p r o n o u n c e d : ko-keen-uh).
c o t y p e s (n.) i n t a x o n o m y , w h e n t w o o r m o r e t y p e - s p e c i m e n s are d e s i g -
n a t e d for a s p e c i e s , a n d all o f t h e m are c o n s i d e r e d t o b e e q u a l l y r e p r e -
sentative o f t h e s p e c i e s , t h e y are c a l l e d c o t y p e s . S o m e t i m e s , i n f o r m e r
y e a r s , t h e t e r m " s y n t y p e " w a s u s e d (cf.: h o l o t y p e ; p a r a t y p e ) .
cyclicity (n.) in stratigraphy, t h e r e g u l a r r e p e t i t i o n of a p a t t e r n or o r d e r i n g
in s e d i m e n t a r y strata.

death assemblage (n.) fossils i n a s t r a t u m t h a t r e p r e s e n t r e m a i n s o f o r g a n - D

isms that d i e d a t s o m e t i m e b e f o r e final b u r i a l , a n d u s u a l l y a c c u m u -
lated f r o m v a r i o u s o r i g i n a l l o c a t i o n s [cf.: life a s s e m b l a g e ) .
dentition (n.) c o l l e c t i v e n a m e g i v e n t o t h e t e e t h a n d s o c k e t s a l o n g t h e
hingeline of an articulate brachiopod or p e l e c y p o d ; the teeth and
s o c k e t s s e r v e t o p r e v e n t t h e v a l v e s f r o m b e i n g t w i s t e d a p a r t by, for
e x a m p l e , a p r e d a t o r ; n o t e that t h e t e r m " d e n t i t i o n , " in this s e n s e , has
n o t h i n g to do with the "dentition" of a jawed vertebrate such as your-
self (cf.: s o c k e t ; t o o t h ) .
deposit feeding (n., adj.) i n g e s t i o n o f f o o d m a t e r i a l s l o c a t e d o n o r w i t h i n
t h e s e d i m e n t (cf: s u s p e n s i o n f e e d i n g ; filter f e e d e r ) .
diagenesis (n.) c h e m i c a l , p h y s i c a l , o r b i o l o g i c a l c h a n g e s o c c u r r i n g i n
s e d i m e n t s a n d r o c k s s u b s e q u e n t t o initial f o r m a t i o n a t relatively l o w
t e m p e r a t u r e s a n d p r e s s u r e s (cf: m e t a m o r p h i s m ) .
diagnosis (n.) a s p e c i a l k i n d of d e s c r i p t i o n t h a t tells h o w i n d i v i d u a l s of a
g i v e n t a x o n differ f r o m all o t h e r m e m b e r s o f taxa a t t h e s a m e level i n
t h e L i n n a e a n h i e r a r c h y . F o r e x a m p l e , t h e d i a g n o s i s o f a s p e c i e s tells
h o w t o differentiate i t f r o m all o t h e r s p e c i e s i n t h e s a m e g e n u s .
dimorphism (n.) t h e e x i s t e n c e i n a t a x o n o f t w o p h e n o t y p i c f o r m s ; t h e
term is u s e d e s p e c i a l l y in r e f e r e n c e to s e x u a l dimorphism (cf:
polymorphism).
dinoflagellates (n.) a g r o u p o f m a r i n e p h y t o p l a n k t o n o r g a n i s m s , e a c h o f
w h i c h has o r g a n i c w a l l s , t w o f l a g e l l a , a n d c h l o r o p l a s t s .
distal (adj.) a w a y f r o m t h e c e n t e r o f t h e a n i m a l ; for e x a m p l e , y o u r f i n g e r
tips are d i s t a l , w h e r e a s y o u r s h o u l d e r is p r o x i m a l (cf: a n t e r i o r ; d o r s a l ;
lateral; m e d i a l ; posterior; p r o x i m a l ; v e n t r a l ) .
d i v e r s i t y (n.) t h e n u m b e r of taxa of a p a r t i c u l a r level that o c c u r in a g i v e n
t i m e a n d p l a c e ; the t e r m , u n m o d i f i e d , g e n e r a l l y refers t o t h e n u m b e r o f
species, with the m e a n i n g s o f " g e n e r i c diversity," " f a m i l i a l diversity," a n d
s o o n , d e n o t i n g diversity a t p a r t i c u l a r h i g h e r t a x o n o m i c levels.
d o l o m i t e (n.) 1. a m i n e r a l c o m p o s e d of c a l c i u m m a g n e s i u m c a r b o n a t e ; 2.
a sedimentary rock that consists of the m i n e r a l dolomite, s o m e t i m e s
c a l l e d rock d o l o m i t e or d o l o s t o n e (cf: a r a g o n i t e , c a l c i t e ) .

Glossary 283
 dorsal (adj.) o n o r t o w a r d t h e top o f t h e a n i m a l ; t h e n o u n f o r m o f t h e
c o n c e p t is " d o r s u m . " ( N o t e t h a t w h a t is f u n c t i o n a l l y dorsal in a g i v e n
a n i m a l m a y n o t c o r r e s p o n d t o t h e a n a t o m i c a l dorsal. For e x a m p l e , i n
h u m a n s , the head is a n a t o m i c a l l y anterior, but, as you walk around,
your h e a d is at the top of your body and, h e n c e , is functionally "dor-
sal"; cf: anterior; distal; lateral; m e d i a l ; posterior; p r o x i m a l ; ventral.)
durophagous (adj.) said o f p r e d a t o r s t h a t c r u s h t h e shells o f their prey.

E e c o l o g y (n.) 1 . t h e s t u d y o f t h e e n v i r o n m e n t ; 2 . t h e e n v i r o n m e n t a l n e e d s
of o r g a n i s m s of a given taxon or g r o u p of taxa (cf: autecology;
synecology).
ecosystem (n.) all t h e o r g a n i s m s i n a g i v e n t i m e a n d p l a c e , a l o n g w i t h t h e
c h e m i c a l a n d p h y s i c a l a s p e c t s o f t h e e n v i r o n m e n t i n w h i c h t h e y live
in that t i m e and place.
Edenian (n.) t e r m for t h e l o w e r m o s t stage o f t h e C i n c i n n a t i a n S e r i e s , also
the earliest age of C i n c i n n a t i a n E p o c h ,
e n d o s y m b i o n t (n.) o r g a n i s m l i v i n g w i t h i n t h e b o d y o f a n o t h e r l i v i n g
organism.
eon (n.) l a r g e s t g e o l o g i c t i m e u n i t , a s i n P h a n e r o z o i c E o n , w h i c h c o m -
prises t h e P a l e o z o i c , M e s o z o i c , a n d C e n o z o i c E r a s ; t h e h i e r a r c h y o f
g e o l o g i c t i m e u n i t s , f r o m largest t o s m a l l e s t is: e o n , era, p e r i o d , e p o c h ,
a n d a g e (cf.: a g e , e p o c h , era, p e r i o d ) .
epibole (11.) a s t r a t i g r a p h i c interval c h a r a c t e r i z e d by a b u n d a n c e of a par-
t i c u l a r k i n d o f fossil; s o m e t i m e s c a l l e d a n a c m e z o n e (cf.: b i o z o n e ) .
epibyssate (adj.) said of p e l e c y p o d s in w h i c h t h e byssus is a t t a c h e d to objects
that are b e n e a t h t h e a c t u a l s e d i m e n t - w a t e r interface of the sea floor.
epicole (n.) a f o s s i l i z e d a n i m a l f o u n d a t t a c h e d t o t h e exterior o f a n o t h e r
f o s s i l i z e d a n i m a l , a n d i t i s n o t c l e a r w h e t h e r t h e latter w a s alive w h e n
t h e f o r m e r w a s a t t a c h e d (cf: e p i z o o n ; e p i f a u n a ) .
e p i c o n t i n e n t a l seas (n.) m o s t l y s h a l l o w w a t e r m a r i n e b o d i e s o f w a t e r ex-
t e n d i n g o v e r c o n t i n e n t a l c r u s t (syn., e p e i r i c seas).
epifauna (n.) a n i m a l s l i v i n g on a s u b s t r a t u m s u c h as t h e sea floor (cf: in-
fauna; e p i z o o n ; epicole).
epireliefs (n.) t r a c e fossils on t h e u p p e r s u r f a c e of a s t r a t u m , o c c u r r i n g as
depressions or raised structures.
e p i z o o n (pl., e p i z o a ; n.) an a n i m a l t h a t s p e n d s its life on or a t t a c h e d to t h e
e x t e r i o r o f a n o t h e r l i v i n g a n i m a l ( D a v i s e t al. 1 9 9 9 , 33; p r o n o u n c e d :
ep-pi-zo-un). In former years, the word w o u l d have b e e n written "epi-
z o o n " ; t h e t w o dots o v e r t h e s e c o n d " o " i s c a l l e d a d i a e r e s i s , a n d i t
i n d i c a t e s that t h e s e c o n d " o " i s i n a d i f f e r e n t syllable f r o m t h e f i r s t " o . "
S o m e people incorrectly use the words "epizoan" and "epizoans" as
t h e s i n g u l a r a n d p l u r a l , r e s p e c t i v e l y (cf: e p i c o l e ; e p i f a u n a ) .
epoch (n.) a s u b d i v i s i o n of a g e o l o g i c p e r i o d , e q u i v a l e n t to series in the t i m e -
stratigraphic c l a s s i f i c a t i o n ; t h e h i e r a r c h y o f g e o l o g i c t i m e u n i t s , f r o m
largest t o s m a l l e s t is: e o n , era, p e r i o d , e p o c h , a n d a g e (cf: a g e , e o n , era,
period).

284 Glossary
era (n.) a g e o l o g i c t i m e u n i t , as in t h e P a l e o z o i c E r a , w h i c h i n c l u d e s , a m o n g
others, t h e O r d o v i c i a n P e r i o d ; t h e h i e r a r c h y o f g e o l o g i c t i m e u n i t s , f r o m
largest t o s m a l l e s t is: e o n , era, p e r i o d , e p o c h , a n d a g e (cf.: a g e , e p o c h ,
e o n , period).
eustatic (adj.) g l o b a l , r e f e r r i n g t o c h a n g e s o f sea l e v e l .
e v e n t h o r i z o n or e v e n t b e d (n.) a s e d i m e n t a r y layer f o r m e d b y a short-
t e r m d e p o s i t i o n a l p r o c e s s or d i s t u r b a n c e , s u c h as a layer of volcanic-
a s h , or a s t o r m b e d .
exoskeleton (n.) shell o r c a r a p a c e t h a t e n c l o s e s t h e b o d y o f a n a n i m a l .
external mold (n.) a m o l d o f t h e e x t e r i o r o f a n o b j e c t (cf: i n t e r n a l m o l d ;
steinkern).

facies (n.) a n y c h a r a c t e r i s t i c s o f r o c k s u s e d for r e c o g n i t i o n o f lateral o r F

vertical variations, and usually reflecting the processes f o r m i n g the
r o c k (cf: b i o f a c i e s ; lithofacies).
facultative (adj.) refers to an i n s t a n c e of an o r g a n i s m t h a t is a p a r t y to an
a s s o c i a t i o n that m a y b e c o n v e n i e n t , b u t i s n o t r e q u i r e d for t h e s u r v i v a l
of t h e o r g a n i s m ; if, on t h e o t h e r h a n d , a g i v e n parasite is u n a b l e to
exist o u t s i d e of its p a r t i c u l a r a s s o c i a t i o n w i t h its h o s t , t h e a s s o c i a t i o n
is an o b l i g a t e o n e (q.v.).
f a m i l y (n.) t h e level i n t h e L i n n a e a n h i e r a r c h y b e l o w o r d e r a n d a b o v e
genus.
filter f e e d e r (n.) a n o r g a n i s m that gets its s u s t e n a n c e b y r e m o v i n g m i c r o -
o r g a n i s m s o r o t h e r s m a l l p a r t i c l e s o f o r g a n i c m a t t e r f r o m t h e water.
E x a m p l e s i n c l u d e b r a c h i o p o d s , b r y o z o a n s , c o r a l s , a n d c r i n o i d s (cf:
deposit feeding; suspension feeding).
food chain, food w e b (n.) a series o f o r g a n i s m s c o n n e c t e d b y f e e d i n g rela-
tionships s u c h as predators a n d prey, or g r a z e r s a n d v e g e t a t i o n , r e f l e c t i n g
t h e transfer o f e n e r g y f r o m p r i m a r y p r o d u c e r s t h r o u g h c o n s u m e r s .
formation (n.) a r o c k - u n i t h a v i n g d i s t i n c t i v e l i t h o l o g i c f e a t u r e s a n d m a p -
p a b l e o n a r e g i o n a l s c a l e o f g e o l o g i c m a p p i n g ( g e n e r a l l y , 1:24,000
scale) (cf: g r o u p ; m e m b e r ) .
forms (n., p l . ) a g e n e r a l t e r m u s e d for g r o u p s o f m o r p h o l o g i c a l l y d i s t i n c t
o r g a n i s m s that h a v e n o t b e e n s u f f i c i e n t l y s t u d i e d t o w a r r a n t t h e i r b e -
ing called "subspecies" or "varieties," both of w h i c h terms m a y have
formal taxonomic implications.
fossil r e c o r d (n.) t h e e v i d e n c e o f a n c i e n t life a s p r e s e r v e d i n t h e r o c k s o f
the Earth's crust.
fossil s u c c e s s i o n (n.) a series of fossil a s s e m b l a g e s o c c u r r i n g in a v e r t i c a l
stratigraphic s e q u e n c e ,
fossil z o n e see b i o z o n e ) .
fucoid (n.) an a r c h a i c n a m e for c e r t a i n k i n d s of Lebensspuren (= trace fos-
sils); it s t e m s f r o m t h e fact that it o n c e w a s t h o u g h t t h a t t h e y w e r e t h e
r e m a i n s o f a n c i e n t s e a w e e d s related t o t h e w e l l - k n o w n g r o u p o f present-
day s e a w e e d s , Fucus.

Glossary 285
generic n a m e (n.) t h e n a m e o f t h e g e n u s t o w h i c h a n o r g a n i s m b e l o n g s ;
it is t h e first w o r d of t h e s p e c i e s n a m e [cf.: s p e c i e s n a m e ; specific n a m e ;
trivial n a m e ) .
g e n u s (pl., g e n e r a ; n.) t h e l e v e l i n t h e L i n n a e a n h i e r a r c h y b e l o w f a m i l y
and above species.
grainstone (n.) a l i m e s t o n e c h a r a c t e r i z e d by f r a g m e n t s in g r a i n - t o - g r a i n
c o n t a c t , a n d less t h a n 1 p e r c e n t of interstitial m u d (cf.: p a c k s t o n e ) .
group (n.) a r o c k - s t r a t i g r a p h i c u n i t c o m p o s e d o f t w o o r m o r e f o r m a t i o n s .
growth-lines (n.) c o n c e n t r i c r i d g e s , g r o o v e s , or striations on a shell that
m a r k t h e s u c c e s s i v e p o s i t i o n s o f f o r m e r m a r g i n s o f t h e shell o r v a l v e .
guild (n.) a g r o u p of s p e c i e s i n d i v i d u a l s of w h i c h u s e food r e s o u r c e s in a
similar manner.

hardground (n.) a s e d i m e n t a r y b e d s u r f a c e t h a t w a s c e m e n t e d o r lithified

b e f o r e final b u r i a l ; it c o m m o n l y is e n c r u s t e d or b o r e d by o r g a n i s m s .
holotype (n.) in t a x o n o m y , a s i n g l e t y p e - s p e c i m e n d e s i g n a t e d as r e p r e s e n -
tative of a s p e c i e s (cf: c o t y p e ; p a r a t y p e ) .
homeomorph (n.) t h e result o f c o n v e r g e n t e v o l u t i o n i n w h i c h t w o o r m o r e
e v o l u t i o n a r y l i n e a g e s h a v e e v o l v e d s o that t h e o r g a n i s m s a t t h e e n d s
of these lineages look alike.
h o s t (n.) " a n a n i m a l o r p l a n t h a v i n g a parasite o r c o m m e n s a l h a b i t u a l l y
l i v i n g i n o r u p o n it" ( S i m p s o n a n d W e i n e r 1989,1336).
hyporeliefs (n.) t r a c e fossils o n t h e l o w e r s u r f a c e o f a s t r a t u m , o c c u r r i n g
as d e p r e s s i o n s or raised s t r u c t u r e s .

Iapetus Sea (n.) a n c i e n t o c e a n that o c c u p i e d t h e a p p r o x i m a t e p o s i t i o n o f

t h e p r e s e n t - d a y A t l a n t i c d u r i n g early P a l e o z o i c t i m e .
ichnofacies (n.) a s e d i m e n t a r y r o c k c h a r a c t e r i z e d b y p a r t i c u l a r t r a c e
fossils.
i c h n o f o s s i l (n.) a t r a c e fossil (q.v.); p r o n o u n c e d : Ik-no-fossil,
ichnology (n.) t h e s t u d y o f t r a c e fossils.
impression (n.) o u t l i n e o r i m p r i n t i n s e d i m e n t o r s e d i m e n t a r y r o c k o f a n
o r g a n i s m o r part o f a n o r g a n i s m .
incertae sedis (n.) o f u n k n o w n b i o l o g i c a l a f f i n i t i e s ( L a t i n , " i n c e r t u s " ,
" d o u b t f u l , u n s e t t l e d , u n c e r t a i n " , + " s e d e s , sedis", " p l a c e " ) .
incolent (n.) a n a n i m a l t h a t lives i n o r o c c u p i e s t h e shell o f a n o t h e r a n i -
m a l after that o t h e r a n i m a l h a s d i e d ; for e x a m p l e , a trilobite that has
f o u n d shelter i n a n e m p t y c e p h a l o p o d shell o n t h e sea f l o o r . ( A c c o r d -
i n g t o t h e O x f o r d E n g l i s h D i c t i o n a r y [ S i m p s o n a n d W e i n e r 1989],
" i n c o l e n t " is an o b s o l e t e a n d rare w o r d that d a t e s f r o m 1597, w h e n it
m e a n t s i m p l y "an i n h a b i t a n t , " f r o m t h e L a t i n " i n c o l o , i n c o l e r e , " m e a n -
i n g " t o i n h a b i t . " ) (cf.: i n q u i l i n e ) .
infauna (n.) t h e o r g a n i s m s that live b e n e a t h t h e s e d i m e n t - w a t e r i n t e r f a c e ;
t h e a d j e c t i v a l f o r m is " i n f a u n a l " (cf: e p i f a u n a ) .
i n q u i l i n e (n.) a n a n i m a l t h a t lives i n t h e n e s t o r a b o d e o f a n o t h e r (cf:
incolent).

Glossary
internal m o l d (n.) a m o l d o f t h e interior o f a n o b j e c t ; t h e i n t e r n a l m o l d o f
a shell s o m e t i m e s is c a l l e d a s t e i n k e r n (q.v.) (cf.: e x t e r n a l m o l d ) ,
isochronous (adj.) e q u i v a l e n t i n a g e .

K-bentonite (n.) a p o t a s s i u m - r i c h c l a y m i n e r a l f o r m e d by c h e m i c a l altera- K

tion of v o l c a n i c ash d e p o s i t e d in salt water,
kingdom (n.) t h e l e v e l i n t h e L i n n a e a n h i e r a r c h y a b o v e p h y l u m .

l a g d e p o s i t (n.) s e d i m e n t a r y a c c u m u l a t i o n r e m a i n i n g after e r o s i o n o f L
finer-grained size fractions or types of s e d i m e n t .
lateral (adj.) on or toward t h e side of t h e a n i m a l (cf: anterior; distal; d o r s a l ;
m e d i a l ; posterior; p r o x i m a l ; v e n t r a l ) .
Lebensspur (pl., Lebensspuren; n.) a t r a c e fossil (= i c h n o f o s s i l ) . ("Lebens-
spur" is a G e r m a n w o r d that literally m e a n s " t r a c e of life"; t h e G e r m a n
w o r d " S p u r " i s related t o t h e a r c h a i c E n g l i s h w o r d "spoor.")
life a s s e m b l a g e (n.) fossils in a s t r a t u m t h a t r e p r e s e n t r e m a i n s of o r g a n -
isms that w e r e l i v i n g a t t h e t i m e o f f i n a l b u r i a l , u s u a l l y a t o r n e a r t h e
site of b u r i a l (cf: d e a t h a s s e m b l a g e ) .
lithofacies (n.) c h a r a c t e r i s t i c s of a s e d i m e n t a r y r o c k b a s e d on r o c k - t y p e
c o n t e n t (cf: b i o f a c i e s , facies).
lithology (n.) t h e n a t u r e o f a p a r t i c u l a r k i n d o f r o c k , i n c l u d i n g s u c h p r o p -
erties a s m i n e r a l l i c c o m p o s i t i o n a n d t h e s i z e o f t h e p a r t i c l e s o r crystals
o f w h i c h t h e r o c k c o n s i s t s ; t h e w o r d s o m e t i m e s i s u s e d for t h e d e s c r i p -
tion and systematic classification of rocks, a s c i e n c e m o r e properly
designated "petrography."
lithostratigraphic unit (n.) a r o c k - u n i t c h a r a c t e r i z e d by a p a r t i c u l a r rock-
t y p e or suite of r o c k - t y p e s (cf: b i o s t r a t i g r a p h i c u n i t ; t i m e - s t r a t i g r a p h i c
unit).
longitudinal (adj.) w i t h r e s p e c t to b r y o z o a n s , r e f e r r i n g to a s e c t i o n c u t
p a r a l l e l t o t h e l e n g t h o f z o o e c i a l t u b e s (Bassler 1953, G 1 1 , G 1 8 ) (cf:
t a n g e n t i a l ; transverse).
lophophore (n.) a c i l i a t e d , f o o d - g a t h e r i n g s t r u c t u r e f o u n d in b r a c h i o p o d s ,
e c t o p r o c t s , a n d s o m e o t h e r a n i m a l s ; a l t h o u g h t h e s e s t r u c t u r e s are
similar in a p p e a r a n c e and function in various groups of a n i m a l s , they
may not have a c o m m o n evolutionary origin.

mantle (n.) t h e tissue of a b r a c h i o p o d or m o l l u s c that s e c r e t e s t h e m a t e r i a l M

that c o m p r i s e s the s h e l l ; it is p a r t of t h e m a n t l e - c o m p l e x , a n d it g e n e r -
ally e n c l o s e s o r lies o n b o t h sides o f t h e m a n t l e - c a v i t y .
marker bed (n.) a s t r a t u m w i t h u n i q u e r o c k - t y p e , fossil c o n t e n t , o r o t h e r
f e a t u r e m a k i n g it u s e f u l for c o r r e l a t i o n or r e c o g n i t i o n of strata.
mass extinction (n.) g l o b a l d i s a p p e a r a n c e o f m a n y g r o u p s o f o r g a n i s m s
o v e r a relatively short t i m e i n t e r v a l .
Maysvillian (n.) t e r m for t h e m i d d l e s t a g e o f t h e G i n c i n n a t i a n S e r i e s , also
the middle age of C i n c i n n a t i a n E p o c h .

Glossary 287
 medial (adj.) o n o r t o w a r d t h e m i d d l e o f t h e a n i m a l , that is, toward t h e
plane of s y m m e t r y of the a n i m a l . S o m e p e o p l e , to avoid potential
c o n f u s i o n w i t h t h e statistical t e r m " m e d i a n , " u s e t h e t e r m " m e s i a l "
(cf.: a n t e r i o r ; d i s t a l ; d o r s a l ; l a t e r a l ; p o s t e r i o r ; p r o x i m a l ; ventral).
megacycle (n.) r e p e a t e d set o f s e d i m e n t a r y strata o f a b o u t o n e t o t w o m e -
ters in t h i c k n e s s .
member (n.) l i t h o s t r a t i g r a p h i c s u b d i v i s i o n o f a f o r m a t i o n .
mesodermal (adj.) p e r t a i n i n g t o t h e m i d d l e layer o f tissue i n t h e b o d y plan
o f t r i p l o b l a s t i c a n i m a l s (those h a v i n g t h r e e e m b r y o n i c tissue layers:
ectoderm, mesoderm, and endoderm).
metamorphism (n.) p r o c e s s e s o f c h e m i c a l o r p h y s i c a l alteration o f rocks
u n d e r t h e i n f l u e n c e o f h e a t , p r e s s u r e , o r b o t h (cf: d i a g e n e s i s ) .
metazoans (n.) a l t e r n a t i v e t e r m for " a n i m a l s . "
Milankovitch cycle (n.) r e p e a t e d p a t t e r n o f e v e n t s o r strata o n a f r e q u e n c y
s i m i l a r t o p e r i o d i c i t y o f o n e o r m o r e v a r i a t i o n s o f t h e E a r t h ' s orbit,
s u c h a s o b l i q u i t y o r e c c e n t r i c i t y . N a m e d after t h e Y u g o s l a v m a t h e m a t i -
c i a n M i l u t i n M i l a n k o v i t c h (1879-1958).
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.) a n a l t e r n a t i v e for " r e p l a c e m e n t "
(q.v.).
mold (n.) a n i m p r e s s i o n o r o t h e r n e g a t i v e r e p l i c a o f t h e internal o r exter-
n a l parts o f a n o r g a n i s m f o r m e d b y s e d i m e n t e n c l o s i n g t h e o r g a n i s m
(specifically, an e x t e r n a l m o l d ) or e n c l o s e d in a c a v i t y w i t h i n t h e or-
g a n i s m (specifically, an i n t e r n a l m o l d or steinkern) (cf: cast).
mutualism (n.) a n a s s o c i a t i o n b e t w e e n o r g a n i s m s o f t w o s p e c i e s i n w h i c h
e a c h r e c e i v e s s o m e b e n e f i t f r o m t h e a s s o c i a t i o n (cf: a e g i s m ; c o m m e n -
salism; parasitism; symbiosis).

N nektonic (adj.) swimming within a body of water (cf: benthic;

planktonic).
nomenclature (n.) t h e s c i e n c e o f n a m i n g t h e g r o u p s into w h i c h o r g a n i s m s
are classified (cf: t a x o n o m y ) .
nomen dubium (pl., nomina dubia; n.) in t a x o n o m y , a n a m e o f a t a x o n
t h a t i s s o p o o r l y u n d e r s t o o d a s t o b e o f d o u b t f u l status. T h e p h r a s e
literally m e a n s " d u b i o u s n a m e . "
nomen nudum (n.) in t a x o n o m y , a n a m e o f a t a x o n t h a t w a s m e n t i o n e d ,
b u t for w h i c h n o d e s c r i p t i o n , d e s i g n a t i o n o f a t y p e , illustration, a n d s o
o n , w a s g i v e n . T h e p h r a s e literally m e a n s " n a k e d n a m e . " A nomen
nudum is n o t t h e v a l i d n a m e of a g e n u s or s p e c i e s ; h o w e v e r , it m a y be
o f h i s t o r i c a l interest.

O o b l i g a t e (adj.) refers to an i n s t a n c e of an o r g a n i s m t h a t is a p a r t y to an
a s s o c i a t i o n t h a t is r e q u i r e d for t h e s u r v i v a l of t h e o r g a n i s m ; t h u s , if a
g i v e n parasite is u n a b l e to exist o u t s i d e of its p a r t i c u l a r a s s o c i a t i o n
w i t h its h o s t , t h e a s s o c i a t i o n is an o b l i g a t e o n e (cf: facultative).
obrution deposits (n.) s e d i m e n t a r y layers f o r m e d b y rapid d e p o s i t i o n , u s u -
ally of fine-grained particles so as to s m o t h e r any organisms.

288 Glossary
ontogeny (n.) e v e r y t h i n g that h a p p e n s t o a n o r g a n i s m f r o m t h e b e g i n n i n g
o f its life t o d e a t h . I n c l u d e d w i t h i n " o n t o g e n y " are e m b r y o l o g y , d e v e l -
o p m e n t , g r o w t h , m a t u r a t i o n , a n d s o o n (cf.: t a p h o n o m y ) .
o r d e r (n.) t h e l e v e l i n t h e L i n n a e a n h i e r a r c h y b e l o w c l a s s a n d a b o v e
family.
Ordovician Biodiversification Event (n.) a p p e a r a n c e o f n u m e r o u s m a j o r
groups of skeletonized m a r i n e invertebrate organisms d u r i n g the Or-
d o v i c i a n Period (syn., O r d o v i c i a n R a d i a t i o n ) .
Ordovician Period (n.) g e o l o g i c t i m e u n i t f o l l o w i n g t h e C a m b r i a n P e r i o d
and preceding the Silurian Period.
outcrops (n.) s u r f a c e e x p o s u r e s o f b e d r o c k f o r m e d b y n a t u r a l p r o c e s s e s .

packstone (n.) a l i m e s t o n e c h a r a c t e r i z e d by f r a g m e n t s in g r a i n - t o - g r a i n p
c o n t a c t , w i t h interstitial c a l c a r e o u s m u d (cf: g r a i n s t o n e ) .
paleobathymetry (n.) d e p t h o f a n a n c i e n t b o d y o f water.
paleontology (n.) t h e s t u d y o f a n c i e n t life b a s e d o n e v i d e n c e p r o v i d e d b y
fossils (from t h e G r e e k " p a l a i o s , " m e a n i n g " a n c i e n t , " + " o n , o n t o s , "
m e a n i n g " b e i n g , " " t h i n g , " o r " t h a t w h i c h has e x i s t e n c e , " + " l o g o s , "
m e a n i n g "discourse," and, by extension, "study of"); the word also
c o m m o n l y is spelled "palaeontology."
paleoslope (n.) a c h a n g e i n e l e v a t i o n o f a n a n c i e n t t o p o g r a p h i c a l s u r f a c e
s u c h as t h e sea floor.
Paleozoic Era (n.) o l d e s t m a j o r d i v i s i o n o f t h e P h a n e r o z o i c E o n (from t h e
G r e e k "palaios," m e a n i n g "ancient," + " z o o n , " m e a n i n g "animal").
parasitism (n.) a k i n d o f p r e d a t i o n i n w h i c h o n e a n i m a l lives w i t h i n , at-
t a c h e d to, or as a t e n a n t of a n o t h e r of a d i f f e r e n t s p e c i e s a n d f e e d s on
t h e h o s t , w h i c h g e n e r a l l y is a d v e r s e l y a f f e c t e d , b u t rarely k i l l e d (cf:
a e g i s m , c o m m e n s a l i s m ; predation; host; symbiosis).
paratype (n.) i n t a x o n o m y , w h e n t w o o r m o r e t y p e - s p e c i m e n s are d e s i g -
n a t e d for a s p e c i e s , a n d o n e of t h e m , t h e h o l o t y p e , is i d e n t i f i e d as
b e i n g m o s t r e p r e s e n t a t i v e o f t h e s p e c i e s , t h e o t h e r s are p a r a t y p e s (cf.:
cotype; holotype).
pelagic (adj.) l i v i n g w i t h i n t h e w a t e r c o l u m n , e i t h e r b y s w i m m i n g , drift-
ing, or floating.
period (n.) a g e o l o g i c t i m e u n i t , a s i n t h e O r d o v i c i a n P e r i o d , d u r i n g w h i c h
time the rocks in the C i n c i n n a t i area w e r e deposited; e q u i v a l e n t to
system i n t h e t i m e - s t r a t i g r a p h i c c l a s s i f i c a t i o n ; t h e h i e r a r c h y o f g e o -
l o g i c t i m e u n i t s , f r o m l a r g e s t t o s m a l l e s t is: e o n , era, p e r i o d , e p o c h ,
a n d a g e (cf.: a g e , e p o c h , e o n , era).
Phanerozoic Eon (n.) m a j o r s u b d i v i s i o n o f g e o l o g i c t i m e m a r k e d b y o c -
c u r r e n c e o f a b u n d a n t fossils o f m e t a z o a n s .
phylum (pl., p h y l a ; n.) t h e level i n t h e L i n n a e a n h i e r a r c h y b e l o w k i n g d o m
a n d a b o v e class.
plane of s y m m e t r y (n.) t h e p l a n e o f s y m m e t r y c o m m o n l y i s c a l l e d t h e
" m i d l i n e " o f t h e a n i m a l (see: b i l a t e r a l l y s y m m e t r i c a l ) ,
planispiral (adj.) said of a shell c o i l e d in a s i n g l e p l a n e .

Glossary 289
 planktonic (adj.) s u s p e n d e d w i t h i n o r f l o a t i n g u p o n a b o d y o f water, w i t h
transportation exclusively or primarily by currents and other move-
m e n t s o f t h e w a t e r itself (cf.: b e n t h i c ; n e k t o n i c ) .
polymorphism (n.) t h e e x i s t e n c e i n o n e s p e c i e s o f i n d i v i d u a l s o f m o r e
t h a n o n e s i z e , s h a p e , o r n a t u r e ; i n b r y o z o a n s , for e x a m p l e , t h e r e m a y
b e several different p o l y m o r p h s i n a g i v e n c o l o n y , e a c h o f w h i c h , pre-
s u m a b l y , p e r f o r m e d a different f u n c t i o n (cf.: d i m o r p h i s m ) .
p o s t e r i o r (adj., n.) on or t o w a r d t h e rear of t h e a n i m a l . ( N o t e that w h a t is
f u n c t i o n a l l y p o s t e r i o r i n a g i v e n a n i m a l m a y n o t c o r r e s p o n d t o the
a n a t o m i c a l posterior. For e x a m p l e , i n h u m a n s , t h e b a c k i s a n a t o m i -
c a l l y d o r s a l , b u t , a s y o u w a l k a r o u n d , y o u r b a c k i s a t t h e rear o f y o u r
bod\ a n d . h e n c e , is f u n c t i o n a l l y "posterior"; cf.: .interior; distal; dorsal;
lateral; m e d i a l ; p r o x i m a l ; ventral.)
predation (n.) a r e l a t i o n s h i p in w h i c h o n e a n i m a l c o n s u m e s a n o t h e r organ-
i s m ; t h e c o n s u m e r is c a l l e d a predator, a n d that c o n s u m e d is called the
prey (cf: a e g i s m ; c o m m e n s a l i s m ; m u t u a l i s m ; parasitism; symbiosis).
primary producers (n.) o r g a n i s m s at t h e b a s e of a f o o d c h a i n that form
organic c o m p o u n d s such as carbohydrates from simple components
such as carbon dioxide and water by the process of photosynthesis.
p r i o r i t y (n.) i n t a x o n o m y , t h e c o n v e n t i o n t h a t , w h e n t h e r e i s a n o l d e r
n a m e a n d a y o u n g e r n a m e t h a t b o t h h a v e b e e n u s e d t o d e s i g n a t e the
s a m e t a x o n , t h e o l d e r n a m e b e c o m e s t h e official n a m e o f the t a x o n .
proximal (adj.) o n o r t o w a r d t h e c e n t e r o f t h e a n i m a l ; for e x a m p l e , y o u r
finger tips are distal, w h e r e a s y o u r s h o u l d e r is p r o x i m a l (cf: anterior;
distal; dorsal; lateral; m e d i a l ; posterior; v e n t r a l ) .
pseudobivalved (adj.) said of a shell that is s h a r p l y folded a l o n g t h e dorsal
axis s o a s t o g i v e t h e a p p e a r a n c e o f b e i n g b i v a l v e d , a l t h o u g h c a l c i f i c a -
t i o n is c o n t i n u o u s across t h e a x i s ; t h u s , t h e r e is no t r u e h i n g e ; rostro-
c o n c h m o l l u s c s are p s e u d o b i v a l v e d (cf: b i v a l v e d ; u n i v a l v e d ) .

R radiometric dating (n.) d e t e r m i n a t i o n o f g e o l o g i c a g e t h r o u g h m e a s u r e -

m e n t o f t h e d e c a y o f r a d i o a c t i v e isotopes c o n t a i n e d i n c e r t a i n m i n e r -
als; c o m m o n m e t h o d s are u r a n i u m - l e a d , p o t a s s i u m - a r g o n , a n d car-
bon-14 a n a l y s i s .
recrystallization (n.) t h e process w h e r e b y t h e o r i g i n a l s u b s t a n c e of a shell,
for i n s t a n c e , is r e c o n s t i t u t e d or c o n v e r t e d to crystals or to crystals of a
different n a t u r e ; for e x a m p l e , s o m e m o l l u s c shells consist of the m i n e r a l
a r a g o n i t e , w h i c h , u n d e r c e r t a i n t a p h o n o m i c c o n d i t i o n s , may, over t i m e ,
convert to the mineral calcite. Aragonite and calcite have the same
c h e m i c a l c o m p o s i t i o n ( C a C O J , b u t h a v e a different t h r e e - d i m e n s i o n a l
crystal s t r u c t u r e .
relative age (n.) t i m e o f o c c u r r e n c e o f a n e v e n t i n g e o l o g i c t i m e relative
to a n o t h e r , in t h e s e n s e of o l d e r or y o u n g e r (cf: a b s o l u t e age).
r e p l a c e m e n t (= m i n e r a l i z a t i o n ) (n.) a t a p h o n o m i c p r o c e s s in w h i c h t h e
a c t u a l s u b s t a n c e o f a n o r g a n i s m i s r e m o v e d bit-by-bit, a n d m i n e r a l
m a t t e r is left in its p l a c e ; a fossil p r o d u c e d by s u c h a p r o c e s s .

290 Glossary
Richmondian (n.) t h e y o u n g e s t s t a g e o f t h e C i n c i n n a t i a n S e r i e s ; a l s o t h e
youngest age of the C i n c i n n a t i a n E p o c h ,
rock-stratigraphic unit (see l i t h o s t r a t i g r a p h i c u n i t ) .

scientific n a m e (see: s p e c i e s n a m e ) . s
sea floor s p r e a d i n g (n.) p r o c e s s b y w h i c h n e w o c e a n i c c r u s t i s f o r m e d b y
u p w e l l i n g o f m o l t e n m a t e r i a l a l o n g a f i s s u r e o r rift.
seismite (n.) r o c k layer f o r m e d o r a l t e r e d b y e a r t h q u a k e s h o c k .
s e r i a l s e c t i o n i n g (n.) c u t t i n g a fossil, for e x a m p l e , a b r a c h i o p o d , i n t o a
n u m b e r of thin, e q u a l l y spaced slices of k n o w n orientation. F r o m
t h e s e s l i c e s , it is p o s s i b l e to r e c o n s t r u c t t h e interior of t h e s h e l l ; in ef-
fect, o n e i s d o i n g a p a l e o n t o l o g i c a l C A T s c a n . I n s p e c i m e n s i n w h i c h
t h e interior o f t h e shell i s f i l l e d w i t h r o c k m a t r i x , t h i s m a y b e t h e o n l y
practical way to see w h a t is inside the shell.
series (n.) a t i m e - s t r a t i g r a p h i c s u b d i v i s i o n of a s y s t e m , s u c h as C i n c i n n a -
tian S e r i e s o r U p p e r O r d o v i c i a n S e r i e s (cf.: s t a g e ; s y s t e m ) .
sequence boundary (n.) s t r a t i g r a p h i c h o r i z o n m a r k i n g t h e b e g i n n i n g o r
t e r m i n a t i o n o f strata d e p o s i t e d d u r i n g a n i n t e r v a l o f sea l e v e l rise o r
fall, u s u a l l y i n d i c a t i n g a c e s s a t i o n of s e d i m e n t a t i o n or i n t e r v a l of e r o -
sion (cf: u n c o n f o r m i t y ) .
s e q u e n c e s t r a t i g r a p h y (n.) t h e s t u d y o f s e q u e n c e s o f s e d i m e n t a r y r o c k s
b o u n d e d b y s u r f a c e s o f n o n - d e p o s i t i o n o r e r o s i o n , u s u a l l y related t o
m a j o r c h a n g e s i n sea l e v e l .
shell pavement (n.) dense accumulation of shells of molluscs or
brachiopods.
siliciclastics (n.) s e d i m e n t s c o n s i s t i n g o f p a r t i c l e s o f clay, m u d , silt, s a n d ,
or coarser material c o m p o s e d of silica-rich materials.
socket (n.) o n e o f t h e d e p r e s s i o n s a l o n g t h e h i n g e l i n e o f a n a r t i c u l a t e
b r a c h i o p o d o r p e l e c y p o d into w h i c h f i t s into a t o o t h o f t h e o p p o s i t e
valve; the teeth and sockets, collectively t e r m e d the dentition, serve to
p r e v e n t t h e v a l v e s f r o m b e i n g t w i s t e d a p a r t by, for e x a m p l e , a p r e d a t o r
(cf: d e n t i t i o n ; t o o t h ) .
s p e c i e s (pl., s p e c i e s ; n.) t h e level i n t h e L i n n a e a n h i e r a r c h y b e l o w g e n u s ;
a s p e c i e s is t h e basic k i n d of o r g a n i s m r e c o g n i z e d by scientists.
species n a m e (n.) t h e f o r m a l n a m e o f a s p e c i e s , c o n s i s t i n g o f b o t h t h e
g e n e r i c n a m e a n d t h e s p e c i f i c (or trivial) n a m e ; c o m m o n l y c a l l e d a
b i n o m e n , b e c a u s e it consists of two n a m e s ; c o m m o n l y called the sci-
e n t i f i c n a m e o f t h e s p e c i e s (cf: g e n e r i c n a m e ; s p e c i f i c n a m e ) .
specific epithet (see: s p e c i f i c n a m e ) .
specific n a m e (n.) t h e s e c o n d w o r d o f a s p e c i e s n a m e ; c o m m o n l y c a l l e d
t h e t r i v i a l n a m e ; s o m e t i m e s c a l l e d t h e " t r i v i a l e p i t h e t " (cf: g e n e r i c
name; species name).
stage (n.) a t i m e - s t r a t i g r a p h i c s u b d i v i s i o n of a series, s u c h as t h e E d e n i a n
S t a g e ; often u s e d for i n t e r c o n t i n e n t a l o r r e g i o n a l c o r r e l a t i o n . T h e r e i s
an e q u i v a l e n c e b e t w e e n a given stage and the time span d u r i n g w h i c h
t h a t b o d y o f r o c k w a s d e p o s i t e d ; for e x a m p l e , t h e r o c k s o f t h e E d e n i a n
S t a g e w e r e laid d o w n d u r i n g t h e E d e n i a n A g e (cf: series; s y s t e m ) .

Glossary 291
 steinkern (n.) p e t r i f i e d s e d i m e n t i n f i l l i n g of a shell or b o d y cavity, an in-
t e r n a l m o l d (from t h e G e r m a n " S t e i n , " m e a n i n g " s t o n e , " + " K e r n , "
m e a n i n g "kernel"); although a G e r m a n n o u n , it c o m m o n l y appears
n e i t h e r c a p i t a l i z e d n o r in italics (cf.: c a s t ; m o l d ) .
storm cycles (n.) a layer of s e d i m e n t a r y r o c k p r o d u c e d d u r i n g a s i n g l e
s t o r m , c o m m o n l y i t c o n s i s t s o f a n interval r e p r e s e n t i n g i n t e n s e distur-
b a n c e o f t h e sea floor f o l l o w e d b y a n interval r e p r e s e n t i n g w a n i n g o f
s t o r m activity.
stratigraphy (n.) the study of layered or stratified rocks (cf: biostratigraphy),
stromatolites (n.) h n e K l a m i n a t e d s e d i m e n t a r y a c c u m u l a t i o n s r e s u l t i n g
from trapping and b i n d i n g of sedimentary particles by cyanobacteria;
usually sheet-like, d o m e - s h a p e d or c o l u m n a r in form; sometimes
called "algal stromatolites," b e c a u s e cyanobacteria o n c e were thought
to be algae.
stromatoporoids (n.) d o m e - s h a p e d , c o l u m n a r , or b r a n c h i n g c a l c a r e o u s skel-
etons, usually with finely laminated mierostructure, formed by calcare-
o u s s p o n g e s , o c c u r r i n g i n O r d o v i c i a n t h r o u g h C r e t a c e o u s rocks.
subspecies (n.) a level i n t h e L i n n a e a n h i e r a r c h y just b e l o w s p e c i e s ; s o m e -
times called variety.
substratum (n.) a s u r f a c e o n w h i c h a n o r g a n i s m m i g h t b e a t t a c h e d ; i t
m i g h t be the surface of a stratum, rock, shell, or other hard object.
( N o t t o b e c o n f u s e d w i t h " s u b s t r a t e , " w h i c h has a p r e c i s e b i o l o g i c a l
m e a n i n g a n d s h o u l d n o t b e u s e d i n this c o n t e x t . )
succession (n.) i n e c o l o g y , t h e r e p l a c e m e n t o f o n e a s s e m b l a g e o f o r g a n -
isms b y a n o t h e r , o f t e n i n w h i c h t h e a c t i v i t i e s o f t h e initial a s s e m b l a g e
of o r g a n i s m s altered t h e e n v i r o n m e n t in s u c h a w a y that a different
assemblage of organisms can occupy the environment; c o m m o n l y
c a l l e d " e c o l o g i c s u c c e s s i o n " (cf: b i o t a l s u c c e s s i o n ; fossil s u c c e s s i o n ) .
s u s p e n s i o n f e e d i n g (n., adj.) (cf: d e p o s i t f e e d i n g ; filter feeder).
symbiont (n.) e a c h o f t h e o r g a n i s m s i n v o l v e d i n a s y m b i o t i c r e l a t i o n s h i p .
symbiosis (n.) a c l o s e a s s o c i a t i o n b e t w e e n o r g a n i s m s o f t w o s p e c i e s ; c o m -
m o n l y a s y m b i o t i c r e l a t i o n s h i p is b e n e f i c i a l to b o t h o r g a n i s m s in-
volved, but parasitism also c a n be considered a symbiotic relationship
(cf: c o m m e n s a l i s m ; m u t u a l i s m ; parasitism).
symmetrical, s y m m e t r y (see: bilaterally symmetrical; plane of
symmetry).
synecology (n.) e c o l o g y o f several o r m a n y s p e c i e s o f o r g a n i s m s t o g e t h e r
in a g i v e n t i m e a n d p l a c e (cf: a u t o e e o l o g y ) .
syntypes (see: c o t y p e s ) .
system (n.) a m a j o r t i m e - s t r a t i g r a p h i c s u b d i v i s i o n , t h e rock record of a p e -
riod of g e o l o g i c t i m e , s u c h as t h e O r d o v i c i a n S y s t e m (cf: series; stage),
s y s t e m a t i c s (see: t a x o n o m y ) .

T 'laconic Orogeny (n.) e p i s o d e o f i n t e n s e c r u s t a l d e f o r m a t i o n that o c -

c u r r e d d u r i n g t h e L a t e O r d o v i c i a n w h e n t h e plate o f o c e a n i c c r u s t
b e n e a t h the Iapetus Sea was subducted b e n e a t h the continental Lau-

292 Glossary
 r e n t i a n plate; n a m e d for t h e ' l a c o n i c M o u n t a i n r e g i o n o f e a s t e r n N e w
York w h e r e this d e f o r m a t i o n first w a s r e c o g n i z e d .
tangential (adj.) w i t h r e s p e c t to b r y o z o a n s , r e f e r r i n g to a s e c t i o n c u t p a r a l -
lel t o t h e s u r f a c e o f t h e z o a r i u m a n d c l o s e e n o u g h t o t h e s u r f a c e o f t h e
colony to show the mature features of the z o o e c i a in cross-section;
s u c h a t a n g e n t i a l s e c t i o n is p e r p e n d i c u l a r to a l o n g i t u d i n a l s e c t i o n
(Bassler 1953, G 1 5 , G18) (cf.: l o n g i t u d i n a l ; transverse).
t a p h o n o m y (n.) 1. e v e r y t h i n g t h a t h a p p e n s to an o r g a n i s m after it d i e s ; 2.
t h e study o f t h o s e p h e n o m e n a that t e n d t o d e s t r o y t h e r e m a i n s a n d
traces of a l i v i n g t h i n g as w e l l as of t h o s e that t e n d to p r e s e r v e t h o s e
r e m a i n s a n d t r a c e s (from t h e G r e e k " t a p h o s , " m e a n i n g " g r a v e " o r
"tomb," + "nomos," meaning "law").
taxobases (sing., taxobasis; n.) c h a r a c t e r i s t i c s o n w h i c h t h e t a x o n o m i c p o -
sition o f a g r o u p o f o r g a n i s m s i s b a s e d ( p r o n o u n c e d : t a x - u h - b a s e -
ease).
taxon (pl., taxa; n.) a f o r m a l b i o l o g i c a l g r o u p to w h i c h an o r g a n i s m is as-
s i g n e d ; h u m a n s , for e x a m p l e , b e l o n g i n a t a x o n a t t h e f a m i l y - l e v e l o f
the L i n n a e a n hierarchy called Hominidae.
taxonomy (n.) t h e s c i e n c e o f a s s i g n i n g o r g a n i s m s t o their p r o p e r b i o l o g i c a l
g r o u p s (alternative n a m e s are c l a s s i f i c a t i o n a n d s y s t e m a t i c s ) .
tempestites (n.) s e d i m e n t a r y r o c k s f o r m e d b y s t o r m p r o c e s s e s , o f t e n ex-
h i b i t i n g features s u c h a s p a r t i c l e - s i z e s o r t i n g , fossil b r e a k a g e , a b r a s i o n ,
and orientation resulting from violent water m o v e m e n t .
time-averaged (adj.) p e r t a i n i n g to s e d i m e n t a r y d e p o s i t s or fossil a s s e m -
b l a g e s that r e p r e s e n t a c c u m u l a t i o n o v e r a relatively l o n g t i m e i n t e r v a l ,
often c o n s i s t i n g o f m i x t u r e s o f fossils f r o m s u c c e s s i v e g e n e r a t i o n s .
time-stratigraphic unit (n.) a b o d y o f r o c k f o r m e d d u r i n g a p a r t i c u l a r
interval of g e o l o g i c t i m e , s u c h as a s y s t e m or series (cf: b i o s t r a t i g r a p h i c
u n i t ; l i t h o s t r a t i g r a p h i c unit).
tooth (n.) o n e o f t h e p r o j e c t i o n s a l o n g the h i n g e l i n e o f a n articulate b r a c h i o -
p o d or p e l e c y p o d that fits into a socket in the opposite valve; the teeth a n d
sockets, collectively t e r m e d t h e d e n t i t i o n , serve to p r e v e n t the valves from
b e i n g twisted apart by, for e x a m p l e , a predator (cf: d e n t i t i o n ; socket).
t r a c e fossil (n.) a fossil that s h o w s t h a t an a n i m a l or p l a n t e x i s t e d , b u t
w h i c h lacks actual remains or replicas of remains; it was m a d e by or is
t h e result o f g e n u i n e a c t i v i t y o f t h e l i v i n g o r g a n i s m , rather t h a n i n v o l -
u n t a r y m o v e m e n t , s u c h a s a shell's b e i n g d r a g g e d across t h e sea f l o o r
b y c u r r e n t s o r w a v e a c t i o n ; i n c l u d e d h e r e are f o o t p r i n t s , trails, bur-
rows, b o r i n g s , t o o t h m a r k s , a n d so on (= i c h n o f o s s i l = Lebensspur)
(cf: b o d y fossil).
t r a n s v e r s e (adj.) w i t h r e s p e c t to b r y o z o a n s , r e f e r r i n g to a s e c t i o n c u t at
right a n g l e s t o t h e d i r e c t i o n o f g r o w t h o f t h e z o a r i u m (Bassler 1953,
G 1 5 , G18) (cf: l o n g i t u d i n a l ; t a n g e n t i a l ) .
trivial n a m e (see: s p e c i f i c n a m e ) .

unconformity (n.) a s u r f a c e s e p a r a t i n g layers or o t h e r b o d i e s of r o c k that u

represents e i t h e r a c e s s a t i o n o f d e p o s i t i o n o r a n interval o f e r o s i o n .

Glossary 293
 univalved (adj.) said of a shell t h a t c o n s i s t s of a s i n g l e v a l v e (cf.: b i v a l v e d ;
pseudobivalved).

V variety (n.) i n f o r m e r t i m e s , e q u i v a l e n t t o t h e t a x o n o m i c r a n k o f s u b s p e -
c i e s , b u t n o l o n g e r r e c o g n i z e d w i t h i n t h e z o o l o g i c a l c o m m u n i t y (In-
t e r n a t i o n a l C o m m i s s i o n o n Z o o l o g i c a l N o m e n c l a t u r e 1999,19); n o w a -
d a y s c o m m o n l y u s e d s p e c i f i c a l l y for k i n d s o f p l a n t s bred deliberately
by horticulturists.
ventral (adj.) o n o r toward t h e b o t t o m o f t h e a n i m a l ; the n o u n form o f the
c o n c e p t is "venter." ( N o t e that w h a t is f u n c t i o n a l l y ventral in a g i v e n a n i -
m a l m a y n o t c o r r e s p o n d t o the a n a t o m i c a l ventral. For e x a m p l e , i n h u -
m a n s , the belly is anatomically ventral, but, as you walk around, your
b e l l y is at t h e front 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 a l l y "anterior"; cf:
anterior; distal; dorsal; lateral; m e d i a l ; posterior; proximal.)

z zone (see b i o z o n e ) .
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 INDEX

Page numbers in italics indicate illustrations. Chlorophyta ("green algae"), 67, 68

Dasyeladaceae, 67, 68
I .ike most indexes, this one is nowhere near as thorough as it might
Anomaloides: A. reticulatus, 68
have been; the tremendous number of items that might have been
Cyclocrinites: C. darwini, 66, 6 7 - 6 8
included would have made the index completely unwieldy.
Ischadites: I. circularis, 68
Certain words and concepts are not separately listed in the
Lepidolites: L. dickhauti, 66, 67, 68
index, because they appear far too many times in the book. Ex-
Cyanophyta (called "blue-green algae," but not actually
amples include Cincinnatian, Ordovician, and Ohio River.
algae). See Cyanobacteria
Although places in the states of Indiana, Kentucky, and Ohio
Cylindrocoelia: C. covingtonensis, 68
are included in the index, every appearance of the name of the
Faberia: F. anomala, 68
whole state is not.
Girvanella. See Cyanobacteria
This volume is not intended as a comprehensive taxonomic oncolites, 67
work. Hence, we have not endeavored to put every genus, species,
Phaeophyta ("brown algae"), 67
and so on, in its complete Linnaean Hierarchy in the index.
Pyrrophyta: dinoflagellates, 69, 239, 248, 283
Rather, we have used the names of upper-level taxa that appear in
Rhodophyta ("red algae"), 67, 68
the text. However, we have departed from this policy in some cases,
Solenopora: S. richmondensis, 68
with the hope of greater clarity and usefulness for the reader.
stromatolites. See Cyanobacteria
Although so-called "common names"things like "brittle a l i g n m e n t of fossils. See taphonomy
star" and "spiny oyster" probably should he sandwiched be- a l l o c h e m , 49, 50
tween quotation marks, we chose not to do this so as not to
Allocotichnus: A. dyeri. See taphonomy: trace fossils
clutter the index more than absolutely necessary.
Allolichas. See Arthropoda: Trilobita: Lichida
Alps, 3
alternata, Rafinesquina. See Brachiopoda: Articulata
alveolatus, Megalograptus. See Arthropoda: C h e l i c e r a t a :
abalones. See Mollusca: Gastropoda: Archaeogastropoda Eurypterida
absolute age, absolute dating. See time Amhonychia. See Mollusca: Pelecypoda
Acadian Orogeny, M o u n t a i n s , 8, 62 American Midland Naturalist, 172
Acanthochaetetes. See Porifera: sclerosponges, coralline American Midcontinent Province. See faunal provinces
sponges American Upper Ordovician Standard, 51
acetate peels. See techniques amoenum, Beloitoceras. See Mollusca: C e p h a l o p o d a
acid dissolution, acid etching. See techniques Amorphognathus. See conodonts
Acidaspis. See Arthropoda: Trilobita: Odontopleurida Amphilichas. See Arthropoda: Trilobita: Lichida
acme zone. See biostratigraphic units: zone: epibole Amsden, T h o m a s W., 54
acritarchs, 5 2 , 6 7 , 6 8 - 6 9 , 239, 247, 279, 309 anacanthus, Arnheimograptus. See Hemichordata:
Multiplicisphaeridium: M. micraulaxum, 6 6 , 67 Graptoloidea
Ordovicium: O. gracile, 66, 67 Anderson, D. R 259
Veryhachium: V. edenense, 66, 67 andersoni, Pontiometra. See Echinodermata: Crinoidea
Actinoceratoidea. See Mollusca: C e p h a l o p o d a a n e m o n e s . See C n i d a r i a : Anthozoa
acutilirata, Platvstrophia. See Brachiopoda Angseesing, J. P. A., 212, 297
Adams County, Ohio, 75, 158, 162, 163, 216, 229, pl. 7 angularis, Glyptocrinus. See Echinodermata: C r i n o i d e a :
aegism. See ecologic associations Camerata
Agassiz, Louis, 21, 260, 273 An n elida, 24, 31, 117, 142-145, 147, 182, 265, 306, 309, 316,
age, dating. See time 319. See also worms
Ager, Derek V., 7 Cornulites, 93, 123, 142, 144, 145, 182, 243, 244, 245, 246,
Aglaspida. See Arthropoda 247, 314, 316
Agnew, ]ohn. 7 1 , 72, 171, 180, pl. 14 myzostomid annelid worms, 182
alata, Ctenobolbina. See Arthropoda: Crustacea: Ostracoda Polychaeta, 143, 144, 204, 205, 212, 221, 224, 238, 247, 248
alata, Nereigenys. See Annelida: scolecodonts serpulids, 144
Albers, H. E., 259 spirorbids, 144
Aldrich, T. H., 259 Protoscolex, 144
Algae, 49, 5 2 , 6 7 - 6 9 , 123, 139, 212, 219, 234, 239, 241, 242, P. omatus, 142
279, 292. See also acritarchs; chitinozoans scolecodonts, 143, 265

323
 Nereigenys: N. alata, PL 5 Eoleperditia, 163
Annual Reviews, 4 Lepcrditicopida, 164
anomala, Faberia. See Algae Palaeocopida, 163
Anomalocrinus. See Echinodermata: Crinoidea: Disparida Podocopida, 163
Anomalodonta. See Mollusca: Pelecypoda Quadrijugator: Q. regularis, 160, 161
Anomaloides. See Algae: Chlorophyta: Dasycladaceae Diplopoda: millipedes, 147
Anthony, John G o u l d , 2 5 9 - 2 6 0 , 261, 269, 276 Insecta, 105, 119, 147, 162, 282
Anthozoa, anthozoans. See C n i d a r i a scorpions. See Chelicerata: Arachnida
Anticosti Island, C a n a d a , 7 2 - 7 3 spiders. See Chelicerata
Aphelognathus. See conodonts ticks. See Chelicerata: Arachnida
Aplaeopliora. See Mollusca Trilobita, 6, 7, 27, 3 1 , 4 0 , 56, 6 0 , 9 2 , 9 3 , 117, 137, 144,145,
Appalachian M o u n t a i n s , basin, 3, 5, 8, 10, 53, 6 1 , 216, 220, 147-157, 1 6 0 - 1 6 1 , 1 6 2 , 1 6 3 , 2 0 2 , 203, 204, 205,
231, 233, pl. 12 207, 210, 212, 219, 220, 223, 224, 225, 2 2 6 , 2 2 8 ,
approximate, Palaeasterina. See Echinodermata: Asteroidea 229, 231, 232, 233, 235, 238,240, 241, 244, 245,
Arachnida. See Arthropoda: Chelicerata 247, 251, 252, 253, 260, 268, 275, 286
aragonite, aragonitic. See c a l c i u m carbonate Acidaspis. See Odontoplcurida
area, Archinacetla. See Mollusca: Monoplacophora Calymene, 149, 150, 296, 305
Archaeogastropoda. See Mollusca: Gastropoda C. meeki. See Flexicalymene meeki
Archinacella. See Mollusca: Monoplacophora C. meeki-retrorsa. See Flexicalymene retrorsa
A r n h e i m Formation, 44, 4 8 , 5 3 , 6 8 , 105, 107, 108, 123, 151, C. senaria. See Flexicalymene senaria
153, 164, 176, 180, 196, pl. 8 C a l y m e n i d a , c a l y m e n i d s , 149, 151, 226, 241
Oregonia M e m b e r , 44, 53, 56, 63, 223. See also Oregonia Flexicalymene, 40, 148, 150, 151, 152, 153, 154, 160,
Formarion 207, 210, 224, 225, 226, 229, 232, 251, 253, 264,
Sunset M e m b e r , 4 4 , 53, 56, 63, 164, 224. See also Sunset 298, 317
Formation F. granulosa, 149, 150, 151,309
Arnheimograptus. See Hemichordata: Graptoloidea F. meeki, 148, 149,150, 151,207, 228, 229, 264,
Arthropoda, 6, 92, 117, 129, 146-164, 204, 205, 221, 222, 298, 317
224, 245, 253, 280, 282. See also Chelicerata: F. retrorsa, 149, 150, 151, pl. 8
Arachnida F. senaria, 149, 153
Aglaspida, aglaspids, 158, 163, 280, 300 Gravicalymene, 225, 226
Neostrabops, 158, 163; N. martini, 158, 163 Cryptolithus, 113, 155, 203, 204, 210, 225, 226, 232,
C h e l i c e r a t a , 27, 161 240, 241, 253
Arachnida, 147, 161 C. tessellatus, 148, 149, 154, 155
mites, 161 Decoroproetus: D. parviusculus, 154, 155
scorpions, 161, 162 Flexicalymene. See C a l y m e n i d a
spiders, 161, 162 Gravicalymene. See C a l y m e n i d a
ticks, 161 lsotelus, 2 7 , 6 0 , 137, 148, 153, 154, 155,203, 2 0 4 , 2 0 7 ,
Eurypterida, 147, 154, 158, 159, 161-163, 235, 238, 241, 210, 224, 225, 226, 232, 235,240, 241, 252, 299,
247, 253, 280, 300, 305 305, 316, 317
Eocarcinosoma: E. batrachophthalmus, 162 J. brachycephalus, 153
Megalograptus, 154, 158, 159, 161, 162, 253, 305, 312 /. gigas, 153
M. aheolatus, 162 /. latus, 149
M. ohioensis, 158, 159, 162 1. maximus, 148, 149, 152, 153, pl. 7
M. shideleri, 162 1. rex, 153, 316
M. welchi, 162 I j c h i d a , lichids
M. williamsae, 162 Mlolichas: A. halli, pl. 6
Xiphosurida: horseshoe crabs, Limulus, 147, 151, 161, Amphilichas: A. halli. See Allolichas halli
163, 280, pl. 3 Odontoplcurida, odontopleurids, 156, 160,251
C h i l o p o d a : centipedes, 147 Acidaspis, 228, 229, 232, 251
C r u s t a c e a , 26, 30, 147, 148, 151, 163, 210, 237, 238, 241 A. cincinnatiensis, 156, 160
barnacles, 145, 188, 248, 250 A. onealli, 271
Isopoda, 148 Primaspis, 92, 160
pillbug, 148 P. crosotus, 156, 160
Malacostraca, 248 Olenida, olenids
crabs, 9 6 , 148, 163, 235 Triarthrus, 146, 147, 160, 226, 253
lobsters, 148, 163, 253 T. eatoni, 154, 155, 160
shrimps, 163 T. spinosus, 160
Ostracoda, 30, 31, 35, 129, 147, 160, 163-164, 221, 235, Phacopida, phacopids
241, 247, 248, 297, 320 Ceraurinus, 220
Ceratopsis: C. chambers!, 160, 16J C. icarus, 157
Ctenobolhina: C. alata, 160, 161 Ceraurus: C. milleranus, 157, 269

324 Index
 Platycoryphe: P. christyi, 157 Bellevue Limestone, J l , 12, 53, 55, 56, 63, 68, 85, 89, 95, 99,
Tricopelta: T. breviceps, 157 125, 174,214, 223, 232, 251. See also M c M i l l a n
Platycoryphe. See Phacopida Eormation
Primaspis. See Odontopleurida Beloitoceras. See Mollusca: C e p h a l o p o d a
Proetus parviusculus. See Decoroproetus parviusculus benthic, benthonic, benthos, 3, 4, 7, 50, 102, 145, 163, 195,
trace fossils, 151-153. See also taphonomy: trace fossils 204, 205, 218, 230, 234, 237, 239, 280, 288, 290
Rusophycus. See taphonomy: trace fossils bentonite. See K-bentonite
Triarthrus. See Olenida Berdan, Jean, 163, 164, 320
Articulata. See Brachiopoda B e r g m a n , C l a e s F., 143, 304
Asaphoidichnus. See taphonomy: trace fossils Bergstrom, Jan, 153, 318
Ascocerida. See Mollusca: C e p h a l o p o d a : Nautiloidea Bergstrom, Stig M., 2 , 5 1 , 62, 196, 306, 308, 319, 321, 331
Ashermann, George (?), 273, 275 Best, Robert, 260, 276
Ashgillian Stage, pl. 2 bibliographies, compilation of, 15, 33, 35
associations. See chapters on individual groups of a n i m a l s ; biclavata, Diplocraterion. See taphonomy: trace fossils
ecologic associations bilix lata, Cyclonema. See Mollusca: Gastropoda
Asteriacites. See taphonomy: trace fossils bilix, Cyclonema. See Mollusca: Gastropoda
Asteroidea. See Echinodermata binomen. See taxonomy: nomenclature: species n a m e
Astraspis. See Chordata: Vertebrata: Agnatha Binomial System of Nomenclature. See taxonomy:
auhurnensis, Platystrophia ponderosa. See Brachiopoda: nomenclature
Platystrophia ponderosa bioclaustration, 156, 210, 243, 281. See also taphonomy:
Audubon, John James, 16, 277 bioimmuration
Aulacera. See Porifera: Stromatoporoidea biocoenose, biocoenosis. See life assemblage
Ausich, W i l l i a m I 186 biofacies. See facies
Austin, George M., 260 bioherm, 7 1 , 72, 7 8 - 8 1 , 85, 9 1 , 131, 242, 281
Austinella. See Brachiopoda: Articulata "bryoherm," 91
Australia, 199, 215, 241, 251. See also Great Barrier Reef Great Barrier Reef, Australia, 9 1 , 241
auteeology. See ecology patch reef, 80, 81
bioimmuration. See taphonomy: bioimmuration
Babcock, Loren E. 153 biostratigraphic units, 4 6 , 281
bacteria, 123, 239. See also cyanobacteria zone, 46
baeri, Charactoceras. See Mollusca: C e p h a l o p o d a b i o z o n e , 4 6 , 195, 2 8 1 , 2 8 4
baeri, Xenocrinus. See Echinodermata: Crinoidea: Camerata epibole, a c m e zone, 60, 281, 284
Bahama Platform, ix, 220 biostratigraphy, 52
balanoides, Enoploura. See Echinodermata: Stylophora biostrome, 81, 281
Baldwin-Wallace College, 29, 30 biota! succession, 4 7 - 4 9 , 6 1 , 281, 292
ball-and-pillow structures, 60, 280 biozone. See biostratigraphic units: zone
Baltica. See paleogeography Bischoff, G. C. O., 182
Bambach, Richard K 237, 238, 239, 300, 315 bivalved a n i m a l s , 281, 290. See also Arthropoda: C r u s t a c e a :
Bardstown Eormation, 2H Ostracoda; Brachiopoda: Mollusca:
barnacles. See Arthropoda: Crustacea Pelecypoda
Barnhart, W . C , 260, 266 Bivalvia. See Mollusca: Pelecypoda
Bassler, Ray S., 18, J9, 20, 21, 25, 30, 33, 3 4 - 3 6 , 48 Blackwell Publishing, 185, 199, 229,
Bassler, Simon Stein, 34 Blanchester Member. See Waynesville Formation
"Bassleroscope," 35 Blastophycus, 212
batrachophthalmus, Eocarcinosoma. See Arthropoda: C h e l i c - Blue Limestone, 47
erata: Eurypterida Blue M i a m i Limestone, 47
Baumiller, Tomasz K., 124, 125, 183, 186 B M H N . See Natural History M u s e u m , London
Bean, W. H 260 Boardman, Richard S., 185
beani, Ceramopora. See Ectoprocta: Cystoporata body fossils, 6, 143, 203, 204, 212, 281, 293. See also taphon-
Bear Creek K-bentonite "zone." See K-bentonite, K-bentonite omy, preservation
beds, K-bentonite "zones" Bolivia, 200
Bear Creek Quarry. See Ohio: Clermont C o u n t y bony fish. See Chordata: Vertebrata: Osteichthyes
bedding index, 49 Boone County, Kentucky, 87, 110, 111, 142, 180
Bedford, Indiana, 10 boring, borings, 74, 7 5 , 7 7 , 9 0 , 9 3 , 119, 123-125, 130, 182,
behavior (of animals). See chapters on individual groups of 203, 206, 210, 211, 236, 242, 243, 244, 293. See
a n i m a l s ; taphonomy: trace fossils also taphonomy: trace fossils
Bell, Bruce M., 180, 189,251 Bouchard, Timothy D., 206,'304
Bell, C M . , 212 bowdeni, Loxoplocus. See Mollusca: Gastropoda: Paupospira
bellerophontids. See Mollusca: Gastropoda bowdeni, Paupospira. See Mollusca: Gastropoda
Bellevue Hill Park, C i n c i n n a t i , Ohio, 21 Bowsher, Arthur L., 121
Bellevue House, C i n c i n n a t i , Ohio, 21 Brachiopoda, 6, 7, 31, 3 2 , 4 0 , 4 1 , 52, 60, 75, 77, 79, 8 7 , 8 8 , 89,

Index 325
 9 2 , 9 3 , 9 8 - 1 1 5 , 119, 123, 124, 125, 127, 128, Ungulate brachiopods, lingulides, 103, 105, 247, 248
129, 130, 137, 144, 162, 172, 178, 179, 182, 188, Petrocrania: P. scahiosa, 40, 104, 105
189, 206, 219, 220, 221, 223, 224, 225, 231, 232, Philhedra: P. laelia, 104, 105, 108
233, 235, 237, 240, 241, 242, 243, 244, 245, 248, Pseudolingula, 104, 105, 225. See also Lingula
250, 251, 252, 253, 260, 261, 272, 274, 280, 281, Schizomania: S. filosa, 104, 105
283, 285, 287, 291, 293 Trematis, 128
Articulata, 40, 93, 100, 101, 102, 103, 105, 112, 130, 261, T. millepunctata, 4 0 , 104, 105
283,291,293 Brachiospongia. See Porifera
Austinella, 112 brachycephalus, lsotelus. See Arthropoda: Trilobita
A. scovillei, 273 Bracken County, Kentucky, 178, 209
Catazyga, 112 Bradshaw, Lael E., 22, 24, 25
C. schuchertana, 111 Brandt, Danita S., 151, 153
Dalmanella, 57, 113, 114, 225, 232, 233, 240, 253 Branson, C . C , 196
D. emacerata, 106 Branson, E. B., 196, pl. 5
Glyptorthis, 112, 220, 224, 233 Branstrator, Jon W., 182, 184, 189
G. insculpta, 107 Braun, E. Lucy, 260
Hebertella, 100, 104, 105, 113, 223, 233, 240, 251 Braun, Frederick, 2 6 0 - 2 6 1 , 270
H. occidentalis, 107 Bretsky, Peter, 231
Hiscobeccus, 112, 220, 224, 240 Brett, Carlton E., 60, 6 1 , 236, 295, 297, 310, 311, 318
H. capax, 111 breviceps, Tricopelta. See Arthropoda: Trilobita: Phacopida
Holtedahlina, 1 1 2 , 2 2 0 , 2 4 0 Bridge, Josiah, 261
Lepidocyclus, 75, 220 British M u s e u m (Natural History). See Natural History Mu-
Leptaena, 40, 112, 220, 224, 233, 240 se um, London
L. richmondensis, 40 brittle stars. See Echinodermata: Ophiuroidea
Onniella, 112, 124 Broaddus, Billie, 29
O. meeki, 60 Brongniart, Alexandre, 61
Orthis: O. scovillei (See Austinella scovillei) Brookville Formation, 44
Orthorhynchula, 112 Excello M e m b e r , 44
Plaesiomys, 108, 112, 220, 224, 233 Liberty Member. See Liberty Formation, M e m b e r
P. subquadrata, 108 Station Hollow M e m b e r , 44
Platystrophia, 109, 112, 113, 223, 224, 232, 233,240, Waynesville M e m b e r . See Waynesville Formation
250,251,252 Brookville, Franklin County, Indiana, 184,196
P. acutilirata, 109 Brown County, Ohio, 123, 128, 151
P. laticosta, 109 Brown, Roland Wilbur, 40
P. ponderosa, 39, 59, 98. 99, 109, 240, 250 Brown, Steve, 155, 156, 157
P. ponderosa auburnensis, 39 "bryoherm." See bioherm; Ectoprocta
P. ponderosa ponderosa, 39 bryozoan. See Ectoprocta
Plectorthis, 1 1 2 , 2 2 4 , 2 4 0 Bucher, Walter H., 24, 4 8 , 123
P. neglecta, 106 Bull Fork Formation, 44, 214
Rafinesquina, 4 0 , 59, 75, 8 7 , 8 8 , 8 9 , 9 9 , 104, 105,110, Burgess S h a l e , 6, 249
112, 113, 114, 189, 223, 224, 232, 233, 235, 240, burrows, burrowing. See taphonomy: trace fossils, ichnofos-
251,252,272 sils, Lebensspuren
R. alternata, 110 Butcher, W i l l i a m , 320
"shingled Rafinesquina beds," 59, 9 9 , 110, 114, 251 Butler County, Ohio, 6 8 , 75, 80, 87, 89,107, 108, 126, 127,
Retrorsirostra, 112, 240 134, 137, 153, 156, 160, 176, 195,229
R. carleyi, 108, 261 Byrnes, Richard M., 261
Rhynchotrema, 112, 220 Byronia. See Byroniida, byroniids
R. dentatum, 111 Byroniida, byroniids, 182, 246. See also Phosphannulus
Sowerbyella, 112, 113, 225, 232, 240, 253 Byronia, 182
S. rugosa, 106 byssus, byssal threads. See Mollusca: Pelecypoda
Strophomena, 112, 113, 114, 224, 232, 233, 240, 247,
252, 261 C I , C 2 , C 3 , C 4 , C 5 , C 6 sequence, 55-56, 63, 214, 231,
Tetraphalerella, 112 232-233, 240. See also sequence stratigraphy
Thaerodonta, 112 Calapoecia. See C n i d a r i a : Anthozoa: Tabulata
7'. rugosa, 40 calcareous. See c a l c i u m carbonate
Thecidellina, pl. 3 calcite, calcific. See c a l c i u m carbonate
Zygospira, 41, 112, 232, 233, 245, 251, 252 c a l c i u m carbonate, 6, 50, 67, 71, 101, 102, 117, 119, 134, 144,
Z. modesta, 111, 178, 179 147, 167, 281
I n a r t i c u l a t a , 4 0 , 9 3 , 101, 102, 103, 108, 115, 128, 225 aragonite, aragonitic, 6, 101, 117, 120, 129, 134, 280, 281,
Leptoholus, 225 283, 290
calcareous, calcareous shell, calcareous skeleton, 3, 56,

Lingula, 5, 103. See also Pseudolingula

326 Index
 6 0 , 6 7 , 7 1 , 72, 79, 13?, 144, 145, 169, 188, 206, Charactoceras. See Mollusca: Cephalopoda
239, 280, 2 8 1 , 2 8 9 , 2 9 2 C h e e t h a m . A l a n H . , 185,298
calcite, calcitic, 6, 7,74, 89, 101, 117, 120, J24, 130, 137, Cheilostomata. See Ectoprocta
139, 163, 167, 168, 188, 280, 281, 283, 290 Cheirocystis. See Echinodermata: Rhombifera
calcium phosphate, 6, 81, 102, 182, 196, 198, 219, 223 Chelicerata. See Arthropoda
Calloway Creek Formation, 214 C h i c a g o , University of. See University of C h i c a g o
Calymene meeki. See Arthropoda: Trilobita: Flexicalymene Chilopoda. See Arthropoda
meeki chitin, chitinous, 6 , 6 9 , 82, 119, 147, 148, 161, 181, 282. See
C a l y m e n i d a , calymenids. Sec Arthropoda: Trilobita also eonchiolin; pseudochitin
Calvptomatida. See hyolithids; Mollusca chitinozoans, 52, 68, 69, 247, 282. See also Algae
C a m b r i a n , xix, 2 , 4 , 6 , 30, 8 2 , 9 9 , 139, 147, 152, 160, 163, Conochitina: C. hirsuta, 66, 67
167, 168, 182, 186, 191, 195, 199, 237, 249, 280, Cyathochitina, 66, 67; C. campanulaeformis, 66, 67
282, 289 Hercochitina: H. turnbulli, 6 6 , 67
" C a m b r i a n Explosion," 2 chitons. See Mollusca: Polyplacophora
"Cambrian Fauna," 2, 237 Chlorophyta. See Algae
Camerata. See Echinodermata: Crinoidea Chondrites. See taphonomy: trace fossils
Cameroceras. See Mollusca: C e p h a l o p o d a : Endoceratoidea Chordata, 38, 169, 196, 198, 282. See also conodonts
camouflage, 9 6 , 280 Urochordata: tunicates, 210, 281. See also Catellocaula
campanulaeformis, Cyathochitina. See chitinozoans Vertebrata
C a m p b e l l County, Kentucky, 59, 109, 203, 209 Agnat ha ("jawless fish"), 199, 200
C a m p b e l l , Kenton S. W., 155, 160 Astraspis: A. desiderata, 199
Canada, 3,215,220, 233,249,253 gnathostome fish ("jawed fish"), 200
Anticosti Island, 7 2 - 7 3 Osteichthyes ("bony fish"), 198
British C o l u m b i a , 6 Reptilia: Virginia, 41
C a n a d i a n Shield, 3, 216 Christy, David, 262
Manitoba, 153 christyi, Platycoryphe. See Arthropoda: 'Trilobita: Phacopida
canadensis, Crewingkia. See C n i d a r i a : Anthozoa: Rugosa chronostratigraphic units. See time-stratigraphic units
C a n a d i a n System, 30, 31 C i n c i n n a t i Arch, 8, 9, 10, 12, 59, 79, 216, 2 2 0 - 2 2 6 , 268, 282.
cancellatus, Sinuites. See Mollusca: Monoplacophora See also Findlav Arch; Kankakee Arch
capax, Hiscobeccus. See Brachiopoda: Articulata C i n c i n n a t i Group, 28, 47, 143, 266, 269, 277
Carboniferous, xix, 4, 23, 191, 198, 261, 270 C i n c i n n a t i Quarterly Journal of S c i e n c e , x, 17, 24, 267, 270
Caritodens. See Mollusca: Pelecypoda C i n c i n n a t i School of Paleontology, 15-36
Carley, S. T., 261 C i n c i n n a t i , Ohio
carfeyi, Retrorsirostra. See Brachiopoda: Articidata Boudinot Avenue and Westwood Northern Boulevard, 151
cadeyi, Rusophycus. See taphonomy: trace fossils C i n c i n n a t i Astronomical Society, 21
Carlson, Sandra J . , 192, 321,322 C i n c i n n a t i C o l l e g e of Pharmacy, 22
Carneyella. See Echinodermata: Edrioasteroidea C i n c i n n a t i Historical Society, 21, 215
carpoids. See Echinodermata C i n c i n n a t i Taw C o l l e g e , 24
Carr, T i m , 54 C i n c i n n a t i M u s e u m C e n t e r ( C M C ) , ix, xv, 21, 72, 75, 85,
Carriker, Melbourne R., 124 87, 89, 9 0 , 9 5 , 105, 110, 111, 120, 123, 126, 134,
casei, Opisthoptera. See Mollusca: Pelecypoda 137, 142, 144, 151, 153, 155, 157, 158, 159, 160,
casei, Plicodendrocrinus. See Echinodermata: Crinoidea: 170, 172, 173, 174, 178, 180, 186, 203, 207, 208,
Cladida 209, 229, 256, 263, pl. 8
Caster, Kenneth E., 25, 27, 28, 30, 31, 3 5 , 4 8 , 9 3 , 138, 151, C i n c i n n a t i M u s e u m of Natural History. See C i n c i n n a t i
158, 159, 162, 163, 186, 191, 209, 212, 215, 259, Society of Natural History
275, 276, 299, 320, pl. 13 C i n c i n n a t i Normal School, 27, 262
Catazyga. See Brachiopoda: Articulata C i n c i n n a t i Observatory, 21
Catellocaula, 9 3 , 1 5 6 , 209, 210, 243, 281. See also bioclaus- C i n c i n n a t i Society for the Promotion of Useful Knowl-
tration; bioimmuration; Chordata: Urochor- edge, 259, 261, 276
data: tunicates C i n c i n n a t i Society of Natural History, 15, 16, 21, 22, 23,
C. vallata, 156, 209, 210, 281 24, 26, 27, 28, 29, 30, 32, 33, 34, 259, 260, 261,
Cenozoic Era, xix, 25, 237, 238, 248, 284 2 6 2 - 2 6 3 , 264, 265, 2 6 6 , 267, 268, 270, 272,
centipedes. See Arthropoda: Chilopoda 275, 276, pl. 13
Cephalopoda. See Mollusca Journal of the Cincinnati Society of Natural History, x,
Ceramopora. See Ectoprocta: Cystoporata 17, 259, 276
Ceratopsis. See Arthropoda: C r u s t a c e a : Ostracoda Eden Park Reservoir, 29
Ceraurinus. See Arthropoda: Trilobita: Phacopida M e d i c a l C o l l e g e of Ohio, 29
Ceraurus. See Arthropoda: Trilobita: Phacopida Ohio M e c h a n i c s Institute, 31, 262
Chaetognatha, 198, 282 Spring Grove Cemetery, 22, 23, 24, 25
chambersi, Ceratopsis. See Arthropoda: Crustacea: Ostracoda Woodward High School, 18, 26, 32, 33, 34
Chappars, Michael S., 25 C i n c i n n a t i , University of. See University of C i n c i n n a t i

Index 327
Gincinnaticrinus. See Echinodermata: C r i n o i d e a : Disparida polyps, 74, 7 7 , 7 9 , 182, 206, 235
Cincinnatidiscus. See Echinodermata: Edrioasteroidea reefs. See biohcrms
cincinnatiensis, Acidaspis. See Arthropoda: Trilobita: Scyphozoa, 74, 82, 182. See also Byroniida
Odontoplcurida C o n u l a r i i d a , 74, 8 1 - 8 2 , 241, 247; but may not be cni-
cincinnatiensis, Diplocraterion. See taphonomy: trace fossils darians, 74, 8 1 - 8 2 , 241, 247
cincinnatiensis, Isorophus. See Echinodermata: Conularia
Edrioasteroidea C. formosa, 80, 82, 267, 269
circular is, Ischadites. See Algae: Chlorophvta: Dasvcladaceae C. miseneri, 270
C l a d i d a . See Echinodermata: C r in o idea sea anemones. See Anthozoa
c h i l l i s . See Mollusca: Pclccvpoda sea fans, 74
Clark, T h o m a s H., 3 sea whips, 74
Clarkson, Euan N. K., 160, 299 coarsening-upward cycle. See cycles, cyclicity
Clarksville M e m b e r . See Waynesville Formation coccinea, luhastraea. See C n i d a r i a : Anthozoa: Scleractinia
class. See taxonomy: l . i n n a e a n Hierarchy cockles. See Mollusca: Pelecypoda
c l . i s s i h i ation. Sec lavonoim C o d e of Stratigraphic Nomenclature. See stratigraphy
clastic ratio, 49. See also shale-to-limestone ratio (loclenterata. Sec I Inidaria
clavacoidea: Leptotrypa. See Ectoprocta: Trepostomata coiling of shell, 117, 119, 120, 132, 134, 139
C l a y s Ferry Formation, 163, 214 planispiral coiling, 119, 120, 139, 289
C l e r m o n t C o u n t y , Ohio, 111, 126, 128, 153, 158, 163, 170, C o l b a t h , G. Kent, 67, 69
186, 203, 207, 208, 209, 261 C o l d Spring, Kentucky, 174
Stonelick C r e e k , 11,254 Goleolus. See hyolitbids
Climacograptus. See Hcmichordata: Graptoloidea collagen, 195, 196
Clinton County, Ohio, 60, 72, 75, 128, 136, 142 colonial organisms, colonies, 189, 281, 290. See C n i d a r i a ,
W i l m i n g t o n , 260, 273, 276 Ectoprocta, graplolites
cluster analysis. See techniques color patterns, 9 6 , 134, 135, 137, 252, pl. 6
C M C . See C i n c i n n a t i , Ohio: C i n c i n n a t i M u s e u m C e n t e r Colorado, 199, 216
C M C IP, See C i n c i n n a t i , Ohio: C i n c i n n a t i M u s e u m C e n t e r C o l u m b i a University Press, 112
C n i d a r i a , 22, 32, 3 8 , 4 1 , 7 1 , 72, 7 3 - 8 2 , 8 5 , 9 1 , 9 3 , 122, 123, C o l u m b i a n University See George Washington University
131, 182, 196, 206, 219, 220, 221, 224, 233, 235, Gomactinia. See Echinodermata: Crinoidea
240, 241, 242, 243, 247, 248, 281, 285. See also Comasterida. See Echinodermata: Crinoidea
Byron iida c o m m e n s a l i s m . See ecologic associations
a n e m o n e s . See Anthozoa c o m m u n i t i e s . See ecology, ecologic needs: ecosystems
Anthozoa, 74, 77 competition, competitive interactions. See ecologic
a n e m o n e s , sea a n e m o n e s , 73, 9 6 , 188 associations
Rugosa, 74, 75, 7 7 , 7 8 , 7 9 , 2 0 6 , 220, 221, 235, 241, 247 composite molds. See taphonomy: molds
Gvathophvlbides, '<>. ~~. 79 conchiolin, 119, 282. See also chitin
C. stellata, 78 cones (the snails). See Mollusca: Gastropoda: Neogastropoda
Grewingkia, 41, 77, 206, 220, 224, 235, 242 C o n n e c t i c u t A c a d e m y of Arts and Sciences, 149
G. canadensis, 74, 75, 77 Conochitin. See chitinozoans
Streptelasma, 11, 220, 224, 235, 240 conodonts, 6, 28, 31, 35, 5 2 , 6 3 , 143, 183, 195, 196-199, 247.
S. divaricans, 75, 77 See also Cbordata
Scleractinia Amorphognathus: A. ordovicicus, 196, 197
luhastraea: T. coccinea, pl. 3 Aphebgnathus: A. grandis, 196, 197
Tabulate, 77, 79, 8 0 , 1 2 3 , 220, 221, 224, 247 Drepanoistodus: D. suherectus, 196, 197
Galapoecia, 76, 7 7 , 7 9 Oulodus: O. oregonia, 196, 197
Foerstephyllum, 76,11, 79 Phragmodus: P. undatus, 196, 197, pl. 5
F. vacuum, 78 Plectodina: P. tenuis, 196, 197
Nyctopora, 76, 77, 79 Prioniodus: P. dychei, 28, 277
Protaraea: P. richmondensis, 76,11, 79, 122, 123, 224, Rhodesognathus: R. elegans, 196, 197
240 C o n r a d , T i m o t h y A., 109, 110, 111, 120, 123, 126, 127, 149
Tetradium, 76,11, 79, 220, 233, 240 Gonstellaria. See Ectoprocta: Cystoporata
hydroids. See Hvdrozoa constellata, Gonstellaria. See Ectoprocta: Cystoporata
Hydrozoa, 74, 93, 242, 245 constrictus, Dystactocrinus. See Echinodermata: Crinoidea:
fire coral, 74 Disparida
hydroids, 74, 281 consumers. See ecology: ecosystems
Siphonophorida Gonularia. See C n i d a r i a : Scyphozoa: C o n u l a r i i d a
Portuguese Man-of-War, 74 C o n u l a r i i d a , conulariids. See C n i d a r i a : Scyphozoa
Porpitidae: Palaeoscia: P. ftoweri, 209, 212. See also convergent evolution. See organic evolution
taphonomy: trace fossils (lomvav Morris, S i m o n . 249
jellyfish. See medusa Cook, W. E., 263
m e d u s a , jellyfish, 7 1 , 74, 182, 209, 212, 251 Cooper, Dan L., 153, 309, pl. 6

328 Index
Cooper, Edward M , 263 eustatic sea-level c h a n g e , 55, 285
coprolites. See feces fining-upward cycle, fining-upward packet, 59
coprophagy. See food m e g a c y c l e , 55, 59, 288
copulation, 162 meter-scale cycles, 55, 59, 61
coquina, 99, 283. See also shell pavements; shingled beds; Milankovitch Cycles, 64, 288
Waynesville Formation: Marble Hill bed shoaling-upward cycles, 53
coral. See Cnidaria storm cycles, 55, 56, 292
coral reef. See bioherm Cycloconcha. See Mollusca: Pelecypoda
Corallidomus. See Mollusca: Pelecypoda Cyclocrinites, See: Algae: Chlorophyta: Dasycladaceae
coralline sponges. See Porifera: sclerosponges Cyclocystoidea. See Echinodermata
Cornulites, Sec: Annelida Cyclonema. See Mollusca: Gastropoda
correlation, 52. See also time Cyclora. See Mollusca: Gastropoda
Corryville Formation, M e m b e r , 12, 21, 53, 55, 60, 63, 68, 80, Cyclostomata. See Ectoprocta
90, 105, 110, 111, 114, 120, 126, 128, 142, 144, Cylindrocoelia. See Algae
151, 153, 158, 163, 170, 174, 175, 178, 180, 186, Cymaronora. See Mollusca: Pelecypoda
203, 207, 208, 209, 211,214, 224, 251. See also: C y n t l u a n a Formation, 123
Grant Lake Limestone; also: M c M i l l a n Cyrroceras. See Mollusca: C e p h a l o p o d a
Formation Cyrtodontula. See Mollusca: Pelecypoda
cotypes. See taxonomy: type specimens Cyrtolites. See Mollusca: Monoplacophora
Covington Croup, 48 Cystaster. See Echinodermata: Edrioasteroidea
Covington, Kentucky, 19, 28, 127, 142, 256, 263, 266, 274 ( A s t o i d c a . Sec Echinodermata

covmgroiiensis, Cylindrocoelia. See Algae cystoids. See Echinodermata

C o w e n . Richard, 199
crabs. See Arthropoda: Crustacea: Malacostraca Dalmanella. See Brachiopoda: Articulata
crateriformis, Lichenocrinus. See Echinodermata: Crinoidea: Dalve, Elizabeth A., 48, 230, 238, 263
Disparida d a m a g e (in life). See injuries
Crawfordsville, Indiana, 23, 261, 270 Dana, James Dwight, 275
Hovey M u s e u m , Wabash College, 23 Darwin, C h a r l e s , 61
Cretaceous, xix, 2, 3,4, 72, 239, 292 darwini, Cyclocrinites. See Algae: Chlorophyta:
Cricoconarida. See also Mollusca; Tentaculitoidea Dasycladaceae
Crinoidea. See Echinodermata darwini, Pasceolus. See Algae: Chlorophyta: Dasycladaceae:
crosotus, Primaspis. See Arthropoda: Trilobita: Cyclocrinites
Odontopleurida Dasycladaceae. See Algae: Chlorophyta
Crustacea. See Arthropoda dating. See time
Cruziana. See taphonomy: trace fossils Dattilo, B e n j a m i n E, 59, 60, 6 1 , 113, 231, 232, 308, 312
cryptolithi, Rusophycus. See taphonomy: trace fossils Davis, Richard Arnold, ix, 14, 89, 212, 229, 249, 255, 298,
Cryptolithus. See Arthropoda: 'Trilobita 304,320
Cryptostomata. See Ectoprocta D C A . See techniques: detrended correspondence analysis
Ctenobolbina. See Arthropoda: Crustacea: Ostracoda ( D C A)
Ctenodonta. See Mollusca: Pelecypoda Dearborn County, Indiana, pl. 3, pl. 5
Ctenostomata. See Ectoprocta death assemblage, 7, 8, 230, 283, 287. See also life assem-
cubichnia. See taphonomy: trace fossils: behavior categories blage; taphonomy
Cucumaria. See Echinodermata: Holothuroidea decadactylus, Glyptocrinus. See Echinodermata: C r i n o i d e a :
Cuffey, Roger J . , 9 1 , 95, 296, 302 Camerata
cultrata, Ambonychia. See Mollusca: Pelecypoda decameter-scale cycles. See cycles, cyclicity
Cupulocrinus. See Echinodermata: Crinoidea: C l a d i d a Deceptrix. See Mollusca: Pelecypoda
Curacao, Netherlands Antilles, pl. 3 Decoroproetus. See Arthropoda: Trilobita
curtus, Merocrinus. See Echinodermata: Crinoidea: C l a d i d a decorus, Neocrinus. See Echinodermata: Crinoidea
curvatum, Gorbyoceras. See Mollusca: Cephalopoda deformation (in life), deformities. See pathology
cuttlefish. See Mollusca: Cephalopoda deformation (post-mortem). See taphonomy
Cuvier, Ceorges, 61 Deiss, C h a r l e s Frederick, 263
Cyanobacteria delicatum, Oncoceras. See Mollusca: Cephalopoda
("blue-green 'algae'"), 67 demissa, Caritodens. See Mollusca: Pelecypoda
Cirvanella, 67 demissa, Pterinea. See Mollusca: Pelecypoda: Caritodens
stromatolites, 67, 292 demissa
Cyanoplryta. See Algae Dendroidea. See Hemichordata
Cyathochitina, See: chitinozoans Dent, Ohio. See Ohio: Hamilton C o u n t y
Cyathophylloides. See C n i d a r i a : Anthozoa: Rugosa dentatum, Rhynchotrema. See Brachiopoda: Articulata
cycles, cyclicity, 52-53, 55-57, 5 8 - 5 9 dentition. See Brachiopoda
coarsening-upward cycle, 59 deposit feeding. See food
decameter-scale cycles, 59 depositional sequences, 52-53, 55-57, 5 8 - 5 9

Index 329
desiderata, Astraspis. See Chordata: Vertebrata: Ag n ath a Dystactophycus. See incertae sedis
detrended correspondence analysis (DCA). See techniques
Devonian, xix, 4, 9, 62, 72, 79, 144, 151, 186, 191, 239, 251 earthquakes, 7, 61, 291. See also seismites
diagnosis. See taxonomy: L i n n a e a n Hierarchy earwigs. See Arthropoda: Insecta
Dickhaut, H. E., 263 eatoni, Triarthrus. See Arthropoda: Trilobita
dickhauti, Lepidolites. See Algae: Chloropbyta: eccentricity (in Earth's orbit). See cycles: Milankovitch Cycles
Dasycladaceae ecdysis, 6 , 9 2 , 9 3 , 117, 147, 148
Diekmeyer, Sharon C. St. Louis, 232, 307 Echinodermata, 24, 26, 144, 145, 1 6 6 - 1 9 2 , 2 1 9 , 2 2 3 , 233,
Diestoceras. See Mollusca: Cephalopoda 237,241,245,246,260,269
Dill Robert F, 54 Asteroidea, 24, 167, 168, 169, 182,183, 184, 185, 186, 188,
Dillsboro Formation, 44, 51 189-190, 208, 210, 235, 241, 247, 248, 259, 260,
d i m o r p h i s m , sexual, 153, 283, 290 265, 266, 274
dinoflagellates. See Algae: Pyrrophyta Asteriacites (See taphonomy: trace fossils)
"dioramas," pl. 13 Fromia: F. nodosa, pl. 9
diplobathrid camerates. See Echinodermata: Crinoidea: Goniasteridae, 189
Camerata Hudsonaster, 189; H. simp/ex, 185, 274
Diplocraterion. See taphonomy: trace fossils Lanthanaster, 189; L. intermedius, 184
Diplopoda. See Arthropoda Ophidiasteridae, 189
disarticulation of hard parts. See taphonomy Palaeaster: P. simplex (See Hudsonaster simplex)
Disparida. See Echinodermata: C r in o idea Palaeasterina: P. approximata, 259, 265; P. speciosa,
divaricans, Streptelasma. See C n i d a r i a : Anthozoa: Rugosa 260,266
diversity, species diversity, taxonomic diversity, 2, 3, 4, 57, 6 1 , Petraster: P. speciosus, 184, 189
69, 103, 113, 115, 118, 120, 130, 131, 137, 147, Promopalaeaster, 189, 235
1 6 1 , 1 6 3 , 1 6 7 , 204, 232, 234, 239, 241, 283 P. dyeri, 184
d'Orbigny, Alcide, 85, 9 0 , 9 5 , 271 P. magnificus, 182, 183
dolomite, 3, 53, 217, 221, 222, 283 P. speciosus, 182, 183
d o m i c h n i a . See taphonomy: trace fossils: behavior categories Salteraster, 189; S. grandis, 184
Donovan, Stephen K., 186 carpoids, 145, 168, 186, 275. See Stylophora
Dorfeuille, Joseph, 263, 272, 274, 276 C r i n o i d e a , 7, 23, 24, 2 8 , 4 3 , 56, 57, 93, 120, 121, 123, 137,
"doughnuts." Sec "W'cichold doughnuts" 144, 1 6 2 , 1 6 6 , 167, 168, 169-178, 180, 181-183,
Drake, Daniel, 16, 2 6 3 - 2 6 4 , 268, 273, 276 186, 188, 212, 223, 224, 225, 226, 231, 232, 236,
Drakes Formation, 44, 138 237, 241, 243, 245, 246, 247, 248, 249, 251, 252,
Preachersville M e m b e r , 214 253, 261, 270, 271, 285
Rowland M e m b e r , 127, 214 C a m e r a t a , 178, 180
Drepanoistodus. See conodonts diplobathrid camerates, 176, 177, 180
drilling. See borings Caurocrinus: G. nealli, 176, 177, 180
Droser, M a r y L., 1,237, 321 monobathrid camerates, 175, 176, 180
Dry Dredgers, 255, 263, 275, 300, 302, 317 Clyptocrinus, 4 2 , 4 3 , 120, 178, 224, 2 3 2 , 2 4 0 , 241,
dubius, hichenocrinus. See Echinodermata: Crinoidea: 252
Disparida G. annularis, 272
d u m b e l l s . See taphonomy: trace fossils: Diplocraterion G. decadactylus, 42, 171, 172, 175, 180
duncanae, Gorbyoceras. See Mollusca: Cephalopoda G. dyeri. See Pycnocrinus dyeri
dunni, Narthecoceras. See Mollusca: C e p h a l o p o d a G. fornshelli, 176, 180
durophagy. See ecologic associations: predation: G. pattersoni, 272
Durrell, Richard H., ix G. richardsoni, 273, 276
Dury, C h a r l e s , 105 Pycnocrinus, 2 4 , 4 3 , 120, 123, 178, 180,224
Dury, Ralph, 105 P. dyeri, 24, 3 7 , 4 0 , 4 2 , 4 3 , 1 7 5 , 176
duseri, Treptoceras. See Mollusca: Cephalopoda: Xenocrinus, 180, 240, 241
Actinoceratoidea X. baeri, 176, 177, 228, 229
Dyche, D. T. D., 28, 277 C l a d i d a , 178
dychei, Prioniodus. See conodonts Cupulocrinus, 178
Dyer, C. B., 14, 2 3 - 2 4 , 29, 31, 3 7 , 4 0 , 80, 82, 184, 185, 208, C. polydactylus, pl. 11
210, 260, 261, 265, 266, 272, 274 Merocrinus, 178
dyeri, Allocotkhnus. See taphonomy, preservation: trace M . curtus, 172, 176,177
fossils Plicodendrocrinus, 178
dyeri, Clyptocrinus. See Echinodermata: C r i n o i d e a : C a m - P. casei, pl. 11
erata: Pycnocrinus Comactinia, 166
dyeri, Promopalaeaster. See Echinodermata: Asteroidea Comasterida, 166
dyeri, Pycnocrinus. See Echinodermata: C r i n o i d e a : Disparida, 173, 174, 178, 186
Camerata Anomalocrinus, 178, 252
Dysracrocr/nus. See Echinodermata: Crinoidea: Disparida A. incurvus, 172, 174

330 Index
 Cincinnaticrinus, 113, 178, 182, 186, 226, 2 3 2 , 2 4 0 , facultative, 285, 288
241,253 inquiline, i n q u i l i n i s m , 286. See also incolent
C. pentagonus, 173 m u t u a l i s m , 234, 236, 288, 290, 292
C. varihrachialus, 172, 173 obligate, 123, 285, 288
Dystactocrinus: D. constrictus, 173, 183 parasite, parasitism, 8 6 , 94, 123, 182, 223, 229, 234, 236,
Ectenocrinus, 113, 178, 226, 2 3 2 , 2 4 0 , 241, 253 243, 246, 280, 281, 282, 285, 286, 288, 289,
E. geniculatus, 173 290, 292
E. simplex, 172, 173, PL 10 predation, 7 , 9 2 , 9 3 , 95, 123, 124, 125, 131, 132, 137, 143,
locrinus, 178, 224 147, 151, 153, 154, 155, 161, 162, 182, 183, 186,
/. oehanus, 271 189,190, 205, 210, 229, 234, 2 3 5 - 2 3 6 , 241, 282,
/. subcrassus, 110, 142, 170, 172, 174, PL 10 283, 284, 285, 289, 290, 291, 293. See also
Lichenocrinus, 178 boring
L. crateriformis, 172 durophagy, 131, 284
L. dubius, 186 symbiosis, 210, 235, 282, 284, 288, 289, 290, 292
L. milleri, 172 endosymbiont, 210, 284
Metacrinus, 182 ecologic succession, 56, 57, 115, 281, 292
Neocrinus: N. decorus, pl. 9 ecology, ecologic needs. See also chapters on individual
Pontiometra: P. andersoni, PL 9 groups of a n i m a l s ; ecologic associations;
Cyclocystoidea, 169, 190-191, 247 environment
7.ygocycloides: Z. magnus, 185 autecology, 229, 280, 284, 292
cystoids, 237. See Rhombifera ecosystems, x, 1, 67, 143, 222, 2 3 3 - 2 4 8 , 284
Echinoidea, 167, 238, 248 c o m m u n i t i e s , 4, 5 , 5 6 , 57, 169, 182,218, 230, 231, 237,
sand dollars, 167 241
sea urchins, 167, 168, 169, 237 consumers, 234, 282, 285, 290
Strongylocentrotus: S. franciscanus, PL 9 food c h a i n , food web, 67, 6 8 , 234, 282, 285, 290
Edrioasteroidea, 5 6 , 6 0 , 114, 169, 182, 186, 188-189, 190, guilds, 237-239, 247, 248, 286
241,243,247,250,251 m e g a g u i l d s , 237, 238
Carneyella, 178, 179, 224 populations, 188, 189
C. pilea, 180, 18J primary producers, 67, 234, 247, 248, 285, 290
Cincinnatidiscus, 40, 41 tiering, 182
Cysfasfer: C . stellatus, 180,181 paleoecology, 6 1 , 136, 188, 189, 230, 231
lsorophus, 189, 224 Economy M e m b e r . See Latonia Formation
J. cincinnatiensis, 40, J28, 180, 181, 189 ecosystems. See ecology
Streptaster, 110, 188, 2 2 4 , 2 5 0 Ectenocrinus. See Echinodermata: Crinoidea: Disparida
S.vorticellatus, 110, 180,181 Ectoprocta, 2 1 , 24, 30, 31, 33, 34, 35, 3 6 , 4 0 , 52, 57, 72, 75,
Eocrinoidea, 168 7 7 , 7 9 , 8 5 - 9 7 , 9 9 , 107, 110, 111, 113, 114, 115,
Holothuroidea, 167, 248 120, 125, 127, 128, 130, 131, 137, 139, 1 4 4 , 1 5 6 ,
Cucumaria: C. miniata, PL 9 162, 178, 180, 182, 188, 196, 2 0 6 , 2 0 9 , 210, 219,
Ophiuroidea, 6, 167, 168, 169, 189, 1 9 0 , 2 2 8 , 229, 247, 248 221, 223, 224, 225, 231, 232, 233, 237, 240, 241,
Ophiothrix, PL 9 242, 243, 244, 245, 247, 248, 250, 251, 252, 253,
Protasterina: P. flexuosa, 190 260, 265, 270, 271, 272, 273, 275, 281, 285, 287,
Taeniaster, 190 290,293
T. spinosus, 185, 190,228, 229 "bryoherms," 91
Rhombifera, rhombiferan "cystoids," 169, 186, 247 "Weichold doughnut," 88, 89, 9 3 - 9 4 , 275
Cheirocystis: C. fultonensis, 178, 179, 186 Cheilostomata, 237, PL 3
Lepadocystis: L. moorei, 178, 179, 186 Cryptostomata: Escharopora, 87, 88
Stylophora, 169, 186, 191-192, 247. See also "carpoids" Ctenostomata: Ropalonaria: R. venosa, 87, 88
Enoploura, 26, 145, 191, 192, 270, 275 {See also Cyclostomata, 107, 110
machaeridians) Cystoporata
E. balanoides, 191 Ceramopora; C. (?) beani, 260; C. nicholsoni, 271
E. popei, 186, 187, 191 Constellaria, 40, 87, 94
Echinoidea. See Echinodermata C. constellata, 275
ecologic associations. See chapters on individual groups of C. florida, 87, 88
animals Fistulipora: F. oweni, 272
aegism, 243, 244, 245, 280, 282, 288, 289, 290 Heteropora: H. pacifica, PL 3
c o m m e n s a l i s m , 95, 123, 144, 189, 229, 234, 236, 243, 244, Trepostomata, 79, 87, 88, 8 9 , 9 0 , 237, 250
245, 246, 280, 282, 286, 288, 289, 290, 292 Heterotrypa, 91
competition, competitive interactions, 182, 234, 236, 237, H.frondosa, 9 0 , 9 5 , 271
282 Eeptotrypa: L. clavacoidea, 120, 121
epizoa, epizoism, 9 3 , 9 4 , 9 6 , 103, 130, 182, 236, 242, 243, Monticulipora
244, 245, 284. See also encrustation M. falesi, 264

Index 331
 M. mammulata, 84, 85, 90, 271
Parvohallopora, 111, 252, 265, 270
P. onealli, 271
P. ramosa, 84, 85, 90, 271
P. rugosa (now included in P. ramosa), 265, 270
Spatiopora, 87, 88, 94
Eden Park, C i n c i n n a t i , Ohio, 29, 263
Eden Park Reservoir, 29
Eden S h a l e , Eden Formation, Eden Group, 47, 4 8 , 51, 52,
296, 301. See also Kope Formation; M i d d l e
(Eden) Shales
edenense, Veryhachium. See acritarchs
Edenian Age, Edenian Stage, 11, 44, 4 6 , 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, 2 0 6 , 235, 316, 321
Elkhorn Formation, 44, 56, 63, 68, 73, 75, 79, 158, 162, 172,
178, 186,214
emacerata, Dalmanella. See Brachiopoda: Articulata
Embree, Jesse, 264
encrustation, 56, 60, 64, 72, 74, 77, 79, 87, 88, 89, 90, 93, 94,
105, 107, 110, 114, 115, 1 2 0 , 1 2 2 , 123, 128,137,
139, 144, 178, 182, 196, 219, 223, 224, 231, 232,
236, 237, 240, 242, 243, 244, 245, 251, 286. See
also epicoles
endobyssate a n i m a l s . See Mollusca: Pelecypoda
F.ndoceras. See Mollusca: C e p h a l o p o d a : Endoceratoidea
Endoceratoidea, endocerids, endoceroids. See Mollusca:
Cephalopoda
endosymbiont. See ecologic associations: symbiosis
Enoploura. See Echinodermata: Stvlophora
environment, 3 - 4 , 5, 45, 52, 67, 69, 7 1 , 81, 189, 204, 2 1 0 - 2 1 1 ,
2 1 5 - 2 2 6 , 2 2 8 - 2 4 8 , 284, 292. See also chapters
on individual groups of a n i m a l s ; ecology, eco-
logic needs; facies; taphonomy: trace fossils:
ichnofacies
oxygen content, 160, 229, 250, 254
salinity, 217, 221, 229, 230
synecology, 229, 230, 280, 284, 292
temperature, 216, 217, 221, 229, 230
turbidity, 92, 229
water movement, 50, 81, 113, 114, 145, 169, 211, 229, 293
Eocarcinosoma. See Arthropoda: Chelicerata: Eurypterida
Eocene, xix, 4
Eocrinoidea. See E c h i n o d e r m a t a
Eoleperditia. SeeArthropoda: Crustacea: Ostracoda
eos, Diestoceras. See Mollusca: Cephalopoda
epeiric sea. See epicontinental sea
epibole. See biostratigraphic units: zone
epibyssate a n i m a l s . See Mollusca: Pelecypoda
epicoles, 93, 284. See also encrustation; epizoa
epicontinental sea, 3, 9, 231, 284
epifauna, epifaunal, 7, 56, 237, 247, 248, 284, 286
epireliefs. See taphonomy: trace fossils
epizoa, epizoism. See ecologic associations
E r i c k s o n . J . Mark, 9 1 , 2 0 6 , 320
Eriksson, M a t s , 143, 145, 307
F.scharopora. See Ectoprocta: Cryptostomata
Eurypterida. See Arthropoda: Chelicerata
332 Index
eustatic sea-level change. See cycles, cyclicity
event beds, event horizons. 5 9 - 6 1 . 224. 225, 285
evolution. See organic evolution
evolutionary faunas, sens!/ Sepkoski, 2, 237; "Modern
Fauna," 2. See also C a m b r i a n , "Cambrian
Fauna"; Paleozoic, "Paleozoic Fauna"
Excello Member. See Brookville Formation
excrement. See feces
exoskeleton, 6, 92, 93, 117, 119, 147, 148, 151, 155, 161,230,
285
extinction: mass extinction, 2, 240, 287
Faber, C h a r l e s , 14, 24, 2 7 - 2 8 , 32, 144, 172, 264, 273, 274,
275, 276
faberi, Technophorus. See Mollusca: Rostroconchia
Faberia. See Algae
facies. See also environment
biofacies, 5 9 - 6 1 , 2 8 1 , 285, 287
lithofacies, 5 9 , 2 8 1 , 2 8 5 , 287
factor analysis. See techniques
facultative association. See ecologic associations
fair-weather wave-base. See wave base: normal wave-base
Fairmount Member. See Fairview Formation
Fairview Formation, 12, 21, 48, 51, 53, 55, 56, 58, 59, 60, 61,
63, 68, 87, 106, 110, 120, 123, 127, 173, 175,
180, 196,214, 224, 225, 232, 251, 252, pl. 5, pl.
10. See also Hill Quarry Beds
la in i it mi it Member, 44
M o u n t Hope M e m b e r , Mt. Hope Member, 44
North Bend Tongue, 44
Fales, J . C , 264
falesi, Monticulipora. See Ectoprocta: Trepostomata
family. See taxonomy: L i n n a e a n Hierarchy
Fascifodina. See taphonomy: trace fossils
faunal provinces: American Midcontinent Province, 199;
North Atlantic Province, 199
feather stars. See Echinodermata: Crinoidea
feces, fecal, 121, 123, 162, 188, 219; coprolites, 203, 283
feeding. See food
Felton, Steven M 68, 120, 123, 127, 137, 140, 275, 314
Fenton, Carroll L a n e , 28, 123, 264
Fenton, Mildred A d a m s , 28, 123, 304
Field M u s e u m of Natural History ( F M N H ) , xv, 172, 184
filosa, Schizocrania. See Brachiopoda: Inarticulata
filter feeder, filter feeding. See food
Findlay Arch, Findlay Branch of Cincinnati Arch, 9. See also
C i n c i n n a t i Arch
Findlay, James, 264
Fine, Ronny L., 89, 9 0 , 9 1 , 3 0 2
fining-upward cycle, fining-upward packet. See cycles,
cyclicity
fire coral. See C n i d a r i a : Hydrozoa
fish. See Chordata: Vertebrata
Fisher, Daniel C, 192, 300
Fisher, Donald W., 144, 145
Fistulipora. See Ectoprocta: Cystoporata
flabellum, Licrophycus. See taphonomy: trace fossils
F l e m i n g County, Kentucky, 72
Flexicalymene. See Arthropoda: Trilobita
flexuosa, Protasterina. See Echinodermata: Ophiuroidea
Florida, pl. 9
florida, Constellaria. See Ectoprocta: Cvstoporata Caurocrinus. See Echinodermata: C r i n o i d e a : C a m e r a t a
Flower, Rousseau H., 43, 133, 134, 138, 295 G e e , Henry, 191
floweri, Fascifodina. See taphonomy: trace fossils generic n a m e . See taxonomy: nomenclature: Binomial Sys-
floweri, Palaeoscia. See C n i d a r i a : Hydrozoa: Siphonophor- tem of Nomenclature
ida: Porpitidae; taphonomy: trace fossils genkulatus, Ectenocrinus. See Echinodermata: C r i n o i d e a :
F M N H . See Field M u s e u m of Natural History Disparida
fodiniclmia. See taphonomy: trace fossils: behavior Ceniculograptus. See Hemichordata: Graptoloidea
categories genus, genera. See taxonomy: I.innaean Hierarchy
Foerste, August F., 18, 2 9 , 4 8 , 7 7 , 7 8 , 109, 149, 150, 151,153, geologic time-scale, xix
162, 207, 229, 264, 270, 271 geologic time-units, 4 5 - 4 6
Foerstephyllum. See C n i d a r i a : Anthozoa: Tabulata Geological Society of A m e r i c a , 5, 11, 30, 32, 36, 54, 59, 133,
food supply. See food 257, 2 7 1 , 2 7 4
food chain, food web. See ecology: ecosystems Penrose M e d a l , 30, 32
food, 4 , 6 7 , 6 8 , 7 1 , 74, 85, 8 7 , 9 2 , 101, 123, 151, 155, 160, 168, geopetal structures, 124
170, 172, 180, 188, 191-192, 198, 211, 229, 234, George Washington University (formerly C o l u m b i a n Univer-
235, 237-238, 239, 253, 280, 282, 283, 285, 286, sity), 22, 35
287, 290. See also chapters on individual Gibson, Bruce, 105, 176
groups of a n i m a l s ; ecologic associations: com- Gibson, Charlotte, 105, 176
mensalism; ecologic associations: predation; gigantea, Anomalodonta. See Mollusca: Pelecypoda
ecology: ecosystems gigas, Isotelus. See Arthropoda: Trilobita
coprophagy, 121, 123,282 Gilbert Formation, 214
deposit feeding, 123, 131, 140, 163, 190, 204, 2 0 5 , 2 1 1 , 2 2 4 , Girvanella. See Cyanobacteria
283, 285, 292 Clyptocrinus. See Echinodermata: Crinoidea: Camerata
filter feeder, filter feeding, 7 1 , 72, 87, 101, 127, 131, 144, Glyptorthis. See Brachiopoda: Articulata
155, 168, 172, 1 8 8 , 2 8 3 , 2 8 5 , 2 9 2 G o l d m a n , Daniel, 195
scavenging, 7, 151, 205, 223, 224, 225 G o n d w a n a . See paleogeography
suspension feeding, 21, 74, 123, 139, 163, 170, 182, 186, Goniasteridae. See Echinodermata: Asteroidea
190, 196, 205, 206, 211, 224, 237, 239, 283, 285, Gorbyoceras. See Mollusca: Cephalopoda
292 Grabau, A m a d e u s W., 18
Ford, John P., 21, 59 gracile striatulum, Cyclonema. See Mollusca: Gastropoda
formation. See lithostratigraphic units gracile, Cyclonema. See Mollusca: Gastropoda
formosa, Conularia. See C n i d a r i a : Scyphozoa: C o n u l a r i i d a gracile, Ordovicium. See acritarchs
fornshelli, Clyptocrinus. See Echinodermata: C r i n o i d e a : graded beds, 60
Camerata G r a h a m , George, 265
Fort Ancient M e m b e r See Waynesville Formation grainstone, 50, 55, 59, 114, 2 8 6 , 289
Fortey, Richard A., 1, 151, 155, 161, 303, 316 Grand Avenue Member. See Kope Formation
fossil succession. See biotal succession grandis, Aphelognathus. See conodonts
Foster, W i l l i a m , 3 1 , 2 6 4 grandis, Salteraster. See Echinodermata: Asteroidea
Fox, W i l l i a m T., 231 Grant Lake Formation, Limestone, 44, 79, 9 0 , 114, 123, 250
franciscanus, Strongylocentrotus. See Echinodermata: Corryville Member. See Corrvvillc Formation, M e m b e r
Echinoidea granulosa, Flexicalymene. See Arthropoda: Trilobita
Franklin County, Indiana, 105, 107, 157, 184, 196, pl. 6, pl. Graptoloidea, graptolites. See Hemichordata
10 Gravicalymene. See Arthropoda: Trilobita: C a l y m e n i d a
Franks' M u s e u m , 265 Great B a h a m a Bank. See B a h a m a Platform
Frey, Robert C, 60, 130, 131, 136, 137, 138, 297, 307 Great Barrier Reef, Australia, 9 1 , 241, pl. 3. See also bioherm
Fromia. See Echinodermata: Asteroidea Great Limestone Deposite, 47
frondosa, Heterotrypa. See Ectoprocta: Trepostomata Grewingk, C. C. A., 41
fucoids. See taphonomy: trace fossils Grewingkia. See C n i d a r i a : Anthozoa: Rugosa
Fulton Beds, Fulton Shale, 144, 186. See also Kope Grinndl, George Bird, 265
group. See lithostratigraphic units
Formation
growth lines, 75, 77, 117, 120, 2 8 6
Fulton, Robert, 265
guilds. See ecology: ecosystems
fultonensis, Cheirocystis. See Echinodermata: Rhombifera
Gunderson, G. O., 82, 309
fungi, 242
Gurley, W. F. E., 25, 157,313
Fusispira. See Mollusca: Gastropoda: Neogastropoda:
gutter casts, 60
Subulites

G a h n , Forest J . , 183, 297 H a i m e , Jules, 265, 270

G a m a c h i a n Stage, 214, 240 Haines, M. P., Mrs., 265
Garrard Formation, 214 Hall, J . H . , 2 5 9 , 265
Gastropoda. See Mollusca Hall, James (of C i n c i n n a t i ) , 265
Gault, W i l l i a m , 265 Hall, James (of New York), 32, 3 5 , 4 2 , 78, 105, 106, 107, 108,

Index 333
 111. 120, 12", 134, 137, 142, 145, 155, 157, 172, Holbrook, R. H., 267
173, 175, 176, 180, 183, 195, 207, 210, 229, 262, boles (in shells). See taphonomy: trace fossils: borings
265-266,272,276,277 Holland, Steven M., 53, 55, 5 9 , 6 3 , 113, 120, 130, 204,
Hall, John W., Jr., 2 5 9 , 2 6 5 , 2 6 6 2 1 5 - 2 2 6 , 229, 231, 232, 233, 241, 255, 312, pl.
11.ill.mi. Anthony 3, 4 12
halli, Alloliehas. See Arthropoda: Trilobita: Lichida I lolothuroidca. See Echinodermata
halli, Amphilichas. See Arthropoda: Trilobita: Lichida: Alloli- holotype. See taxonomy: type specimens
chas halli I loltedahlina. See Brachiopoda: Articulata
Hallopora. See Ectoprocta: Trepostomata: Parvohallopora homeomorphy. See organic evolution: convergent evolution
1 lamilton Comity, Ohio, 10, 8 9 , 9 0 , 106, 142, 151, 155, 156, horseshoe crabs. See Arthropoda: Chelicerata: Xiphosurida
173, 180, 186, 277, pl. 10 Hosea, L. M., 24, 267
'Trammel Eossil Park, Sharonville, 12, 124, 257 host, 77, 86, 87, 88, 9 3 , 9 4 , 103, 123, 144, 182, 189, 210, 234,
I l a m m o n d , W. E., 266 235, 236, 242, 243, 244, 245, 246, 282, 285, 286,
Hand, Greg, 306 288, 289
hardgrounds, 6 0 , 6 4 , 105, 124, 125, 178, 188, 203, 2 0 6 , 211, Hovey M u s e u m , Wabash College. See Crawfordsville,
219,250, 2 5 1 , 2 8 6 Indiana
I larding Sandstone, 199 Howe, A. J . , 267
Harper, George W., 17, 18, 26, 27, 32, 3 3 - 3 4 I [udson River C r o u p , 266, 269, 277
Harris, Frank W., 5 6 , 5 7 , 114, 115, 231 I ludsonaster. See Echinodermata: Asteroidea
Harris, G. D., 25 I luffman D a m . near Davton, ()hio, 153
Harris, I., 259, 266 Hughes, Nigel C , 82, 153
I larvard University, xv, 18, 23, 32, 260, 273, 274 humerosum. See Mollusca: Gastropoda: Cyclonema
M u s e u m of Comparative /oology ( M C Z ) , xv, 18, 23, 32, humerosum, Cyclonema. See Mollusca: Gastropoda
2 6 0 , 2 7 3 , 2 7 4 , pl. 13 Hunda, Brenda, 151
Harvey, J . W 260, 266 Hunter, W. H. H., 267
Haucr, Kendall, 311 huronensis, Labechia. See Porifera
Hay, Helen B 44, 53, 55, 56, 57 hurricanes. See storms
Hayden, F. V., 266, 268 I lutchinson, G. Evelyn, 229
Uebertella, S e e : Brachiopoda: Articulata Huxley, T. H., 267
I ledcen, Stanley. i\ hydroids. See C n i d a r i a : Hydrozoa
I [eimbrock, W i l l i a m , 67 I Ivdrozoa. hvdrozoans. See C Inidaria
Helcionopsis. See Mollusca: Monoplacophora I lyinan, L. H., 167
Hemichordata, 169, 196 Hyolithes. See hyolithids
Dendroidea, 195 hyolithids, 145, 247. See also Mollusca: Calyptomatida
Muslivograptus, 196 Coleolus, 145
graptolites, Craptoloidea, 52, 63, 161, J94, 195-196, 240, Hyolithes. 145
241,247,251 hyporeliefs. See taphonomy: trace fossils
Amheimograptus: A. anacanlhus. 196
Climacograptus. See Geniculograptw lapetus O c e a n , Sea. See paleogeography
V.enkuhgmptus, 196; C. typicalis. 194, 195 icarus, Ceraurinus. See Arthropoda: Trilobita: Phacopida
Orthograptus: (). quadrimucronatus, 196 Ice Age, ice cap, ice sheet, ix, 10, 53, 216, 217, 240, 269
Pterobranchia, 196; Rhahdopleura, 196 ichnofossils. See taphonomy, preservation: trace fossils
I lendy, Austin, xiv ichnogenus, ichnogenera. See taphonomy: trace fossils
I leran, M a g g i e , xiv ichnospeeies. See taphonomy: trace fossils
Hercochitina. See chitinozoans ICZN. See International C o d e of Zoological Nomenclature
I Icrrlinger, Andrew, 267 Illinois Geological Survey, 31
Herzer, Henry, 28, 30, 267 impedichnia. See taphonomy: trace fossils: behavior categories
Heteropora. See Ectoprocta inaequahile, Cameroceras. See Mollusca: Cephalopoda:
Heterotrypa. See Ectoprocta: Trepostomata Endoceratoidea
High Bridge C r o u p (middle Ordovician], 67. 139, 164 Inarticulata. See Brachiopoda
Tyrone I .iinestone. 139 incertae sedis, 211-212, 286. See also Palaeoscia
Highland County. Ohio, 151, 153 Blastophycus, 212
1 Mil Quarry Beds, 47, 225. See also Fairview Formation Dystactophycus, 212
Hill, H. H., 267 incolent, 286. See also ecologic associations
Himalayas, 3 incurvus, Anomalocrinus. See Echinodermata: Crinoidea:
Hints, Olle, 145 Disparida
Hirnantian Stage, 63, pl. 2 index fossils, 195, 255
lunula, Conochitina. See chitinozoans Indiana
Hiscoheccus. See Brachiopoda: Articulata Bedford, 10
Hitz bed, 214 Crawfordsville, 23, 2 6 1 , 2 7 0
Hogstrom, Anette E. S., 145, 307 Hovey M u s e u m , Wabash College, 23

334 Index
 Dearborn County, pl. 3, pl. 5 C a m p b e l l County, 59, 109, 203, 209, pl. 5
Franklin County, 105, 107, 157, 184, 196, pl. 6, pl. 10 Cold Spring, 174
Brookville, 184, 196 Covington, 19, 28, 127, 142, 256, 263, 266, 274
Indiana Geological Survey, 51, 214, 255, 263 F l e m i n g County, 72
Jefferson County, 111 Kenton County, 67, 87, 9 0 , 110, 123, 209
Madison, 7 7 , 7 9 , 2 2 1 , 2 6 6 , 2 6 9 Kentucky Geological Survey, 10, 51, 214, 256
Richmond, 47, 120, 265, 270, 272 Lexington, 40, 67, 123, 272
W a y n e County, 67, 75, 228 Transylvania C o l l e g e , University, 40, 272
infauna, infaunal, 7, 131, 139, 153, 204, 205, 237, 238, 239, Madison County, 77
247, 248, 284, 286 Marion County, 120
injuries, d a m a g e (in life), 137, 183, 186, 235, 236. See also Maysville, 11, 5 9 , 7 9 , 9 1 , 180
ecologic associations: predation Nelson County, 78
inquiline, inquilinism. See ecologic associations Trimble County, 124, 129
insculpta, Glyptorthis. See Brachiopoda: Articulata Bedford, 129
Insecta. See Arthropoda kentuckyensis, Rhytiodentalium. See Mollusca: Scaphopoda
intermedins, Lanthanaster. See Echinodermata: Asteroidea Kesling, Robert V., 178
International C o d e of Zoological Nomenclature ( I C Z N ) , xi Kiessling, Wolfgang, pl. 2
International Geological Correlation Programme, 63, 255 kingdom. See taxonomy: L i n n a e a n Hierarchy
International Stratigraphic C o d e . See stratigraphy Kjellesvig-Waering, Erik N., 158, 159, 162, 300
interspecific interactions. See ecologic associations Knell, Simon J . , 265
locrinus. See Echinodermata: Crinoidea: Disparida Kope Formation, II, 19, 21, 44, 49, 51, 52, 53, 55, 56, 57, 59,
Ischadites. See Algae: Chlorophyta: Dasycladaceae 6 0 , 6 1 , 6 7 , 6 8 , 6 9 , 106, 113, 120, 123, 127, 130,
Ischyrodonta. See Mollusca: Pelecypoda 142, 144, 149, 151, 155, 156, 160, 173, 176, 178,
isochronous surfaces. See time 180, 186, 190, 196, 203, 208, 209, 211,214, 219,
Isopoda. See Arthropoda: Crustacea 225, 231, 232, 253, pl. 5. See also Eden Shale;
Isorophus. See Echinodermata: Edrioasteroidea Latonia Formation
Isorthoceras. See Mollusca: Cephalopoda Fulton M e m b e r , 144, 186
isotopes, 62, 290. See also time: absolute age: radiometric Grand Avenue M e m b e r , 44
dating W e s s e l m a n Tongue, 4 4
Kozlowski, Roman, 196
Jacobson, Stephen R., 69 Kritsky, G e n e , 267
James Book Store, 14, 17, 22, 31
James, Joseph A., 17 Lahechia. See Porifera
James, Joseph R, 2 2 - 2 3 , 29, 203, 207, 210 laelia, Philhedra. See Brachiopoda: Inarticulata
James, U. P., 14, 17-22, 2 3 , 2 8 , 29, 31, 209, 210, 260, 261, lag deposit, 56, 114,287
264, 2 7 1 , 2 7 7 land plants. See Plantae, plants
jamesi, Lepidocoleus. See Machaeridia Lanthanaster. See Echinodermata: Asteroidea
Jefferson County, Indiana, 111 lata, Cyclonema hilix. See Mollusca: Gastropoda: Cyclonema
jellyfish. See C n i d a r i a : medusa hilix
Jennette, David C, 55, 59, 60 Lathrop, J. P., 268
Johnson, T h o m a s T., 229, pl. 7 laticosta, Platystrophia. See Brachiopoda
Journal of Geology, 55 Latonia Formation, 4 4 , 51. See also Eden Formation; Kope
journal of the Cincinnati Society of Natural History. See C i n - Formation
cinnati Society of Natural History Economy M e m b e r , Economy beds, 44, 144
junior synonym. See taxonomy: nomenclature: synonym; McMicken Member, 44
taxonomy: nomenclature: synonymy Southgate M e m b e r , 4 4
Jurassic, xix, 4, 239 latus, Isotelus. See Arthropoda: Trilobita
Laurentia. See paleogeography
K-bentonite, K-bentonite beds, K-bentonite "zones," 62, 63, laurentiensis, Sanctum. See taphonomy: trace fossils: borings
163, 287 layer-cake geology, 45
Bear Creek K-bentonite "zone," 62 Lebanon Beds, 47; renamed Richmond, 47
Millbrig K-bentonite bed, 62 L e b a n o n , Warren County, Ohio, 28, 176, 260, 267, 271, 273,
Seneca County, Ohio, 6 2 - 6 3 274, 277
Kankakee Arch, Kankakee Branch of the C i n c i n n a t i Arch, 9. Lebensspuren. See taphonomy, preservation: trace fossils
See also Cincinnati Arch Lepadocystis. See Echinodermata: Rhombifcra
Kaplan, Peter, 124, 125 I.eperditicopida. See Arthropoda: C r u s t a c e a : Ostracoda
Kemper, Willis, 268 Lepidocoleus. See M a c h a e r i d i a
Kenton County, Kentucky, 67, 87, 90, 110, 123, 209 Lepidocyclus. See Brachiopoda: Articulata
Kentucky Lepidolites. See Algae: Chlorophyta: Dasycladaceae
Boone County, 87, 110, 111, 142, 180 Leptaena. See Brachiopoda: Articulata
Bracken County, 178, 209 Leptoholus. See Brachiopoda: Inarticulata

Index 335
Leptotrypa. See Ectoprocta: Trepostomata
l.esquerenx, Leo, 266, 268
Letton, Ralph. See Letton's M u s e u m
I.etton's M u s e u m , 268
Lexington Limestone, 52, 62, 123, 140, 163, 199, 214, 220
Lexington Platform, 216, pl. 12
Lexington, Kentucky, 4 0 , 67, 123, 272
Liberty Eormation, M e m b e r , 44, 4 8 , 53, 56, 63, 68, 72, 77,
7 8 , 79, 105, 107, 108, 111, 134, 162, 176, 180,
211, 214, 224, Pi 11. See also Brookville
Eormation
Lichenocrinus. See Echinodermata: C r i n o i d e a : Disparida
Licrophycus. See taphonomy: trace fossils
life assemblage, 8, 230, 283, 287. See also ecology: ecosys-
tems: c o m m u n i t i e s
life habits. See chapters on individual groups of a n i m a l s
limestone
grainstone, 50, 55, 59, 114, 286, 289
packstone, 50, 55, 59, 114, 286, 289
wackestone, 50
Limper Geological M u s e u m See: M i a m i University
limpets. See Mollusca: Gastropoda: Archaeogastropoda
Limulus. See Arthropoda: Chelicerata: Xiphosurida
L i n d a h l , Josua, 33, 268
Lingula. See Brachiopoda: Lingula
I .innaean I licrarchy. See taxonomj
L i n n a e u s , Karl. See taxonomy
Linne, Karl von. See taxonomy: L i n n a e u s
Liospira. See Mollusca: Gastropoda
lithofacies. See facies
lithology, 46
lithostratigraphic units, 46. See also names of particular units
formation, 46
group, 4 6
member, 4 6
Li Xing, 1, 303
lobsters. See Arthropoda: C r u s t a c e a : Malacostraca
Locke, John, 153, 156, 160, 268, 269, 274
Lockeia. See Mollusca: Pelecypoda; taphonomy: trace fossils
Loeblich, Alfred R Jr., 69
longitudinal thin-sections. See techniques: thin-sections
Lophophora, lophophorates. See Brachiopoda, Ectoprocta
Lophospira. See Mollusca: Gastropoda
Lorenz, D. M., 231
Lorraine Group, 47, 48. See also Maysvillian Stage
Loxoplocus bowdeni. See Mollusca: Gastropoda: Paupospira
luniformis, Diplocraterion. See taphonomy: trace fossils
Lyell, C h a r l e s , x, 5, 17, 170, 260, 268, 269
Lyon, Sidney S., 269
Lyrodesma. See Mollusca: Pelecypoda
m a c h a e r i d i a n s , 142, 192. See also Echinodermata: Stylo-
phora: Enoploura
Lepidocoleus, 145
L. jamesi, 142, 145, 276
M a c k e , W i l l i a m B., 158, 163, 300
Madison County, Kentucky, 77
Madison, Indiana, 77, 79, 221, 2 6 6 , 269
magnificus, Promopalaeaster. See Echinodermata: Asteroidea
magnus, Zygocycloides. See Echinodermata: Cyclocystoidea
Malacostraca. See Arthropoda: Crustacea
336 Index
M a l a y s i a , pl. 3
mammulata, Monticulipora. See Ectoprocta: Trepostomata
Manitoba, C a n a d a , 153
Manitoulinoceras. See Mollusca: Cephalopoda
Maquoketa Group, 81
Marble Hill bed. See Waynesville Eormation
M a r c h a n d , Paul, pl. 13
Marion County, Kentucky, 120
marker beds, 59, 60, 287
M a r t i n , W a y n e D., 49, 50, 5 6 , 9 9 , 114, 115, 218, 256, 298,
307, 315
martini, Neostrabops. See Arthropoda, Aglaspida
Mason County, Kentucky: Maysville, 11, 59, 79, 9 1 , 180
mass extinction. See extinction: mass extinction
Mastigograptus. See Hemichordata: Dendroidea
Mather, W i l l i a m W., 268, 269, 277
Maysville, Mason County, Kentucky, 11, 59, 79, 91, 180
Maysvillian Stage, 11, 2 1 , 4 4 , 4 6 , 51, 5 5 , 6 3 , 6 7 , 6 8 , 7 2 , 79,
80, 81, 89, 109, 123, 125, 130, 139, 144, 145,
151, 153, 155, 158, 160, 163, 182, 184, 191, 203,
207, 208, 209, 219, 220, 223, 233, 287. See also
Lorraine Group
McGraw-Hill C o m p a n i e s , 238
McKinney, Frank K., 94
M c M i c k e n 1 hill, 21. See also University of Cincinnati
M c M i c k e n Member. See Latonia Formation
M c M i l l a n Formation, 44, 48
Bellevue Limestone Member, Bellevue Tongue. See Bel-
levue Limestone
Corryville Member. See Corryville Formation
M o u n t Auburn M e m b e r , 21, 44, 53, 55, 59, 63, 68, 160,
214, 2 2 3 , 2 5 0
M C Z . See Harvard University
M e d i c a l College of Ohio, C i n c i n n a t i , Ohio, 29
medusa. See C n i d a r i a
M e e k , F. B., 24, 3 7 , 4 2 , 4 3 , 4 7 , 9 9 , 106, 109, 126, 127, 137,
149, 150, 156, 160, 170, 172, 174, 175, 176, 178,
182, 184, 186, 191, 229, 266, 269, 270, 277
meeki, Calymene. See Arthropoda: Trilobita: Vlexicalymene
meeki
meeki, Flexicalymene. See Arthropoda: Trilobita
meeki, Onniella. See Brachiopoda
meeki-retrorsa, Calymene. See Arthropoda: Trilobita: Flexica-
lymene retrorsa
megacycle. See cycles, cyclicity
m e g a g u i l d s . See ecology: ecosystems: guilds
Megalograptus. See Arthropoda: Chelicerata: Eurvptcrida
M e h l , M. G., 196, pl. 5
member. See lithostratigraphic units
Merocn'nus. See Echinodermata: Crinoidea: Cladida
Mesozoic Era, xix, 9, 25, 131, 237, 238, 239, 248, 284
Messing, C h a r l e s G., pl. 9
Metacrinus. See Echinodermata: Crinoidea
M e t a z o a , m e t a z o a n , 1, 2, 74, 288, 289
meter-scale cycles. See cycles, cyclicity
Meyer, David L., 113, 249, 299, 308, 311, 318
M i a m i University, Oxford, Ohio, xv, 22, 256, 262, 263, 274
Carl E. Limper Geological M u s e u m ( M U G M ) , xv, 72, 77,
7 8 , 87, 105, 106, 107, 108, 109, 111, 120, 123,
126, 127, 134, 137, 156, 157, 174, 175, 176, 184,
195, 229, 256
 Miamitown Shale, 1 2 , 2 1 , 53, 60, 124, 125, 129 Endoceras, 134
Michigan, University of. See University of M i c h i g a n Corbyoceras, 138
Mickleborough, John, 26, 27, 153, 155, 262, 276, 313 G. curvatum, 137, 138
micraulaxum, Multiplicisphaeridium, S e e : acritarchs G. duncanae, 134, 135
microscopy. See techniques: microscopy lsorthoceras, 137
i n f r a s t r u c t u r e , 6, 6 8 , 7 2 , 144, 166, 168, 192, 292 Manitoulinoceras
Middle (Eden) Shales, 47 M. tenuiseptum, 137
Milankov itch Cycles. See cycles, cyclicity M. williamsae, 137
Millbrig K-bentonite bed. See K-bentonite, K-bentonite beds, Narthecoceras: N. dunni, 138
K-bentonite "zones" nautiloid cephalopods, 60, 87, 88, 8 9 , 9 0 , 94, 132-139,
millepunctata, Trematis. See Brachiopoda: Inarticulata 2 0 5 , 2 2 8 , 229, 235, 236, 238, 241, 244, 245, 249,
Miller, Arnold I., 113, 231, 232, 305, 308, 312 2 5 1 , 2 5 2 , 264, pl. 8 (includes
Miller, C. A., 269 Actinoceratoidea, Endoceratoidea, and
Miller, S. A., 14, 17, 23, 2 4 - 2 5 , 28, 2 9 , 4 2 , 6 8 , 80, 82, 126, Nautiloidea)
127, 134, 137, 142, 144, 157, 160, 161, 176, 182, Nautiloidea, 245
184, 185, 203, 204, 205, 2 0 6 , 208, 209, 210, 259, Ascocerida, 137
260, 261, 265, 266, 267, 270, 272, 274, 275 Nautilus, 95, 117, 131, 132, 134, 137, 139, 241. pl. 4
milleranus, Ceraurus. See Arthropoda: Trilobita: Phacopida Orthocerida, 137
milleri, Eichenocrinus. See Echinodermata: C r i n o i d e a : octopus, 117, 131, 139
Disparida Oncoceras, 137
milleri, Technophorus. See
Mollusca: Rostroconchia O. delicatum, 137
millipedes. See Arthropoda: Diplopoda Orthonyhyoceras, 43. See also Treptoceras
Milne-Edwards, Henri, 265, 270 Schuchertoceras
miniata, Cucumaria. See Echinodermata: Holothuroidea O.obscurum, 137, 138
Minnesota Geological Survey, 31 squid, 117, 118, 131, 137, 139,252
M i n t z , Leigh W., 178, 310 trace fossils, 138. See also taphonomy: trace fossils
Miocene, xix, 4 Treptoceras, 43, 136, 137, 223,228, 229. See also
Misener, John, 270 Orthonyhyoceras
Misener, S. R., 270 T. duseri, 60, 134, 135, 136, 137, 138,228, 229
miseneri, Conularia. See C n i d a r i a : Scyphozoa: C o n u l a r i i d a Zittelloceras: Z. russelli, 137; Z. williamsae, 137
mode of life, 5,7, 52, 74, 160, 169, 188. See also chapters on chitons. See Polyplacophora
individual groups of a n i m a l s C r i c o c o n a r i d a , 145. See Tentaculitoidea
modesta, Isygospira. See Brachiopoda: Articulata Gastropoda, 7, 24, 32, 101, 116, 117, 118, 119-125, 127,
modiolaris, Modiolopsis. See Mollusca: Pelecypoda 129, 137, 139, 182, 223, 225, 231, 235, 237, 238,
Modiolopsis. See Mollusca: Pelecypoda 240, 241, 243, 244, 246, 247, 248, 282
Mohawkian Stage, 167 Archaeogastropoda, 123 (See also Cyclonema;
Mohr.Paul, 261,270 Naticonema)
molds, composite. See taphonomy: molds abalones, 119, 123
molecular clocks, 62. See also organic evolution limpets, 119, 123, 139
Mollusca, 26, 3 1 , 8 7 , 9 3 , 9 9 , 101, 116-140, 144, 145, 162, 204, periwinkles, 123
206, 219, 223, 224, 225, 231, 233, 241, 244, 245, bellerophontids, 120
259, 260, 2 6 1 , 2 8 2 , 2 8 8 , 2 9 0 , 2 9 1 Cyclonema, 120, 123, 144, 176, 223, 252
anatomy and morphology: tentacles, 119, 131, 132, 138, 252 C. bilixlata, 122, 123
Aplacophora, 119,280 C. gracile striatulum, 122, 123
Calyptomatida, 145. See hyolithids C. humerosum, 120, 122, 123
Cephalopoda, 29, 43, 8 7 , 8 8 , 9 3 , 9 4 , 9 5 , 9 6 , 101, 118, 119, C. sublaeve, 122, 123
131-138, 139, 144, 154, 196, 205, 220, 223, 225, C. varicosum, 122, 123
231,235, 238, 241, 247, 248, 249, 251, 252, 260, Cyclora, 223
264, 270, 280, 286 Liospira, 225
Actinoceratoidea, actinocerids, actinoceroids, 245 Lophospira, 223, 225
Beloitoceras: B. amoenum, 136, 137 Loxoplocus: L. bowdeni. See Paupospira
Charactoceras, 138 Naticonema, 120
C. haeri, 136, 137 Neogastropoda, 123
cuttlefish, 131 cones, 123
Cyrtoceras: C. vallandighami, 275 murexes, m u r i c e s , 123
Diestoceras, 138 Subulites (Fusispira), 122, 123
D. eos, 136, 137 whelks, 123
Endoceratoidca, endocerids, endoceroids, 134, 137, Paupospira
154,245 P. bowdeni, 125, 127
Cameroceras, 137, 225 P. moorei, 122, 123, 127
C. inaequabile, 134, 135, 137 P. tropidophora, 127

Index 337
 platyceratids, 120 monobathrid camcratcs. Sec Echinodennata: Crinoidea:
Ptcropoda, 145 Camerata
Salpingostoma: S. richmondensh, 120, 12 J Monoplacophora. See Mollusca
slugs, sea slugs, 117, 119 Montgomery County, Ohio, 72, 157
Monoplacophora, 93, 119, 120, 125, 139, 223, 244, 247 monticule. See Ectoprocta: anatomy and morphology
Archinacella: A. area, 120, 121 Monticulipora. See Ectoprocta: Trepostomata
Cyrtolites, 125, 139 Moore, Richard B., 17, 270, 279
C. ornatus, 120, 121 Moore, T i m , 38
Helcionopsis: H. striata, 120, 121 moorei, Lepadocystis. See Echinodermata: Rhombifera
Neopilina, 139 moorei, Paupospira. See Mollusca: Castropoda
Sinuites, 225 Morris, Robert W., 123
S. cancellatus, 120, 121 Morrow, Warren C o u n t y . Ohio, 175
Pelecypoda, 7, 92, 93, 98, 99, 100, 101, 112, 113,117, 118, Mount Auburn M e m b e r , Formation, 21, 44, 53, 55, 59, 63,
119, 123,125, 128-131, 139, 144, 157, 168, 189, 68, 160,214, 223, 250. See also M c M i l l a n
206, 210, 223, 225, 231, 235, 237, 238, 240, 241, formation
242, 244, 245, 251, 269, 276, 280, 281, 283, 284, Mount I lope Member. See Fairview Formation
291,293 mountain building, orogeny. See tectonic activity
Ambonychia, 88, 89, 126, 129, 130, 131, 223, 225, 240, mudstone, 50, 217, 218, 219, 222, 224, 225
241, 252 M l ' C M . Sec Miami I University, ()hio, ( h l n r d
A. cultrata, 126 multipartitum, Petalichnus. See taphonomy: trace fossils

Anomalodonta: A. gigantea, 126 Multiplicisphaeridium. See acritarchs

Caritodens, 128, 130, 223, 241, 251 murexes, murices. See Mollusca: Castropoda:
C. demissa, 126 Neogastropoda
cockles, 129 mussels. See Mollusca: Pelecypoda
Corallidomus: C. scobina, 128, 206 m u t u a l i s m . See ecologic associations
Ctenodonta, 129, 131 mvtilids. See Mollusca: Pelecypoda
Cycloconcha, 129 myzostomids. See Annelida
Cymatonota, 129
C. typicalis, 40 nannoplankton, 239
Cyrtodontula, 129 Narthecoceras. See Mollusca: Cephalopoda
Deceptrix, 129, 225 Naticonema. See Mollusca: Gastropoda
giant c l a m . See Tridacna National Academy of Sciences, 30, 32, 274
Ischyrodonta: 1. truncata, 126, 131 National M u s e u m of Natural History ( U S N M ) , xv, 19, 21,
l.ockeia: 1,. siliquaria, 209, 210 75, 120, 127, 128, 153, 173, 175, 176, 182, 184,
l.yrodesma, 129 185, 266. See also Smithsonian Institution;
Modiolopsis, 129, 130, 131, 224, 225, 251 United States National M u s e u m
M. modiolaris, 127 Natural History M u s e u m , London, xv, 23, 26
mussels, 22, 32, 33, 117, 129, 131, 261 natural selection. See organic evolution
mvtilids, 131 nautiloid ccphalopod. See Mollusca: C e p h a l o p o d a
Opisthoptera, 129 Nautiloidea. See Mollusca: C e p h a l o p o d a
O. casei, 126 Nautilus. See Mollusca: Cephalopoda: Nautiloidea
oysters, 114, 117, 129, 131 nealli, Caurocrinus. See Echinodermata: Crinoidea:
Pseudocolpomya, 131 Camerata
Pterinea, 129 neglecta, Plectorthis. See
Brachiopoda: Articulata
P. demissa. See Caritodens demissa nekton, nektonic, 7, 138, 198, 280, 288, 290
Rhytimya, 225 Nelson County, Kentucky, 78
scallops, 112, 113, 117, 129 Neocrinus. See Echinodermata: Crinoidea
spiny oysters. See Spondylus Neogastropoda. See Mollusca: Castropoda
Spondylus, 131 Neopilina. See Mollusca: Monoplacophora
Tridacna, 131 Neostrabops. See Arthropoda: Aglaspida
Vanuxemia: V. waynesvillensis, 305 Nereigenys. See Annelida: scolecodonts
Polyplacophora, 117, 118, 119, 139 Nestor, Robert, 153
chitons, 117, 139, 145, 248 New C a l e d o n i a , pl. 4
Rostroconchia, 119, 127, 139, 247, 290 New England, 3, 215
Technophorus, 139 New York, 3, 17, 23, 27, 35,47, 4 8 , 149, 155, 160, 257, 265,
T. faberi, 127 2 6 6 , 2 7 1 , 2 7 6 , 293
T. milleri, 127 New York Geological Survey, 23, 32
Scaphopoda, 119, 139, 140 New York State M u s e u m , 180
Rhytiodentalium: R. kentuckyensis, 140 Newberry, John Strong, 24, 270, 271, 276, 277
tusk shells. See Scaphopoda Newton, A . J . , 270
molting. See ecdysis Nicholson, H. Alleyn, 75, 77, 270-271

338 Index
nicholsoni, Ceramopora. See Ectoprocta: Cystoporata Ohio State University, C o l u m b u s , Ohio, xv, 2, 51, 256, 271
Nickles, John M., 1 8 , 1 9 , 2 0 , 27, 30, 32-33, 3 4 , 4 7 , 52, 230, Orton G e o l o g i c a l M u s e u m ( O S U ) , xv, 67, 128, 134, 229
259, 260, 261, 263, 265, 266, 267, 269, 270, 272, ohioensis, Megalograptus. See Arthropoda: Chelicerata:
275, 277 Eurypterida
Nitecki, Matthew H 68 Oldroyd, John David, 231
nodosa, Fromia. See Echinodermata: Asteroidea Oligocene, xix, 4
nomen duhium (pl., nomina dubia), 42, 288. See taxonomy: Oncoceras. See Mollusca: Cephalopoda
nomenclature oncolites. See Algae
nomen nudum (pl., nomina nuda), 42, 288. See taxonomy: onealli, Acidaspis. See Arthropoda: Trilobita:
nomenclature Odontopleurida
nomenclature. See taxonomy onealli, Parvohallopora. See Ectoprocta: Trepostomata
normal wave-base, fair-weather wave-base. See wave base Ophidiasteridae. See Echinodermata: Asteroidea
North Atlantic Province. See faunal provinces Ophiothrix. See Echinodermata: Ophiuroidea
North Bend Tongue. See Fairview Formation Opisthoptera. See Mollusca: Pelecypoda
numerosum, Trachomatichnus. See taphonomy: trace fossils Orbigny, Alcide d', 85, 90, 95, 271
nutrition. See food orbital parameters. See cycles: Milankovitch C y c l e s
Nyctopora. See C n i d a r i a : Anthozoa: Tabulata order. See taxonomy: L i n n a e a n Hierarchy
Ordovician: "Ordovician Biodiversification Event," 2, 289;
O ' N e a l l J . K., 271 "Ordovician Radiation," 1, 2, 289
Obata, Tadahiro, 43,317 ordovicicus, Amorphognathus. See conodonts
obligate association. See ecologic associations Ordovicium. See acritarchs
obrution deposits, 60, 288 Oregonia Formation, M e m b e r , 44, 53, 56, 63,214, 223. See
obscurum, Schuchertoceras. See Mollusca: C e p h a l o p o d a also Arnheim Formation
occidentalis, Hebertella. See Brachiopoda: Articulata oregonia, Oulodus. See conodonts
octopus. See Mollusca: Cephalopoda organic evolution, 1, 2, 39, 6 1 , 62, 67, 82, 105, 115, 118, 131,
Odontopleurida, odontopleurids. See Arthropoda: Trilobita 134, 139, 163, 167, 195, 196, 229, 233, 237, 267,
Oeh, George, 271 282, 286, 287. See also durophagy; molecular
oehanus, locrinus. See Echinodermata: Crinoidea: Disparida clocks
Ohio convergent evolution, 105, 282, 286
Adams County, 75, 158, 162, 163, 216, 229, pl. 7 evolutionary relationships. See chapters on individual
Brown County, 123, 128, 151 groups of a n i m a l s
Butler County; 68, 75, 80, 87, 89, 107, 108, 126, 127,134, homeomorphy, 105, 282, 286
137, 153, 156, 160, 176, 195, 2 2 9 organic evolution by natural selection, 134
Clermont County, 111, 126, 128, 153, 158, 163, 170, 186, ornatus, Cyrtolites. See Mollusca: Monoplacophora
203, 207, 208, 209, 261 ornatus, Protoscolex. See Annelida
Bear Creek Quarry, 62 orogeny (mountain building). See tectonic activity
Stonelick Creek, 11, 254 Orthocerida. See Mollusca: C e p h a l o p o d a : Nautiloidea
Clinton County, 6 0 , 7 2 , 75, 128, 136, 142 Orthograptus. See Hemichordata: Graptoloidea
Dent, 176,180 Orthonybyoceras. See Mollusca: C e p h a l o p o d a :
Hamilton County, 10, 8 9 , 9 0 , 106, 142, 151, 155, 156, 173, Actinoceratoidea
1 8 0 , 1 8 6 , 277, PL 10 Orthorhynchula. See Brachiopoda: Articulata
Highland County, 151, 153 Orton Geological M u s e u m . See Ohio State Universitv
Huffman Dam (near Dayton), 153 Orton, Edward, 2 4 , 4 7 , 52, 268, 271, 276
Lebanon, 28, 176, 260, 267, 271, 273, 274, 277 Orton, Edward, Jr., 271
Montgomery County, 72, 157 Osgood, Richard G., Jr., 138, 203, 204, 205, 206, 208, 209,
Morrow, 175 210,211,212
Oxford, 105, 153, 155, 156, 256, 269, pl. 11 Osteichthyes ("bony fish"). See Chordata: Vertebrata
Point Pleasant, 225 Ostracoda. See Arthropoda: Crustacea
Portsmouth, 10 O S U . See Ohio State University: Orton Geological M u s e u m
Preble County, 77, 105, 107, 108, 109, 111, 123, 126, 137, Oulodus. See conodonts
178, 184 Owen, David Dale, 261, 269, 272
Trammel Fossil Park, Sharonville, 12, 124, 257 oweni, Fistulipora. See Ectoprocta: Cystoporata
Seneca County, 63 O w e n s , R. M., 151, 155, 161, 305
Warren County, 60, 105, 126, 136, 172 Oxford, Butler County, Ohio, 105, 153, 155, 156, 256, 269,
Waynesville, 120, 182, 184, 185, 256, 259, 266 pl. 11
W i l m i n g t o n , 260, 273, 276 oxygen content. See environment
Ohio Division of Geological Survey, 92, 256. See also Ohio oysters. See Mollusca: Pelecypoda
Geological Survey ()zarkian System. 30, 31
Ohio Geological Survey, 23, 269, 276. See also Ohio Divi-
sion of Geological Survey pacifica, Heteropora. See Ectoprocta
Ohio M e c h a n i c s Institute, C i n c i n n a t i , Ohio, 31, 262 packstone, 50, 55, 59, 114, 2 8 6 , 289

Index 339
Palaeasterina. See Echinodermata: Asteroidea
Palacocopida. See Arthropoda: C r u s t a c e a : Ostracoda
Palaeophycus. See taphonomy: trace fossils
Palaeoscia. See C n i d a r i a : Hydrozoa: Siphonophorida: Por-
pitidae; laphonomv: trace lossils
Palau, I'alau islands, pl. 3, pl. 9 '
p a l e o b a t h y m e t r y , 6 1 , 113, 289
Palcoccne, Palaeocene, xix, 4
Paleodictyon. See taphonomy: trace fossils
paleoecology. See ecology
paleoenvironmental interpretation. See environment
paleogcographv
Baltica, 6 2 , 2 1 5 , 2 1 6
G o n d w a n a , 215, 216
Iapetus O c e a n , S e a , 3, 62, 216. 286, 292
Laurentia, 62, 215, 216, 238, /'/. J
paleogeographic maps, 30, 32, pl. 1, pl. 2, pl. 12
Paleotcthys O c e a n . 216
Panthalassic O c e a n , 216
Queenston Delta, 216, 220, PL 12
Rodinia, 216
Paleontological Research Institution, 75, 87, 158, 159, 173,
186, 203, 209, 257
Paleontological Society, 2, 30, 32, 36, 172, 252, 257, 274
Schuchcrt M e d a l , 32
The Paleontologist, 17, 22
paleoslope, 55, 289
paleosynecology. See ecology: synecology
P a l e o l e t l n s ( ) c c a n . See p a l c n g c o g r a p l n
Paleozoic Era, xix, 1, 2, 8, 9, 31, 34, 35, 45, 4 6 , 67, 72, 121,
123, 131, 138, 139, 182, 183, 199, 237, 238, 239,
241, 250, 277, 279, 284, 285, 286, 289
"Paleozoic Fauna," 2, 237
Palmer, T i m o t h y ) . , 124, 2 0 6 , 209, 210, 281, 298, 322
P a n a m a , /'/. 9
Panthalassic O c e a n . See paleogeography
parasite, parasitism. See ecologic associations
paratype. See taxonomy: tvpe specimens
Parsley, Ronald L., 186
parviusculus, Decoroproetus. See Arthropoda: trilobita
parviusculus, Proetus. See Arthropoda: Trilobita:
Decoroproetus
Parvohallopora. See Ectoprocta: Trepostomata
Pasceolus. See Algae: Chlorophvta: Dasvcladaeeae:
Cyclocrinites
pascichnia. See taphonomy: trace fossils: behavior categories
patch reef. See bioherin
pathology, 93, 182
deformation (in life), deformities, 103, 182
Patterson, W i l l i a m ) . , 272
pattersoni, CAyptocrinus. See Echinodermata: Crinoidea:
Camcrata
Pattersonia. See Porifera
Patzkowsky, Mark E., 63, 232, 233, 241, 308
Paul, Christopher R. C. 190. 191
Paupospira. See Mollusca: Gastropoda
Peaslee, John B., 27
pedicle. See Brachiopoda: anatomy
Pelecypoda. See Mollusca
Pennsylvania A c a d e m y of S c i e n c e , 95
Penrose M e d a l . See Geological Society of America
340 Index
pentagonus, Cincinnaticrinus. See Echinodermata: Cri-
noidea: Disparida
pera, Petroxestes. See taphonomy: trace fossils: borings
Pericharax. See Porifera
periwinkles. See Mollusca: Gastropoda: Archaeogastropoda
Permian, xix, 4, 74, 77, 121, 145, 182, 186
Petalichnus. See taphonomy: trace fossils
Peter, M a r k , 151, 157
Petraster. See Echinodermata: Asteroidea
Petrocrania. See Brachiopoda: luarticulata
Perroxesfes. See taphonomv: trace fossils: borings
Phacopida, phacopids. See Arthropoda: Trilobita
Phaeophyta. See Algae
Phanerozoic Eon, 2 , 4 , 4 6 , 62, 239, 284, 289
Philhedra. See Brachiopoda: luarticulata
Pholadomorpha. See Mollusca: Pelecypoda
Phosphannulus, 182. See also byroniids
phosphatization. See taphonomv: phosphatization
Phragmodus. See conodonts
phytogeny, phylogenetic, 198. See also organic evolution
phvlum, phvla. See taxouomv: l.innacan Hierarchy
pilea, Carneyella. See Echinodermata: Edrioasteroidea
pillbug. See Arthropoda: Crustacea: Isopoda
Plaesiomys. See Brachiopoda: Articulata
planispiral coiling. See coiling of shell
plankton, planktonic, 7, 52, 6 8 , 69, 74, 145, 172, 195, 234,
239, 280, 282, 288, 290
microplankton, 52, 239
nannoplankton, 239
phytoplankton, 6 8 , 6 9 , 239, 283
zooplankton, 74
Plantae, plants: land plants, 239, 268, 270 '
platyceratids. See Mollusca: Gastropoda
Platycoryphe. See Arthropoda: Trilobita
Platystrophia. See Brachiopoda: Articulata
Plectodina. See conodonts
Plectorthis. See Brachiopoda: Articulata
Pleistocene, ix, xix, 4, 10
Plicodendrocrinus. See Echinodermata: Crinoidea: Cladida
Pliocene, xix, 4
Plummer, John T , 272
Podocopida. See Arthropoda: Crustacea: Ostracoda
Point Pleasant Formation, 19, 62, 209, 214, 225. See also
River Quarry Beds
Pojeta, John, Jr., 127, 128, 206, 308
polar ordination. See techniques
Polychaeta, polychaete worms. See Annelida
polvdactvlus, Cupulocrinus. See Echinodermata: Crinoidea:
Cladida
polymorphism, 89, 283, 290. See also dimorphism
polyp. See C n i d a r i a
polypide. See Ectoprocta: anatomy and morphology: zooid
PoKplacophora. See Mollusca
Polyzoa. See Ectoprocta
ponderosa auhurnensis, Platystrophia. See Brachiopoda
ponderosa, Platystrophia. See Brachiopoda
Pontiometra. See Echinodermata: Crinoidea
Pope, John K . , 4 8 , 145
popei, Enoploura. See Echinodermata: Stylophora
populations. See ecology: ecosystems
Porifera, 24, 31, 38, 6 8 , 7 1 - 7 3 , 79, 123, 241, 242, 247, 248, 292
 Brachiospongia: B. tuberculata, 72, 73
Labechia: L. huronensis, 72, 73
Pattersonia: P. tuberosa, 72, 73
Pericharax, pl. 3
sclerosponges, coralline sponges, 7 1 , 72
Acanthochaetetes: A. wellsi, pl. 3
Stromatoporoidea, 6 8 , 72, 128, 2 0 6 , 220, 221, 233, 242,
243, 245, 247, 292
Aulacera, 72
A. undulata, 72, 73
Portsmouth, Ohio, 10
Portuguese Man-of-War. See C n i d a r i a : Hydrozoa:
Siphonophorida
post-mortem transportation, 114, 137
Potter, Paul Edwin, 8, 11, 59, 106
Preachersville Member. See Drakes Formation
Preble County, Ohio, 77, 105, 107, 108, 109, 111, 123, 126,
137, 178, 184
Precambrian, xix, 67, 7 1 , 151
predation. See ecologic associations
predator-prey interactions. See ecologic associations
preservation. See taphonomy
primary producers. See ecology: ecosystems
primary productivity, primary production, 2 3 8 - 2 3 9 , 282. See
also ecology: ecosystems
Primaspis. See Arthropoda: Trilobita: Odontopleurida
Prioniodus: P. dychei. See conodonts
priority. See taxonomy: nomenclature
Probasco, Henry, 272
Proetus parviusculus. See Arthropoda: Trilobita:
etus parviusculus
Promopalaeaster. -See Echinodermata: Asteroidea
pronunciation of scientific n a m e s . See taxonomy:
nomenclature
Protaraea. See C n i d a r i a : Anthozoa: Tabulata
Protasterina. See Echinodermata: Ophiuroidea
Protoscolex. See Annelida
pseudochitin, 69
Pseudocolpomya. See Mollusca: Pelecypoda
Pseudolingula. See Brachiopoda: Inarticulata
Pterinea demissa. See Mollusca: Pelecypoda: Caritodens
demissa
Pterobrancbia. See Hemichordata
Pteropoda. See Mollusca: Gastropoda
pudicum, Rusophycus. See taphonomy: trace fossils
Pycnocrinus. See Echinodermata: Crinoidea: C a m e r a t a
pyritization. See taphonomy
Pyrrophyta. See Algae
Quadrijugator. See Arthropoda: Crustacea: Ostracoda
quadrimucronatus, Orthograptus. See Hemichordata:
Graptoloidea
Queenston Delta. See paleogeography
radiometric dating. See time: absolute age
Rafinesque, C. S., 40, 272
Rafinesquina. See Brachiopoda: Articulata
ramosa, Parvohallopora. See Ectoprocta: Trepostomata
Reba Formation, 214
recrystallization, 6, 8, 94, 290. See also taphonomy:
recrystallization
reef. See bioherm
regeneration, 173, 182, 183, 186, 198, 236
regularis, Quadrijugator. See Arthropoda: C r u s t a c e a :
Ostracoda
Reinhardt, E., 272
relative age, relative dating, 46
repichnia. See taphonomy: trace fossils: behavior categories
reticularis, Anomaloides. See Algae: Chlorophyta:
Dasycladaceae
retrorsa, Calymene. See Arthropoda: Trilobita: Flexicalymene
retrorsa
Retrorsirostra. See Brachiopoda: Articulata
rex, lsotelus. See Arthropoda: Trilobita
Rbahdopleura. See Hemichordata: Pterobrancbia
Rhodes, F. H. T, 196
Rhodesognathus. See conodonts
Rhodophyta. See Algae
Rhombifera, rhombiferan "cystoids." See Echinodermata
Rhynchotrema. SeeBrachiopoda: Articulata
Rhytimya. See Mollusca: Pelecypoda
Rhytiodentalium. See Mollusca: Scaphopoda
Richards, R. Peter, 102
Richardson, J . M . , 273
richardsoni, Clyptocrinus. See Echinodermata: C r i n o i d e a :
Camerata
Richmond Group, R i c h m o n d i a n Stage, 4 0 , 4 4 , 4 6 , 47, 4 8 ,
51, 6 2 , 6 3 , 6 8 , 72, 73, 75, 77, 78, 79, 81, 89, 105,
120, 124, 125, 130, 137, 138, 139, 145, 151, 153,
156, 157, 158, 160, 162, 164, 178, 180, 182, 184,
Decoropro- 185, 186, 189, 191, 196, 2 0 6 , 208, 220, 232, 233,
260, 291, pl. 7
richmondense, Stromatocerium. See Algae: Rhodophyta:
Solenopora richmondensis
richmondensis, Peptaena. See Brachiopoda: Articulata
richmondensis, Protaraea. See Cnidaria: Anthozoa: 'Tabulata
richmondensis. Salpingostoma. See Mollusca: Gastropoda
richmondensis, Solenopora. See Algae: Rhodophyta
richmondensis, lentaculites. See Tentaculitoidea
R i c h m o n d i a n Invasion, 60, 81, 138, 232-233, 240, 241
Richter, Rudolf, 204
Riddell, John L., 273
rippled beds, 60
River Quarry Beds, 47, 225. See also Point Pleasant
Formation
Robison, Richard A., 144
rock-stratigraphic units. See lithostratigraphic units
Rodinia. See paleogeography
Roeininger, C a r l , 273
Ropalonaria. See Ectoprocta: Ctenostomata
Ross, Reuben J . , Jr., 150, 151, 160
Rostroconchia. See Mollusca
Rowell, Albert J . , 185, 298
Rowland M e m b e r . See Drakes Formation
rugosa, Parvohallopora. See Ectoprocta: Trepostomata
Rugosa, rugosans. See C n i d a r i a : Anthozoa
rugosa, Sowerbyella. See Brachiopoda: Articulata
rugosa, Thaerodonta. See Brachiopoda: Articulata
Runncgar, Bruce, 127
Rush, Jerry, 89, 155, 156, 157
Rusophycus. See Arthropoda: Trilobita: trace fossils; taphon-
omy: trace fossils
Index 341
russelli, 7,itteIloceras. See Mollusca: Cephalopoda
Sacco, W i l l i a m K., pl. 3
salinity. See environment
Salpingostoma. See Mollusca: Gastropoda
Salteraster. See Echinodermata: Asteroidea
Saluda Formation, M e m b e r , 44, 48, 49, 53, 79, 81, 138, 162,
214, 221. See also Whitewater Formation
Sanctum. See taphonomy: trace fossils: borings
sand dollars. See F c h i n o d e r m a t a : Echinoidea
Sanner, JoAnn, 19, 21
Sansom, Ivan ) . , 199
scabiosa, Petrocrania. See Brachiopoda: Inarticulata
Scaphopoda. See Mollusca
scavenging. See food
Schafer, W i l h e l m , 2 0 4
Schizomania. See Brachiopoda: Inarticulata
S c h l e m m e r , C h a r l e s , 273, 275
S c h m i d t , David A., 186, 303
Schuchert, C h a r l e s , 20, 28, 30, 31-32, 33, 35, 184, 264, 273,
274, 275
schuchertana, Catazyga. See Brachiopoda: Articulata
Schuchertoceras. See Mollusca: C e p h a l o p o d a
S c h u m a c h e r , Gregory A., 44, 178, 312, 318
Schweizcrbarfschc Science Publishers, 67
scientific n a m e . See taxonomy: nomenclature: species n a m e
Scleractinia. See C n i d a r i a : Anthozoa
sclerosponges. See Porifera
scobina, Corallidomus. See Mollusca: Pelecypoda
scolecodonts. See Annelida
scorpions. See Arthropoda: Chelicerata: Arachnida
Scotese, C h r i s , pl. 1
Scott, W i l l i a m B e r r y m a n , 273
Scoville, S. S., 273
scovillei, Orthis. See Brachiopoda: Articulata: Austinella
scovillei
Scyphozoa, scyphozoan. See C n i d a r i a
sea a n e m o n e s . See C n i d a r i a : Anthozoa
sea c u c u m b e r s . See Echinodermata: Holothuroidea
sea fans. See C n i d a r i a
sea level, sea-level c h a n g e , sea-level curves, 3, 4, 9, 53, 55, 56,
59, 6 1 , 80, 214, 216, 217, 219, 221, 240, 285, 291
sea lilies. See Echinodermata: Crinoidea
"sea scorpion." See Arthropoda: Chelicerata: Eurypterida
sea slugs. See Mollusca: Gastropoda
s c i s i , i i Sec Echinodermata: Asteroidea
sea urchins. See Echinodermata: Echinoidea
sea whips. See C n i d a r i a
sea-floor spreading, 3, 216, 291
Seilacher, Adolf, 114, 204, 205, 210, 211
seismites, 6 0 - 6 1 , 291
senaria, Calymene. See Arthropoda: Trilobita: Flexicalymene
senaria
S e n e c a County, Ohio, 63
Sepkoski, John J . , Jr., 2
S E P M . T h e Society for S e d i m e n t a r y Geology (originally the
Society of Economic Geologists and M i n e r a l o -
gists), 57, 158
sequence stratigraphy, 53, 54-57, 59. See also C I , C 2 , C 3 ,
C4, C5, C6 sequence
sequence boundaries, 55, 6 1 , 291
342 Index
serpulid worms. See Annelida: Polychaeta
sexual dimorphism. See dimorphism, sexual
Seychelles, Indian O c e a n , pl. 9
shalc-to-lhnestone ratio, 232. See also elastic ratio
Shaler, Nathaniel Southgate, 23, 2 7 3 - 2 7 4
shedding. See ecdysis
shell coiling. See coiling of shell
shell pavements, shingled beds, 59, 60, 99, 110, 114, 251. See
also Brachiopoda: Articulata: Rafinesquina
Shideler, W i l l i a m Henry, 28, 262, 263, 265, 268, 274
shideleri, Megalograptus. See Arthropoda: Chelicerata:
Fury pterida
S h i m i z u , Saburo, 43
shingled beds. See shell pavements
Shirley, J . , 150
Shrake, Douglas L., 9 2 , 9 3 , 317
shrimps. See Arthropoda: Crustacea: Malacostraca
S i g m a G a m m a Fpsilon, 91
siliciclastics, 3, 220, 291
silicification. See taphonomy
siliquaria, Lockeia. See Mollusca: Pelecypoda
Silurian, xix, 2 , 4 , 9 , 35, 5 6 , 7 2 , 7 9 , 1 6 1 , 1 9 5 , 217, 237, 2 4 0 , 2 8 9
simplex, Ectenocrinus. See Echinodermata: Crinoidea:
Disparida
simplex, Hudsonaster. See Echinodermata: Asteroidea
simplex, Palaeaster. See Echinodermata: Asteroidea: Hudson-
aster simplex
Singh, R a m a n J . , 9 5
Sinuites. See Mollusca: Monoplacophora
slugs. See Mollusca: Gastropoda
Smith, Andrew B., 190, 191
S m i t h , M. Paul, 199, 316
S m i t h , M o y a M., 199, 316
S m i t h , W i l l i a m , 6 1 , 281
Smithsonian Institution, xv, 19, 21, 30, 153, 269
snails. See M o l l u s c a : (Gastropoda
soft-tissue preservation, 79, 142, 143, 168, 190. See also
taphonomy
Solenopora. See Algae: Rhodophyta
Southgate Member. See Latonia Formation
Sowerbyella. See Brachiopoda: Articulata
Spatiopora. See Ectoprocta: Trepostomata
species. See taxonomy: L i n n a e a n Hierarchy
species diversity. See diversity
species n a m e . See taxonomy: nomenclature
specific n a m e . See taxonomy: nomenclature
speciosa, Palaeasterina. See Echinodermata: Asteroidea
speciosus, Petraster. See Echinodermata: Asteroidea
speciosus, Promopalaeaster. See Echinodermata: Asteroidea
Sphenothallus, 93, 243, 244
spiders. See Arthropoda: Chelicerata: Arachnida
spinosity, 69, 89, 123, 131, 145,148, 153, 155, 160, 162, 163,
167
spinosus, Taeniaster. See Echinodermata: Ophiuroidea
spinosus, Triarthrus. See Arthropoda: Trilobita: Olenida
spiny oyster. S e c M o l l u s c a : P e l e c y p o d a : Spondvlus
spirorbid worms. See Annelida: Polychaeta
Spondylus. See Mollusca: Pelecypoda
sponges. See Porifera
Spreiten. See taphonomy: trace fossils
Spring Grove Cemetery, C i n c i n n a t i , Ohio, 22, 23, 24, 25
Sprinkle, James, 1 6 8 , 1 8 5 , 1 9 1 , 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 Cnidaria: Anthozoa: Rugosa
stellatus, Cystaster. See Echinodermata: Edrioasteroidea
stelliforme, Asteriacites. See taphonomy: trace fossils
sterlingensis, Tentaculites. See Tentaculitoidea
Stevens, W. J . , 274
Stingy Creek Formation, 214
Stokes, W i l l i a m Fee, 1
Stonelick Creek, Clermont County, Ohio, II, 254
storm wave-base. See wave base
storms, 7, 56,57, 5 9 , 6 0 , 6 3 , 6 4 , 77, 8 1 , 9 2 , 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
stratigraphic code. See stratigraphy: C o d e of Stratigraphic
Nomenclature
stratigraphy. See also sequence stratigraphy
C o d e of Stratigraphic Nomenclature, 49, 51
International Stratigraphic C o d e , 49
Streptaster. See Echinodermata: Edrioasteroidea
Streptelasma. See C n i d a r i a : Anthozoa: Rugosa
striata, Helcionopsis. See Mollusca: Monoplacophora
striatulum, Cyclonema gracile. See Mollusca: Gastropoda:
Cyclonema gracile
Strimple, Harrell L., 172, 173, 320
Stromatocerium richmondense. See Algae: Rhodophyta: Sole-
nopora riclunondensis
Stromatoporoidea. See Porifera
Strongylocentrotus. See Echinodermata: Echinoidea
Strophomena. See Brachiopoda: Articulata
Stunner, W i l h e l m , 153
Stylophora. See Echinodermata: carpoids
suhcrassus, locrinus. See Echinodermata: Crinoidea:
Disparida
suherectus, Drepanoistodus. See conodonts
suhlaeve, Cyclonema. See Mollusca: Gastropoda
subquadrata, Plaesiomys. See Brachiopoda: Articulata
subspecies. See taxonomy: L i n n a e a n Hierarchy
Subulites [Fusispira). See Mollusca: Gastropoda:
Neogastropoda
succession. See ecologic succession
Sumrall, Colin D., 178, 185, 250
Sunset Formation, Member, 44, 53, 56, 63, 164,214, 224. See
also Arnheim Formation
superposition. See time: relative age, relative dating
suspension feeding. See food
S w a d l e v , W C , 129
Sweet, Walter C, 51, 199, 321
symbiosis, symbiont. See ecologic associations
symmetry, 77, 98, 99, 100, 101, 130, 145, 167, 168, 169, 189,
190, 191, 196, 198, 280, 288, 289, 292
synecology. See ecology
synonym, synonymy. See taxonomy: nomenclature
syntypes. See taxonomy: type specimens: cotypes
Systema Naturae. See taxonomy: L i n n a e u s
systematics. See taxonomy
Tabulata. See C n i d a r i a : Anthozoa
Taconian. See Taconic Orogeny
l a c o n i c Orogeny, M o u n t a i n s , 3, 5, 8, 62, 216, 220, 233,
2 9 2 - 2 9 3 , pl. 12
Taeniaster. See Echinodermata: Ophiuroidea
tangential thin-sections. See techniques: thin-sections
Tanners Creek Formation, 44
T a p a n i l a , Leif, 210
taphonomy, 1 , 4 - 8 , 2 7 , 4 6 , 52, 56-57, 60, 7 9 , 9 2 , 101, 102,
118, 119, 120, 125, 127, 129-130, 132, 133,
134-135, 137, 147, 148, 151, 153, 155, 163, 169,
188, 224, 231, 282, 289, 290, 293. See also chap-
ters on individual groups of a n i m a l s ; death
assemblage
a l i g n m e n t , 77, 142, 145, 178, 196, pl. 8, pl. 10
bioimmuration, 209, 236, 242, 243, 244, 281, 322. See also
Catellocaula
body fossils, 6, 143, 203, 204, 212, 281, 293
casts, 6, 210, 282, 288, 292
deformation (post-mortem), 8, 130
disarticulation, 7, 8, 114, 188, 223, 224
molds
composite molds, 120, 121, 129, 130
external molds, 6, 117, 129, 281, 285, 287, 288. See also
bioimmuration
internal molds, 6, 101, 117, 120, 121, 122, 123, 125, 126,
127, 129, 134, 135, 136, 137, 139,228, 229, 285,
287, 288, 292
Steinkern. See internal mold
phosphatization, 223. See also c a l c i u m phosphate
pyritization, 153, 160, 168, 190
silicification, 163
soft-tissue preservation, 79, 142, 143, 168, 190
trace fossils, ichnofossils, Lebensspuren, 6, 24, 56, 60,77, 138,
143,151,155, 202-212, 2 2 4 , 2 2 6 , 2 3 5 , 2 6 1 , 2 8 1 ,
284,285, 286,287,293. See also incertae sedis
Allocotichnus: A. dyeri, 204
Asaphoidichnus: A. trifidum, 202, 203, 204
Asteriacites: A. stelliforme, 2 0 8 , 210
behavior categories, 2 0 4 - 2 1 1
c u b i c h n i a , 207, 2 0 8 , 209, 210, 211
d o m i c h n i a , 202, 203, 205, 206, 2 0 8 , 209, 210, 211
fodinichnia, 202, 2 0 3 , 2 0 5 , 2 0 8 , 211
i m p e d i c h n i a , 210, 211
pascichnia, 202, 2 0 3 , 2 0 4 , 211
repichnia, 202, 203, 2 0 4 , 2 1 1
Blastophycus. See incertae sedis
borings, 74, 7 5 , 7 7 , 9 0 , 9 3 , 123-125, 130, 182, 203, 206,
2 1 0 , 2 1 1 , 2 3 6 , 2 4 2 , 2 4 3 , 2 4 4 , 293
Corallidomus. See Mollusca: Pelecypoda
Petroxestes: P. pera, 206
Sanctum, 93, 244
S. laurentiensis, 206, 210
Trypanites, 75, 7 7 , 9 3 , 2 0 6 , 210,
242, 244
burrows, burrowing, 6, 7, 103, 130, 131, 138, 151, 152,
Index 343
 153, 155, 189, 190, 2 0 3 - 2 1 2 , 217, 218, 221, 222, subspecies, 39, 285, 292, 294
223, 224, 225, 226, 235, 238, 293 taxon (pl., taxa), xi, 38, 39, 42, 43, 293
Chondrites, 56, 205, 211, 224, 226 type-species, 40, 42, 43
Chondrites type-A, 205, 2 0 8 variety. See subspecies
Chondrites type-B, 205, 2 0 8 Linnaeus, Karl, 38; Systema Naturae, 38
Chondrites t y p e - C , 205, 2 0 8 nomenclature
Cruziana, 202, 203,211 Binomial System of Nomenclature, 38, 204
Diplocraterion, 60, 206, 211, 224, 226, 247 binomen. See species n a m e
D. biclavata, 2 0 6 , 2 0 9 generic n a m e , 37, 41, 42, 43, 286, 291
D. cincinnatiensis, 2 0 6 , 2 0 8 , 2 0 9 scientific n a m e . See species n a m e
D. luniformis, 206 species n a m e , xi, 37, 4 1 , 4 3 , 198, 203, 281, 286, 291
Dystactophycus. See incertae sedis specific n a m e , 37, 39, 43, 286, 291, 293
epircliefs, 204, 284 trivial n a m e . See specific n a m e
Fascifodina: F. floweri, 138 derivation ol n a n u s , 40
fucoids, 203, 211, 285 International C o d e of Zoological Nomenclature
holes (in shells). See borings (ICZN),xi
hyporeliefs, 2 0 4 , 2 0 7 , 2 1 2 , 286 priority, 43, 290
ichnofacies, 2 1 0 - 2 1 1 , 2 8 6 pronunciation of scientific n a m e s , 41
ichnogenus, ichnogenera, 204, 211 synonym, synonymy, 151; junior synonym, 153, 182.
ichnospecies, 204, 2 0 6 , 2 1 0 , 2 1 1 See also priority
Licrophycus: L. flahellum, 261 type specimens, 22, 2 5 , 4 2 , 158, 260, 266, 272, 274, 286
l.ockeia: L. siliquaria, 209, 210. See also Mollusca: cotypes, 42, 283, 286, 289, 292
Pelecypoda holotype, 42, 72, 158, 173, 178, 182, 184, 186, 209, 283,
Palaeophycus, 204, 207, 224, 226 286, 289
Palaeoscia: P. floweri, 209, 212. See also Cnidaria: paratype, 42, 158, 159, 186, 283, 286, 289
Hydrozoa: Siphonophorida: Porpitidae syntypes. See cotypes
Paleodictyon, 202, 203, 204, 205 techniques. See also chapters on individual groups of animals
Petalichnus: P. multipart Hum, 204 acetate peels, 95, 96
Petroxestes. See borings acid dissolution, acid etching, 68, 9 6 , 143, 163, 195, 196,
Rusophycus, 151, 152, 155,202, 203, 204, 210, 226, 235. 223
See also Arthropoda: Trilobita cluster analysis, 231
R.carleyi, 155,207,210 detrended correspondence analysis (DCA), 231
R. cryptolithi, 155, 210 factor analysis, 231
R. pudicum, 152, 1 5 3 , 2 0 7 , 2 1 0 health, safety, 96
Sanctum. See borings microscopy, 35, 36, 50, 89, 97, 187, 198, 275
Spreiten, 206, 209. See also Diplocraterion polar ordination, 231
Trachomatichnus: T. numerosum, 204 thin-sections, 2 9 - 2 0 , 31, 95, 9 6 - 9 7 , 275
Trichophycus ("turkey tracks"), 138, 211, 224, 226 longitudinal, 77, 95, 9 6 - 9 7 , 287, 293
T. venosum, 202, 203, 205 tangential, 95, 9 6 - 9 7 , 287, 293
Trypanites. See borings transverse, 95, 9 6 , 97, 287, 293
I appan, I lelen, 69 'Vechnophorus. See Mollusca: Rostroconchia
Tate Formation, 214 tectonic activity, 3, 5, 8, 1 0 , 5 3 , 6 1 , 6 2 , 215, 217, 233, 292
taxon, taxa. See taxonomy: L i n n a e a n Hierarchy Teichert, C u r t , 43
taxonomic diversity. See diversity temperature. See environment
taxonomy, 3 7 - 4 3 , 89, 198, 2 0 3 - 2 0 4 , 282, 283, 286, 288, 289, tempestite, 60, 293
290, 292, 293. See also chapters on individual Tentaculites. See Tentaculitoidea
groups of a n i m a l s Tentaculitoidea, tentacuhtoids, 144-145, 247
based on evolutionary relationships, 39 Tentaculites
Linnaean Hierarchy, 3 8 - 3 9 T. richmondensis, 142, 145
class, 38, 39, 282, 289 T. sterlingensis, 145
diagnosis, 42, 283 tenuis, Plectodina. See conodonts
family, xi, 38, 39, 285, 286, 289 tenuiseptum, Manitoulinoceras. See Mollusca: Cephalopoda
genus, genera, xi, 38, 3 9 , 4 0 , 4 2 , 4 3 , 283, 285, 286, 288, Terrill Formation, 214
291. See also ichnogenus, ichnogenera tessellatus, Cryptolithus. See Arthropoda: Trilobita
ichnogenus, ichnogenera. See taphonomy: trace fossils Tetradium. See C n i d a r i a : Anthozoa: Tabulata
ichnospecies. See taphonomy: trace fossils Tetraphalerella. See Brachiopoda: Articulata
k i n g d o m , 38, 287, 289 Thaerodonta. See Brachiopoda: Articulata
order, 38, 39, 282, 285, 289 thanatocoenose, thanatocoenosis. See death assemblage
phylum, phyla, 38, 118-119, 282, 287, 289 T h e Society for S e d i m e n t a r y Geology. See S E P M
species, xi, 37, 38, 3 9 , 4 2 , 4 3 , 283, 286, 291, 292. See Thecidellina. See Brachiopoda: Articulata
also ichnospecies thin-sections. See techniques: thin-sections

344 Index
Thompson, Esther H., 120 undatus, Phragmodus. See conodonts
ticks. See Arthropoda: Chelicerata: Arachnida undulata, Aulaeera. See Porifera: Stromatoporoidea
tiering. See ecology: ecosystems United States Department of Agriculture, 22
time. See also geologic time-units United States Geological Survey, 20, 22, 29, 30, 32, 33, 35,
absolute age, absolute dating, 4 6 , 6 1 , 62, 63, 279, 290 51, 2 6 1 , 2 6 6
radiometric dating, 1 , 4 6 , 6 2 , 6 3 , 279, 290 United States National M u s e u m , 22, 30, 32, 35, 260, 261. See
isochronous surfaces, 46 also National M u s e u m of National History;
relative age, relative dating, 45, 4 6 , 6 1 , 62, 195, 281, 290. Smithsonian Institution
See also biotal succession University of C h i c a g o , 22, 25, 28, 262
superposition, 46 Walker M u s e u m , 2^
time-averaged accumulations, 7, 230, 231, 234, 293 University of C i n c i n n a t i , 14, 16, 18, 20, 21, 22, 25, 26, 27, 28,
time-environment d i a g r a m , 240 29, 32, 33, 34, 3 5 , 4 8 , 8 0 , 9 0 , 9 3 , 110, 138, 142,
time-scale, geologic. See geologic time-scale 151, 155, 156, 170, 180, 203, 209, 225, 256, 260,
time-stratigraphic units, 46 261, 263, 264, 272, 275, 298. See also M c -
Tobin, Rick C, 4 4 , 5 3 , 54, 55, 56, 57, 312 Micken Hall
Townsend, M . ) . , 212, 297 University of M i c h i g a n , xiii, 178, 273
trace fossils. See taphonomy Utica Group, Utica S h a l e , 4 7 , 4 8 , 160
Trachomatichnus. See taphonomy: trace fossils
Trammel, R. L., 12 vacuum, Foerstephyllum. See C n i d a r i a : Anthozoa: Tabulata
transverse thin-sections. See techniques: thin-sections Vail, Peter R., 4, 320
Transylvania College, University. See Kentucky: Lexington V a l l a n d i g h a m , George L., 259, 275
Treatise on Invertebrate Paleontology, xi, 35, 144, 145, 255, vallandighami, Cyrtoceras. See Mollusca: C e p h a l o p o d a
297, 300, 301, 304, 307, 308, 319, 322 vallata, Catellocaula. See Catellocaula
Trematis. See Brachiopoda: luarticulata Van Cleve, J. W., 275
Trepostomata. See Ectoprocta Vanuxemia. See Mollusca, Pelecypoda
Treptoceras. See Mollusca: Cephalopoda: Actinoceratoidea varihrachialus, Cincinnaticrinus. See Echinodermata: Cri-
Treptoceras duseri shale. See Waynesvillc Formation noidea: Disparida
Triarthrus. See Arthropoda: Trilobita: Olenida varicosum, Cyclonema. See Mollusca: Gastropoda
Triassic, xix, 4, 82, 199 variety. See taxonomy: L i n n a e a n 1 Iierarchy: subspecies
Trichophycus. See taphonomy: trace fossils Vaupel, Ernst H., 28, 32, 273, 275
Tricopelta. See Arthropoda: 'Trilobita: Phacopida venosa, Ropalonaria. See Ectoprocta: Ctenostomata
Tridacna. See Mcrllusca: Pelecypoda venosum, Trichophycus. See taphonomy: trace fossils
trifidum, Asaphoidichnus. See taphonomy: trace fossils Verne, Jules, 40
Trilobita. See Arthropoda Veryhachium. See acritarchs
Trimble County, Kentucky, 124, 129 Virginia, 3, 162, 315
Bedford, 129 Virginia. See Chordata: Vertebrata: Reptilia
trivial n a m e . See taxonomy: nomenclature: specific n a m e volcanic ash beds, 1, 7,62, 63, 163, 285, 287. See also K-bentonite
Trollope, Frances, 274 volcanic eruptions, 7, 62
tropidophora, Paupospira. See Mollusca: Gastropoda vorticellatus, Streptaster. See Echinodermata:
truncata, Ischyrodonta. See Mollusca: Pelecypoda Edrioasteroidea
Trypanites. See taphonomy: trace fossils: borings V u l i n e c , Kevina, 7 4 , 8 6 , 100, 117, 125, 147, 195
Tubastraea. See C n i d a r i a : Anthozoa: Scleractinia
tuberculata, Brachiospongia. See Porifera Wabash C o l l e g e . See Crawfordsville, Indiana
tuberosa, Pattersonia. See Porifera wackestone. See limestone
tunicates. See Chordata: Urochordata W a h l m a n , Gregory P, 120
turbidity. See environment Walcott, C h a r l e s D., 25, 199
"turkey tracks." See taphonomy: trace fossils: Trichophycus Walker M u s e u m . See University of C h i c a g o
turnhulli, Hercochitina. See chitinozoans Warder, John Aston, 275
Turner, Peter, 199, 316 Warn. John M . , 172, 173, 182
tusk shells. See Mollusca: Scaphopoda Warren County, Ohio, 60, 105, 126, 136, 172
Twenhofel, W. H., 274 Warshauer, Steven M., 163
Twitchell, George B., 273, 274-275 Washington, pl. 3, pl. 9
type specimen. See taxonomy Washington, George, 106
type-species. See taxonomy: L i n n a e a n Hierarchy water movement. See environment
typicalis, Cymatonota. See Mollusca: Pelecypoda W a u g h , David A , 9 1 , 304
typicalis, Geniculograptus. See Hemichordata: Graptoloidea wave base, 221. 223. 22>
Tyrone Limestone. See High Bridge Group normal wave-base, fair-weather wave-base, 56, 2 2 1 , 222,
223, 240
Ulrich, E. O, 6, 18, 19, 20, 2 8 - 3 1 , 32, 33, 34, 3 5 , 4 7 , 6 7 , 6 8 , storm wave-base, 56, 221, 240. See also storms
87, 111, 120, 123, 126, 127, 142, 143, 144, 160, W a y n e County, Indiana, 67, 75, 228
172,173, 176, 261, 263, 267, 271, 273, 274, 277 W a y n e State University, 176

Index 345
Wavncsvillc Formation, 48, 53, 56, 60, 63, 75, 79, 87, 107, "Lower Member," 44, 2/4
111, 115. 120, 124, 126, 128, 129, 130, 136, 138, "Upper Member," 44, 214
142, 149, 150, 151, 153, 157, 160, 162, 172, 176, Saluda Member. See Saluda Formation, Member
180, 184, 196,2J4, 225, 229, 307, 314, 317, pl. 6, Whitfield, R. P., 134, 137, 142, 145, 229, 2 7 6 - 2 7 7
pl. 10. See also Brookville Formation: W a j nes- W h i t t i n g t o n , Harry B., 160
x ilk- Member Whittlesey, C h a r l e s , 269, 277
Blanchester M e m b e r , 44 Willet, 268, 277
Clarksville M e m b e r , 4 4 \\ illiains. I Icnn Shaler. 23
Fort Ancient M e m b e r , 44 williamsae, Manitoulinoceras. See Mollusca: Cephalopoda
Marble Hill bed, 124, 125, 127, 129 williamsae, Megalograptus. See Arthropoda: Chelicerata:
"Treptoceras duseri shale," 60, 136, 137, 138 Furypterida
Waynesville, Ohio, 120, 182, 184, 185, 256, 259, 266. See williamsae, Zittelloceras. See Mollusca: Cephalopoda
Warren County, Ohio W i l m i n g t o n , Ohio, 260, 273, 276
waynesvillensis. Vanuxemia. See Mollusca. Pelecypoda Wilson, Mark A., 124, 209, 210, 281, 296, 298, 311, 315
Weaver, T h o m a s , 75, PI S W i l s o n , W i l l i a m W., 277
W e e d o n , M . J . , 82, 309 wilsoni. Polygnathus, 277
Weichold, C h a r l e s , 93, 275 Woodward High School, C i n c i n n a t i , Ohio, 18, 26, 32, 33, 34
"Weichold doughnut." See Ectoprocta Woodward, Henry, 26
"Weichold ring." See Ectoprocta: "Weichold d o u g h n u t " worms and "worms," 5, 6, 77, 86, 103,142-145, 152, 153, 155,
W e l c h , James R., 182 182, 196, 206, 210, 222, 224, 241, 242, 244, 246,
Welch. I. B . 276 282. See also Annelida
welchi, Megalograptus. See Arthropoda: Chelicerata: Worthen, A. H., 47, 126, 137, 170, 172, 174, 266, 268, 269,
Furypterida 277,311
wellsi, Acanthochaetetes. See Porifcra: sclcrosponges, coral-
line sponges Xenocrinus. See F c h i n o d c r m a t a : Crinoidea: C a m e r a t a
Wcsschnan Tongue. See Kope Formation Xiphosurida. See Arthropoda: Chelicerata
Wessels, C h a r l e s , 276
Western Academy of Natural Sciences, 15, 16, 17, 21, 259, Yale University, 20, 32, 260, 274, 275
260, 261, 262, 263, 265, 269, 272, 273, 275, 276 Yochelson, Ellis L. 34, 35, 36, 124, 300
Western M u s e u m , 16, 260, 263, 264, 265, 268, 274, 276
Wetherby, A. G., 17,18, 26-27, 29, 3 2 , 2 6 5 , 2 7 0 , 272,273, 276 /.itk'lloceras. See Mollusca: Cephalopoda
whelks. Sec Mollusca: Castropoda: Neogastropoda /.one. See biostratigraphic units
Whitewater Formation, 44, 48, 49, 53, 56, 63, 67, 68, 75, 79, zooid. See Ectoprocta, graptolites
109. I l l , 120, 123, 126, 127, 134, 137, 1 3 8 , 2 0 8 , Zygocycloides. See Echinodermata: Cyclocystoidea
214, 223 Zygospira. See Brachiopoda: Articulata

346 Index
ABOUT THE AUTHORS

DAVID L . M E Y E R i s Professor o f G e o l o g y a t t h e U n i v e r s i t y o f C i n c i n n a t i . H i s
r e s e a r c h interests are c h i e f l y i n t h e f i e l d o f i n v e r t e b r a t e p a l e o n t o l o g y , a n d
they extend to studies of living and fossil reefs, paleoecology, and
taphonomy.

R I C H A R D A R N O L D DAVIS i s Professor o f B i o l o g y a n d G e o l o g y a t t h e C o l l e g e
of M o u n t St. Joseph in C i n c i n n a t i . His r e s e a r c h interests f o c u s on fossil
a n d l i v i n g c e p h a l o p o d s , s y m b i o s e s a n d s i m i l a r r e l a t i o n s h i p s i n t h e fossil
r e c o r d , a n d t h e history o f t h e g e o l o g i c a l s c i e n c e s .

S T E V E N M . H O L L A N D i s Professor o f G e o l o g y a t t h e U n i v e r s i t y o f G e o r g i a ,
Athens. His research interests are i n s t r a t i g r a p h y , p a l e o n t o l o g y , a n d
paleoecology.