A Sea without Fish: Life in the Ordovician Sea of the Cincinnati Region (Life of the Past)

A SEA WITHOUT FISH LIFE IN THE ORDOVICIAN SEA OF THE CINCINNATI REGION David L. Meyer and Richard Arnold Davis With a ...

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A SEA WITHOUT FISH LIFE IN THE ORDOVICIAN SEA OF THE CINCINNATI REGION

David L. Meyer and Richard Arnold Davis With a chapter by Steven M. Holland

Indiana

University

Bloomington

C?

Press Indianapolis

This book is a publication of Indiana 601

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© 2009 by Richard Arnold Davis and David Lachlan Except chapter All

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Meyer

Holland

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Cataloging-in-Publication

Data

Meyer, David L. 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.

p. cm. — Includes ISBN 2.

bibliographical

978-0-253-35198-2

references (cloth:

and

alk.

Fossils—Ohio—Cincinnati Region.

index.

paper) I.

Davis,

1. R. A.

II. Title. QE726. 2. M49 560'.

2008

17310977178—dc22 2008020036

1

2 3 4 5

14

Holland.

(Life of the past)

13

12

11

10 09

Paleontology—Ordovician. (Richard Arnold),

date-

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 , reg 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 xiil

PREFACE 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

VII

THE OTHER BIVALVES

9 117

10 143

11 147

12 167

13 195

14 203

15 215

16 229

Molluscs HARD, BUT WITH A SOFT CENTER

Annelids and Worm-Like Fossils <

Arthropods TRILOBITES AND OTHER LEGGED CREATURES

Echinoderms A WORLD UNTO THEMSELVES

Graptolites and Conodonts OUR CLOSEST RELATIVES?

Type-Cincinnatian Trace Fossils TRACKS, TRAILS, AND BURROWS

Paleogeography and Paleoenvironment BY STEVEN M HOLLAND

Life in the Cincinnatian Sea <

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 Pleistoc 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 southwestern 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

Journal of the Cincinnati Society of Natural History.

Paleontologist,

No single

and

library

the

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 richness 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 ecosystems 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 geolo 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 literature 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 By

international

logical Nomenclature

Groups

of Organisms

a g r e e m e n t of z o o l o g i s t s , is

the

International Code of Zoo-

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 Fossils 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 a n d the Glossary

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 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 anatomic 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 stud 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 artistic 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 reprod 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 Bergstrom ( 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 History 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 disc 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 University 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 Bradley D e l i n e , and Austin Hendy.

xiv

Acknowledgments

Buick, Katherine Bulinski,

REPOSITORIES OF FOSSILS ILLUSTRATED IN THIS BOOK

BMNH C M C IP FMNH

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

University,

of Comparative Z o o l o g y

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 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 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

Museum

A SEA WITHOUT FISH

Figure

1. 1.

7999 Geologic Time Scale.

Reprinted by permission of the Geological Society of America.

order to read this chart in stratigraphic order, of the Precambrian column, umn,

adding

the bottom of the Paleozoic column

the bottom of the Mesozoic column

zoic column

to

the

read the columns from bottom

to

to

In

starting at the bottom

the top of the Precambrian

the top of the Paleozoic column,

top of the Mesozoic column.

to top,

col-

and the bottom of the Ceno-

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 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

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

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 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 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

the

interior,

none

bluffs at Cincinnati, Here

the

cian

Ohio.

Upper Ordovi-

rocks are almost

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

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

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.

as perfectly fossils

can

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,

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

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 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 richness 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 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 -

and even

common Almost in

the

fossils more

than every

pebbles. museum

the world has speci-

mens from

this locality.

W i l l i a m Lee S t o k e s

cinnati was in the S o u t h e r n H e m i s p h e r e . 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

1960, 1 8 8 - 1 8 9

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 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 ( 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

[The

Ordovician

represents largest

one

major

radiation] of the 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

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

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

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

the

appearance

for the

to

econext

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

families

through

Phanerozoic.

the

The

uppermost the sum

metazoan heavy

curve of the

"evolutionary

depicts three

faunas, "

each

shaded

while

the stippled por-

tion

below the

curve

total

represents

diversity

not

for by the each

residual

accounted

three compo-

nent faunas. for

differently,

Taxa listed

evolutionary

fauna are those taxa that contribute

most

heavily

to the diversity of that fauna. I = Cambrian Fauna,

II = Paleozoic

Fauna, and III = Modern Fauna.

From

Sepkoski

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

(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

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

of The

Pale-

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; Seil 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 surv 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

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

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

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

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 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 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

are

crust,

accustomed of North

of

the past when nent

was

so

beneath

could have

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

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 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 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

one

earth's

us to imagine a time in

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

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 -

firma,

and it is difficult for

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 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 Amer-

to

our contisubmerged

the sea

that fish

swum

directly

the Atlantic

Ocean

the Pacific Ocean,

from Hudson Bay to the

Gulf of Mexico.

such a

million years ago the

Yet

time did exist 450

epeiric sea

when

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,

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

from

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. 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,

1960, 6 8

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 redistributed 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 skeletons 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

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 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 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

will see,

dence of the early, jawless 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

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.

elsewhere.

These nutrients m a y have

Introduction

Cincinnatian

rocks contain no fossil evi-

3

Late Ordovician

Figure 1. 3.

Global sea

level curves

for the

Phanerozoic.

A.

curve, B. (1977).

Hallam

Vall et al. curve

From Hallam

(1984) and reprinted by permission Reviews. more

of Annual According

recent

to

studies

(Miller et al. 2006),

maxi-

mum rise of sea level in the

Cretaceous

lower than mates, 50 m level,

was

these

reaching

esti100 m ±

above present sea 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 add 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 diversity 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. Upper

Thickness of

Ordovician

strata

in relation to the ancestral

Appalachian

Moun-

tains (tectonic land)

that

was

uplifted during

the

Late

Ordovician

Orogeny.

Taconic

Contours

lines of equal rock ness (isopachs). (1951,

are thick-

From Kay

figure 4) and re-

printed by permission the

of

Geological Society

of

America.

The

organic

remarkably

preserved

for so

a

rock,

argillaceous stone,

significance of variable preservation can be gained by c o n s i d e r i n g aspects 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 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.

appears

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

compact lime-

Its

deposition

to have gone on

very tranquilly, as the Lingula

has been met with

in its natural and erect position,

as if enclosed in

mud when alive, standing

Nature of the laving O r g a n i s m

a blue

not unlike the lias

of Europe.

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

well

ancient

especially those

occurring in

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

remains

here are

on

or still

its peduncle.

C h a r l e s Lyell 1845, 49 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 fossilization 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 softb 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 bia 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 (burr 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 seaw 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 calcite, 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 feat 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 arag 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 . Prese 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

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

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

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 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 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 details o f R o m a n life.

paleoecologist

the living

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

in

the churchyard,

then

Derek V. A g e r 1963, 184

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

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 trilobites 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 settleout 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 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

and

only after many

visits by grave robbers.

volcanic eruptions than about day-to-day processes.

cycle of scavenging, decay, and destruction.

vil-

lage but only the bodies

Thus, processes of mortality are of 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 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

must

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 Life

1.

Life a s s e m b l a g e

Death assemblage

Articulation

good

disarticulated

Breakage

rare

common

Abrasion

rare

common

Preserved in life position

maybe

not

Size-sorting

uncommon

possible

Characteristics of

Assemblages

Death

and

Assemblages

History

often

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 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

. . . our search for a mechanism range

forces us

far beyond

Cincinnati consider

region the

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

the and

geologic

tory of much North

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

to

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 his-

of eastern

America,

especially

continental

collisions

referred to as the

Taconic,

Acadian,

Alleghenian

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 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 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 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 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

and

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

orogenies.

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 fossils 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 associated 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 branches, Arch

(through

the

Kankakee

(through

and its

the Findlay Ohio) and Arch

Indiana).

Shaded areas

depict

of Ordovician

bed-

rock;

heavy lines

indicate

Silurian-Devonian that defines and

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 subm 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

out-

crop

9

contact

the Findlay

Kankakee

branches.

Figure 1. 6. east-west from

Diagrammatic

cross

across Ohio.

the to

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

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 -

Cincinnati

near Portsmouth,

From Potter (1996)

and

reprinted

sion

of the

Geological

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 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

near Bedford in

south-central Arch

section

by permis-

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 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 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 dist 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

Kentucky

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).

Survey.

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 , prev 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 virtually 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.

Streambed

A.

Natural exposure of Cincinnatian strata in the bed of Stonelick Creek,

outcrops

have

wide surfaces

about fossil distribution and orientation. by participants in a

1981

Here,

the

beds

(bedding

fossil-rich

Limestone (Maysvillian Stage).

B.

Clermont County,

that provide

west of Maysville,

Mason County,

(Edenian Stage) Kope Formation,

Photo by Paul E.

Ohio.

information

exposed and examined

One of the most extensive roadcut

1. 3 km (0. 8 miles) in length and about 75 m (250 ft) high,

leading to the Ohio River,

posed are the lower Cincinnatian

planes)

Bellevue Limestone is

Geological Society of America field trip.

exposures of Cincinnatian strata, Route 3071,

of fossiliferous

Kentucky.

and the highest strata are

Potter.

Introduction

11

along Kentucky

The lowest strata exthe Bellevue

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,

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

Ohio.

This is a ten acre hillslope where

construction

posed

four

formations:

the

stone,

Fairview

Miamitown

Bellevue

s h o w s that t h e a b u n d a n c e o f 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 i s t h e result o f m a n y i n t e r a c t i n g 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 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

Lime-

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

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 -

Trammel donated

cepts of paleontology and geology originated from research on C i n c i n n a -

mation. R. L.

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. T h e foregoing overview

fossiliferous

Formation, Shale,

ex-

The

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

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,

educa-

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

geology ogy.

paleontol-

The park has easy

access, and

Ordovician

and parking

includes

space,

interpretive

signage

and

in situ

fossil shell pave-

ment.

Because

abundance face

a

of the

of fossils,

collecting

specimens

protected,

is

sur-

of small permitted.

12

A Sea without Fish

Figure 2. 1.

Members of the Cincinnati School of Paleontology who were amateur paleontologists: B.

James, publisher and owner of the James Book Store. for. D. ing.

C.

B.

Dyer;

who,

after he retired as a maker of soap and candles,

Photograph of Dyer from an old album in

Davis); all others

from

A Sea without Fish

C.

A.

U.

University of Cincinnati.

P.

Charles Faber, real-

devoted himself to fossil collect-

the possession of Richard Arnold Davis (© Richard Arnold

the Department of Geology,

14

S. A. Miller, attorney.

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 assiduously 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 an

A.

C o v e r of

7 8 4 9 publication of

the

Western

Natural

Academy of

Sciences

pub-

lished by U. P. James, a member

of the

School

of

Cincinnati

Paleontology, B.

and his brother. Cover

of

the

Quarterly Science,

Journal volume

num-

1874, by S. A. Miller,

member of the

nati

School

ogy.

C.

Journal nati

Cincin-

of PaleontolCover of the

of

the

Society

of

History,

volume

ber 7.

D.

CincinNatural 1,

num-

Cover of The

Paleontologist, 4,

of 1,

7, published in Janu-

ber ary, a

Cincinnati

Number

published in July

1879

by U. P. James, a member of the of

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

Cincinnati School

Paleontology.

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. Woodward High School, M. Nickles. Ulrich, Bassler).

C.

where he

George W.

Harper,

"Friendly enemies": left to right, August F. Foerste, Amadeus W.

during the International Geological Congress in Photograph

B.

Washington,

of Wetherby courtesy of the Museum

President and Fellows of Harvard College); University of Cincinnati;

"Friendly enemies,"

78

long-time principal of

facilitated the start of the careers of students Ray S.

that of Harper, from

A Sea without Fish

the

D.C,

Bassler and John

Grabau, and Edward O.

1934 (picture taken by Ray S.

of Comparative Zoology,

Harvard University (©

courtesy of the Archives and Rare Books Library,

Department of Geology,

University of Cincinnati.

Figure 2.4.

Field work in the early days.

exposure of the Kope Formation,

Cincinnati,

and Point Pleasant Formations on exposures are now underwater. seum of Natural History,

A.

John M. Nickles (left) and Ray S. Bassler, collecting from an

Ohio,

1900.

B.

E.

O.

Ulrich at the contact between the Kope

the banks of the Ohio River below Covington,

(A and B

Smithsonian

from Bassler Archive,

Institution,

Washington,

Kentucky,

Department of Paleobiology,

D.C,

1901.

courtesy of JoAnn Sanner.)

Science in the Hinterland

19

These

National Mu-

Figure

2.5.

Four Cincinnatians

Professor, Yale University. National Museum. ology,

D.

who became leading professional paleontologists.

B. John M. Nickles, U. S. Geological Survey, E.

O.

Ulrich,

U. S.

University of Cincinnati.)

20

A Sea without Fish

Geological Survey.

1942.

A. C.

Charles Schuchert,

Ray S. Bassler, U. S.

(All photographs from the Department of Ge-

Figure 2.6. crops in where

Urban out-

Cincinnati,

members

Cincinnati

of the

School

found

their inspiration.

A.

Bellevue House, site

on

The the

of the present Belle-

vue Hill Park, ca.

1895.

The

section

stratigraphic

exposed the

below

Kope

spans

begins

in

Formation,

the entire Fairview

Formation

and

Miam-

itown Shale (a small "step" below crest), is

and

topped by the Bellevue

Limestone.

Clifton

Av-

enue runs below the posure,

was desig-

the

type section

nated as of the

ex-

which

Fairview Formation

by Ford (1967).

(Image

courtesy

Cincinnati

of

the

Historical Society

Library,

Cincinnati

Museum

ter.)

Exposure of

B.

Maysvillian

Cen-

strata

(prob-

ably Corryville and Mt. Auburn

Formations)

corner and

of

Clifton

Calhoun

(right),

at

Avenue

Street

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 University of Cincinnati lege of Law.

Bassler Archive, ment

of

National

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,

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,

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,

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

Depart-

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

Museum

of Nat-

Smithsonian Washington,

courtesy

Sanner.)

Col-

(From the

of JoAnn

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 interests 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 bec 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-tod 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 , professor 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 Castle, Worcestershire,

England.

C. B. Dyer

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 lifet 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 fortunate 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 society 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 pap 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 indiv 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 can

Mesozoic

American Scientists

and

Cenozoic

Geology and

(1889,

Palaeozoic Fossils

Geology

and

Palaeontology for the

with s u p p l e m e n t s in

1892

(1877),

Palaeontology

Use of Amateurs,

and

1897).

North Ameriand

(1881),

Students,

North and

' 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 professionals, 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 reported 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 professor 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 borrow 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;

1951,1981,1982, 2002; M e r r i l l 1924;

Caster

Utgaard

pers. c o m m . ; C r o n e i s Sherborn

1940;

Anon.

1963; C u f f e y , 1875,1878.)

Davis, and

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, Pennsylvan 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 app 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 University 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 MagneticCity, 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-aut 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 B;

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, 1956; W e t h e r b y 1879a, 1879b, 1880, 1881; A n o n . 1876, 1878,

Nickles

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 exa 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 sec 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 posit 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 1876—1877 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 , propositions 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 janifor, 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 proteg 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. Onco 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 MaryC 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 lessi 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 proxim 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 disputeas 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 extraordinarily 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 stratig 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 rev 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 professional 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 byN e w York, Harvard, and Yale Universities. He was a m e m b e r of the National A c a d e m y of S c i e n c e s , and was elected president both of the Paleontological 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 rec 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" (Nickles 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 Bryozoa ( 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 bibliography 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

notated Bibliography of Economic

of

Geology,

North and

American the

Geology,

t h e An-

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 presid 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 facilitated 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 Invertebrate 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 classification 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 lifet 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

Retrospect

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 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 standard 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 individuals 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 org 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 , especially 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 growi 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 related 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 chord 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 Species

Felis 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 hierarchy. 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 characteristics 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 together 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 (alt 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,

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 -

half-Latin,

and always

difficult

pronounce,

many

ished scorch

terms

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

unpol-

that would

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, 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 .

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] 1992,

Journey to

Centre

of the

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

the

Earth,

type-Cincinnatian called 4

Rafinesquina

after C. S. R a f i n e s q u e (1783-1840),

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 Cincinnatidiscus 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 richmondensis 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 trilobite Flexicalymene c o m m o n l y is f o u n d flexed into a b a l l . b r a c h i o p o d Petrocrania

scabiosa

The i n a r t i c u l a t e

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 antepenultim 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 nomen 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 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 -

"Musical N a m e s "

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 Glyptocrinus, 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

name—in

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 genericn 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

From Schumacher (1984, logical Survey.

figure 2),

1955 through

This chart shows stratigraphic subdivisions

researchers for different parts of the Cincinnati Arch region. were based largely on differences in Lineback (1966),

Hatfield (1968),

fossil content.

Gray (1972),

from his study.

In

the Hatfield column,

of the

Cincinnatian

Subdivisions in

A Sea without Fish

Series proposed by different

the Caster et al.

(1955) column

and Lee (1974) columns were based on general

Hay (1981) and Tobin the

(1986) used both lithologic as well

vertical lines indicate parts of the section

Units separated by jagged lines indicate lateral changes in

44

From Davis and Cuffey (1998).

Broader subdivisions such as those of the Brown and

Peck (1966),

characteristics of the rocks (lithology) and bedding. as paleontologic aspects.

1986.

courtesy of the Ohio Department of Natural Resources Division of Geo-

rock

characteristics

excluded (faces)

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 howit 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 reg 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 different. 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 , resulting 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 intervals 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 layers 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 Period 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 (usually 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 ,

Lithostratigraphic

their size, shape, and a r r a n g e m e n t .

The fundamental 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 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

Units

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 fourfold 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 excellent 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 Lorraine 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 conditions prevailing at the close of the Trenton to the time of the Lower Richmond, 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 differentiate 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 vertic 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 (Figure 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 postW 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 percent

composition

allochem

of

fraction

nal

groups and algae in

the

upper (Saluda

Whitewater and lower tion

and

Formations) (Kope

Forma-

to Saluda Formation)

parts Series.

of the

Cincinnatian

(Allochems

fragments

are

of fossils

other discrete grains limestones.) from 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 classif 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 revision 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 following 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 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

or in

Adapted (1975),

tesy of Wayne D.

1975; T o b i n 1982).

the

of fau-

cour-

Martin.

Figure 4 . 3 . semblages. as seen

Variation in limestone deposition in relation to benthic fossil asSmaller circles

not the same. tive

depict

the

in petrographic thin section

bonate sediment

of skeletal grains (mud),

has skeletal grains in between grains.

A

tergranular spaces.

and

and calcium

the

In

produce a

est level of water motion,

50

of organisms,

A Sea without Fish

(spar).

mud filling

A

types

the relapackstone

the spaces

cement filling "floating" in a

the inmud ma-

and a mudstone has less

causing

resulting in

A grainstone

water movement is

fine-grained sediment either packstones or

water movement at margins

wider variety of limestones.

Martin.

10% grains,

with skeletal grains,

variable

grains to be cemented. D.

limestone

All limestones are fine-grained car-

the interior of a benthic assemblage,

dense growth

More

cement with

wackestone has skeletal grains

to be deposited along wackestones.

carbonate

grainstone is grain-supported with A

10% grains.

other allochems,

contact (grain-supported)

trix (mud-supported) with more than retarded by

of different

Different types of limestone will form depending on

contributions

than

character

under the microscope.

of assemblages

usually reflects

by which mud is winnowed,

From Martin and Hauer (2006),

will

the high-

leaving only skeletal courtesy of Wayne

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 , geologicresearch 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 lithostratig 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 ass 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 classification

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 southeastern 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 classification 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-stratig 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 chara 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 frees 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 est 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 acritarchs), 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 activelys 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.

Cycles and

2004).

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 transition 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 ) traceable

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, 74).

figure

The base of the Cin-

cinnatian is at the base of the Kope Formation the

in

left-hand column;

Corryville tinues

Formation

above

Limestone,

the

and

the con-

Bellevue the

Waynesville

Formation

overlies

Oregonia

the

Formation

and

continues

to the top of the Cincinnatian tion).

(Saluda

Forma-

Arrows

thickness

indicate

of each

cycle in meters.

major Note that

in each cycle, shale is replaced

by

limestone

toward the top,

indicat-

ing a shallowing transition.

upward

Rock

symbols

are: brick pattern = limestone,

dashed pattern

shale,

wavy pattern =

wavy-bedded

limestone,

rhombic pattern mite,

=

=

dolo-

small circles = nod-

ular limestone.

More re-

cent work by Holland has recognized

six

major

shoaling-upward or sequences same

over

stratigraphic

cycles the 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

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

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 shallowi 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

three 15.1).

cycles

(see

Figure 4 . 5 . Society

Geological

of America

Trip, 1981.

A.

Field

Left to

right: leader Rick C. Tobin, Carr, den, Dill.

participants

Tim

Thomas W. Amsand Robert F. B.

Professor

Wayne A. Pry or with poster natian cycles

illustrating

Cincin-

shoaling-upward and

fades

relationships.

54

A Sea without Fish

Figure 4.6. scales

A.

of

Three

Cincinnatian

stratigraphic

cyclicity.

From Tobin (1982,

figure

30). In storm cycle, basal bed with oblique

wavy and lines

indicates

cross-stratification duced

by

currents

pro-

storm-induced

and

waves.

B.

of three

concepts

cinnatian cles.

of Cin-

meter-scale

cy-

From Holland et al.

(1997, of

Comparison

figure 1), courtesy

the Journal

ogy.

of Geol-

In each column,

width

of bed pattern

dicates

lithology

in-

as

shown in scale (pack = packstone, stone).

grain

Arrows

= grainindicate

thickness of cycle in ters.

Tapering

beside cate ing

(wider = shallower). facies

transported

contain ma-

from

place

than

proximal

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 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 transitions 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 The R i c h m o n -

55

indi-

of shallow-

terial

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

Rocks, Fossils, and Time

column

Distal

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

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 .

each

direction

me-

wedges

farther

of formation fades.

Figure 4.7. deposition across

A.

Storm

and

the

erosion

Cincinnatian

sea floor. From Tobin (1982,

figure 37).

Note

that hurricane rotation correct

for

the

Hemisphere. tions in

B. From

Tobin

figure 36).

indicate ment

Varia-

Cincinnatian

storm cycles. (1982,

is

Southern

layers

of sedi-

deposited

storm events.

Arrows

by

single

Sh = shale,

L = laminated unit, W = whole

fossil limestone,

= siltstone,

S

F = fragmen-

tal limestone, FL = finegrained limestone,

SL

=

storm layer, Ls = any limestone.

Chondrites

is

a trace fossil. A = abundant, 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 , WaynesvilleLiberty-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 reco 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 Harris 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 , edrioaste 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 . of a by

soft

Colonization

mud substratum

Dalmanella

(low di-

versity) is followed by the development pavement of

of a and

erect

shell succession

bryozoans.

Higher diversity is achieved in community

the by

mature crinoids

growing on a firm substratum. Martin

printed by permission

MATURE COMMUNITY

the tary

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 Form 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 .

From Harris and (1979) and re-

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

Society

for

Geology.

of

Sedimen-

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

a middle c o m p o n e n t with thin interbedded limestones (packstones) and

mational

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 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 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

Kope-Fairview contact

fortrace-

able over broad area of Cincinnati A.

Arch.

Geological Society of

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

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

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

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 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 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 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 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 -

Kope

along

examining

Kentucky

in

Route

445 near the Ohio River, Campbell tucky. Kope

County, B.

Formation

overlying

Ken-

Contact of with

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,

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.

Cincinnati,

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 lithostratig 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 during the phase of C i n c i n n a t i a n Lithostratigraphy. Exceptions were the "shingled 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 disc 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 variety 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 soc 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 stratig 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 inc 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 original 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 superposition 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 geologicsuccession of strata was established by r e c o g n i z i n g that strata with characteristic 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 radioactive 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 possible 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 occurring 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-prod 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 (Bergs 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 building) 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 Form 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 1991).

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

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 missing. 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 my—445 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 seq 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 thickness. 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 product 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 Hiatus 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 acc 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 processes ( 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,

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,

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 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 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

Kope

Wayne County, ana.

A.

edenense 569

all Ede-

Formation, Indi-

Veryhachium

Colbath,

(from

x

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.

B. Ordovicium

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-

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,

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,

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

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 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 . 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 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

x 507 (from Col-

4]).

C.

cisphaeridium micraulaxum 1038 A-C

reprinted by permis-

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.

SchweizerNote: 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,

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 D-F.

D.

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 -

C.

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

x indicates the factor.

Cincinnatian

zoans,

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 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

Cyathochitina sp.

OSU 32534,

x

747 (from

Miller [1976, plate 5, ure

6]).

E.

tina

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

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]).

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 -

tina

turnbulli

hirsuta

ure 1]). tian

to a l g a e ( B l a c k w e l l et al. 1984).

dickhauti

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 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

Laufeld, x 287 (from

G-l. G.

Edenian, tion,

William

collection, Kope Forma-

Kenton

County,

Kentucky, x 3.8.

67

al-

Lepidolites

Ulrich,

Heimbrock

fig-

Cincinna-

dasycladacean

gae.

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

13, Conochi-

Miller [1976, plate 8,

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

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

Jenkins,

x 516 (from F.

OSU 32542,

fig-

Hercochi-

OSU 32568,

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,

cf.

campanulaeformis,

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

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

chitino-

all Maysvillian.

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 ,

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

x

Colbath

[1979, plate 7, figure 10]). sion of E.

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.

Colbath,

(from

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

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

fig-

Multipli-

H.

Cy-

clocrinites (Miller), collection, Mt.

darwinii

Stephen

Auburn I.

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 ,

Maysvillian,

Butler County, 1.3.

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-

Felton

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

Formation, Ohio,

most c o m m o n dasyclad, found in the Bellevue Limestone and throughout

x

C. darwinii, sur-

face detail of H,

polygo-

nal facet diameter ~0.4

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

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 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 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

mm.

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 taxon 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 m a t i o n , a n d Faheria anomala

Ulrich

from

the

u p p e r F a i r v i e w For-

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 prov 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-resistant 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 biostratigraphic 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 represent 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 phytoplankton 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 different 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 different 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 flagell 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 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

Figure 6 . 1 . A simple sponge,

showing a

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

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

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 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),

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

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-

a

magnified

view of incurrent

Drawing

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 discovered 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 sclcrosponges 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

cross-

of the body wall. canals

collar cells and chambers. by John

Agnew.

Figure 6.2.

Reconstruc-

tion of a living stromatoporoid,

modeled on

living

a

sclerosponge.

section

A

is removed to

show internal laminae the skeleton. occupies

Living

only

most layer,

the

of

tissue upper-

with excur-

rent canals radiating oscula.

Magnified

shows

surface

tissue

microstructure nae.

from

inset

of

and

lami-

Compare to Figure

6.3D,

below.

John

Agnew.

Drawing by

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 Figure 6.3.

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,

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-

poroids.

A.

A cylindri-

cal

stromatoporoid,

acera

Aul-

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

undulata

(Billings),

MUGM 29618,

Richmondian,

horizon

B.

culata

tuber-

James,

holotype,

CMC IP 209, dian,

Richmon-

Clinton County, C.

Ohio,

tuberosa

(Beecher),

MUGM,

Maysvillian, County, section poroid,

(Billings),

634A2t, Ohio,

x

huron-

MUGM

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-

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 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 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, Aulacera 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 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

Richmondian,

Montgomery

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

( 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

stromato-

Labechia

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

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

Polished cross-

of a

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

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

Fleming

Kentucky, D.

0.3.

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

d e t a i l a n d illustrated b y R i g b y ( 1 9 9 6 ) .

x

A sponge, Pat-

tersonia

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 -

c a n r e a c h 2 8 c m (11

Liberty Formation,

0.3.

ensis

x

A large sponge,

Brachiospongia

often m a y be overlooked b e c a u s e they can resemble bryozoans or corals, 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, 0.3.

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 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 .

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

County,

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

x 0.3.

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-

tion

Cincinnatian

of the

solitary rugose

coral,

Grewingkia

canaden-

sis,

showing

with

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 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 . 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

the polyp

extended

Drawing

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

tentacles.

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 (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 -

by Kevina

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 ,

Vulinec.

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 repres 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 widee 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.

alla attached to bryozoan, [1982, plate 3, Rafinesquina, 3,

figure 5]). Richmondian,

figure 9[).

mondian,

C.

canadensis

dian, Adams County, CMC IP 45413, 10]).

B.

A.

S. divaricans,

divaricans,

x 0.6,

Ohio,

CMC IP 50667,

Clinton County,

135767,

USNM Ohio,

70211,

several cor-

x 2.0 (from Elias

three coralla attached to margin of brachiopod Indiana,

x

1.3 (from Elias [1982,

two coralla attached to brachiopod Lepidocyclus, x 2.2 (from Elias [1982, plate 3, Weaver

Collection,

inset showing epithecal growth-lines,

typical abraded corallum

reprinted by permission

divaricans (Nicholson),

Wayne County,

USNM 40086,

Whitewater-Elkhorn Formations,

A-C

USNM

Whitewater Formation,

(Billings), Ohio,

Streptelasma

Waynesville Formation,

Whitewater Formation, Butler County,

Grewingkia

mondian,

S.

Richmondian,

with broken rim, Wayne County,

figure 8[).

well-preserved corallum, scale in mm.

E.

G.

D. Richmon-

canadensis,

showing circular borings of Trypanites,

Indiana,

of the Paleontological Research

x

1.0 (from Elias [1982, plate 9, Institution.

Poriferans and Cnidarians

75

plate

Rich-

Richfigure

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 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 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 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 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 . 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-

Figure 6.6. Cincinnatian A-E.

colonial corals.

Corallites as seen on external surface or in cross sections of corallum, from Elias (1998).

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,

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, dian.

B.

lum,

a tabulate,

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 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

Foerstephyl-

Richmondian. a

C.

Cala-

tabulate,

Rich-

D.

mondian. pora,

a

Nycto-

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.

highly abraded, encrusted, and bored (Figure 6.5E). External c o n c e n t r i c

dium, a tabulate,

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.

solitary c o r a l s , Grewingkia p r o b a b l y lived

Richmondian,

upright, partly buried

in soft

sediment with the polyp exposed. S o m e encrustation and b o r i n g took 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 activity 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 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

County,

length (from

Elias [1998, figure 5]). G.

Protaraea

densis

richmon-

Foerste,

ing

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,

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

Coral bed, Madison

Kentucky,

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

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

TetraRich-

of hammer 25 cm

5435,

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).

E. F.

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

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

co-

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 ,

poecia,

A. a

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 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

at

same scale x 3.7. A - E

a

MUGM

tabulate encrust-

brachiopod shell,

mation,

Liberty For-

Preble County,

Ohio, x 1.7.

H. Octago-

nal tool house built ca. 1900 in John Paul Park, 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

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

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

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-

smooth, worn

sion

e p i t h e c a of Grewingkia.

of the

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 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 int 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

the vicinity. Mid-America Society.

Colonial Corals

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

individual polyps) ( F i g u r e s 6.6, 6 . 7 B ) .

of colo-

coral beds

Paleontology

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 c a l l e d the T a b u l a t a .

entirely

77

Figure 6.7. rugosan, (Foerste),

Cincinnatian colonial corals.

Richmondian,

Liberty Formation,

MUGM 5301,

Nelson County,

a

Kentucky, 78

tabulate in

A. Nelson

Cyathophylloides stellata County,

vertical section,

x 0.8. A Sea without Fish

Kentucky,

(Hall),

x 3.

showing tabulae,

MUGM 5285, B.

a colonial

Foerstephyllum vacuum

Richmondian,

Liberty Formation,

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 , Libertv, 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 distances 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; genera of massive c o l o n i a l

tabulates

Hatfield

and Tetradium) and o n e c o l o n i a l r u g o s a n these b e d s ( F i g u r e s 6 . 6 A - F ) .

1968; E l i a s 1998). F o u r

(Foerstephyllum,

Calapoecia,

(Cyathophylloides)

Nyctopora,

are f o u n d in

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 stonewalls 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 rep 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

T h e s e m o u n d s o c c u r in the

of t r e p o s t o m e b r y o z o a n s Maysvillian

Grant

Lake

(Cuffey

1998).

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 subtropical 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, sity of Cincinnati tions,

Maysvillian,

ryville Formation, County, Ohio, x

UnivercollecCorButler 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 certain 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 duri 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). Present-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 foursided, 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 specim 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 often 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 ) . 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

Figure 7.1. ments

A. Frag-

of bryozoan

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

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.

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

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 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 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 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 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

Parramosa

(d'Orbigny),

CMC IP

27957,

Bellevue

stone,

Cincinnati,

LimeOhio.

Scale in mm.

B.

of bryozoan

colony

showing

minute

ings (zooecia) on

aperture serving as both m o u t h and anus.

and

a

Surface 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

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

colo-

the most abun-

the left

cross-section

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.

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

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,

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 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 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

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 -

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." 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 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 ,

Each of the to a

individual animal. tome

bryozoan,

ticulipora

CMC IP 51107,

Bellevue

Limestone.

cinnati,

Ohio.

individual

(zooecia)

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 soc 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 parl 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

TreposMon-

mammulata

d'Orbigny,

of

tube

Cin-

Diameter openings

about 0.2

mm.

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

in

feeding position

(left)

and the other partly retracted (right). by Kevina

Drawing

Vulinec.

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 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 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 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 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 exa 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.

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.

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

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

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 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

Cincinnatian A.

Colonies

bryozoan

Spatiopora

genus

characteristi-

cally form a thin coating on

shells

of orthoconic

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,

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

Cincinna-

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

are

elongated

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 . 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 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-

(monticules)

aligned with

and

the axis of B,

host nautiloid shell. 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,

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.

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

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

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. 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 ( 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 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

Ropalonaria B.

Rafinesquina,

Ohio,

cates sac zooid. Scanning graph

C.

electron

cavities zooids

excavated into

BMNH D.52264,

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

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 . 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

by

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

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

micro-

of polyester cast of

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 -

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

CMC

x 9. Arrow indi-

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

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

Col-

shell of brachio-

from

(1978, plate

x 22.

1,

figures 5,

7) and reprinted by permission

of the

Paleonto-

logical

Research

Institu-

tion.

D.

Cystoporid

bryozoan,

Constellaria

The surface of the c o l o n y m a y be s m o o t h .

florida

Ulrich,

Many, however, bear regularly spaced b u m p s , termed monticules, or regu-

51108,

Fairview Forma-

a t r e m e n d o u s variety i n details.

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 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 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 pattern.

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

tion,

Inset,

showing

monticules,

scale in mm.

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.,

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,

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,

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 .

CrypEsch-

CMC IP

Fairview FormaKen-

tucky.

Escharopora

zooecia

on both sides of

the

thin,

enlargement

showing zooecia, mm.

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

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 -

87

F.

has

bladelike zo-

Inset,

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

Bryozoans

E.

bryozoan,

Boone Co.,

arium.

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

Ken-

enlargement

characteristic

star-shaped tostome

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

CMC IP

Kenton Co.,

tucky.

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).

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

B

Pohowsky

scale

in

Basal surface of bryozoan

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

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 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 present-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 s h a p e , at least in s o m e taxa.

chiopod

Rafinesquina

and overgrown it,

Rich-

ard Arnold Davis

collec-

tion,

Bellevue

Cincinnati, mm.

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

G.

bryozoan

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

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

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 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 , 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

Limestone,

Ohio,

scale in

Trepostome

colony

on

that has

shell of pelecy-

Ambonychia

overgrown it, Arnold

Davis

Cincinnatian, locality

and

Richard collection, horizon

unknown,

and

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.

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

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

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-

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 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 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 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

dian,

donut")

I.

encrusting

tiloid,

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,

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

h e n c e , s e e m t o represent areas o f the

mondian,

i n w h i c h the z o o i d s p e r f o r m e d

bryozoan

("Weichold

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-

colon

Ohio,

Cross-section

ring-shaped

zoarium nut")

("We-

presum-

Hamilton Co.,

x 0.7. of

Ring-shaped

H.

zoarium

do-

nau-

Richard Arnold Upper Rich-

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 . 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-

elliptical zone on

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

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

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-socarefully 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

right

and left sides of ring are

89

replacement shell.

of

Figure 7.4. Large bryozoan colonies from the type-Cincinnatian. tome

bryozoan,

Parvohallopora

ber of Grant Lake Limestone,

Hamilton Co.,

branching

trepostome bryozoan.

mm.

C.

logued,

nati collections, d'Orbigny,

into bryozoan.

disappeared, E.

A. B.

Cincinnatian,

Scale in mm. Monticulipora

Intact colony of a branching trepos-

of Cincinnati

Monticulipora

Corryville Member of Grant Lake Limestone,

CMC IP,

cm in width.

CMC IP uncatalogued,

that has

University

Ohio. Scale in mm.

no horizon or locality data.

nautiloid shell

Note borings

d'Orbigny,

Intact colony of trepostome bryozoan,

Cincinnatian,

probable

ramosa

no horizon D.

d'Orbigny,

d'Orbigny,

Co.,

Ohio.

Intact colony of trepostome bryozoan,

A Sea without Fish

Mem-

Scale in

CMC IP uncata-

Trepostome bryozoan encrusted on

mammulata

Kenton

This colony was excavated and reassembled by Ron Fine.

90

Corryville

or locality data.

mammulata

Hamilton

Corryville Member of Grant Lake Limestone,

collections,

Intact colony of an unidentified

Co.,

University

of Cincin-

Scale 2 cm in length.

Heterotrypa Kentucky.

frondosa Colony about 65

See Cuffey and Fine (2005).

Figure 7.5.

Reconstruc-

tion of the lower part of the zoarium erotrypa

of Het-

sp.,

diameter

about 28 cm.

From

Waugh et al.

(2004).

open

structure

zoarium

would

exposure

of interior

ing surfaces flow,

to

tion

provide

provide

fronds stabiliza-

and attachment

the substratum. Gamma

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 several 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 fragm 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

to

Re-

printed by permission Sigma

feed-

water

while the down-

ward-arching would

The

of the

of

Epsilon.

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 respiratory 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 fragm 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 fragm 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 stabil 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 incorp 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 diff 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 genus 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 , therew 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 creatures. 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 Catellocaula. 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 (2—2 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 recrystall 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 rings 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 cove 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 longest 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 alm 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 relations 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. studies

A.

Detailed

of bryozoans

quire

carefully

thin-sections peels.

or

acetate-

Diagram

orientation and

shows

of sections

terminology

nal

skeletal

used

to

of inter-

structures

identify species.

From Arens (1989,

re-

oriented

and

Cuffey

figure 5),

repro-

duced by courtesy of Roger J. mission

Cuffey, of the

with perPennsylva-

nia Academy of Science.

B.

longitudinal of

Left, thin

Heterotrypa

dosa

(d'Orbigny),

40336,

Bellevue

stone,

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R. J. Singh Collection. Right,

tangential thin

section

of same.

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

Both

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 coati 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 epiz 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 useless, 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 app 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 stand 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 essential 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 ) reported 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

( 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

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 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 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 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-

and D,

Platystrophia

ponde-

rosa

showing

fold.

Drawings from

Meek the

(1873), Ohio

Natural

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

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 maybe 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

of

sulcus

and

courtesy of

Department of

Resources Geological

Divi-

Figure 8.2. section

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

articulate B.

brachiopod. views

A.

of an

of pedicle

Interior valve

(left) and brachial valve (right)

of the

orthid

brachiopod

bertella. Kevina

Cincinnatian

Drawings

He-

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). 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 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 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 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 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 -

by

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

Vulinec.

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 projections 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 soc 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 inart 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 , respire, 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 involves 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 traditionalh 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 articulate 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 inf 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 relat 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 , however, 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 extension 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.

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., 51994,

a case in p o i n t . 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 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 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 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

A.

Pseudo-

CMC IP

Cincinnatian,

ho-

rizon and locality unknown,

x

1.9.

Specimen

oriented

perpendicular

bedding,

presumably

life position

with

to in

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.

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

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,

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 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 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 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 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).

B.

Trema-

millepunctata

Hall,

horizon and locality

rocrania

C.

scabiosa

encrusting

on

Pet(Hall)

Heber-

tellasp.,

MUGM

29461,

Arnheim,

Oxford,

Ohio,

x

D.

1.3.

scabiosa

Petrocrania

(Hall)

encrust-

ing on brachial valve of

Articulates

Rafinesquina

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

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,

( 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,

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 are beastly difficult to tell apart. T h e p r o b l e m is that brachiopods that are 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 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

sp.,

Charlotte

Bruce

Gibson

no.

1042, Franklin

Co., Indiana, approx. 1.5. lia two on

x

E.

Philhedra lae-

(Hall),

MUGM 26219,

specimens

encrusted

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,

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 F.

Lib-

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.

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

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.

Rafinesquina

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 ,

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 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 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

Ohio, x 1.8.

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

pedicle

valve exterior, Formation,

105

Hall,

Corryville

Warren

Co.,

Figure 8.4.

A,

B.

Plectorthis neglecta

Co., Ohio.

A.

Brachial valve exterior.

erata (Hall),

MUGM 24362,

Kope Formation,

valve exterior, both x 2.4.

E,

F.

nati, Ohio. E.

Pedicle valve exterior.

byella

Kope Formation.

rugosa,

106

(James),

B.

MUGM uncatalogued.

Pedicle valve exterior, both x Hamilton Co.,

Ohio.

Sowerbyella rugosa (Meek), F.

Fairview Formation, 1.8. C, D.

C. Pedicle valve exterior. D. MUGM 24564,

Brachial valve exterior, both x 2.2. G.

Courtesy of Paul E.

A Sea without Fish

Potter.

Hamilton

Dalmanella emac-

Kope Formation,

Brachial Cincin-

Slab covered with Sower-

Figure 8.5.

A-D. A.

Indiana.

Hebertella occidentalis (Hall),

scars and triangular pedicle opening. ing, all x tion,

1.2.

E.

Ohio.

E-H.

Brachial valve exterior.

Butler Co.,

Ohio.

x

D.

MUGM

G.

F.

11205,

Pedicle valve exterior.

Waynesville Formation, C.

MUGM 22450,

Pedicle valve exterior, H.

Liberty Formation,

Preble Co.,

x 2.8. G - H , MUGM 29458, Arnheim Forma-

Pedicle valve interior, showing muscle scars, x 2.2.

Franklin Co.,

Pedicle valve interior, showing muscle

Posterior view of articulated valves and triangular pedicle open-

Glyptorthis insculpta (Hall); E-F,

encrusting cyclostome bryozoans, processes,

B.

Brachial valve exterior.

Brachial valve interior,

triangular pedicle opening,

and

showing muscle scars and cardinal

1.8.

Brachiopods

107

Figure 8.6. Ohio.

A-C.

A.

valve exterior, (Hall),

x.2.1.

C.

F.

(Hall),

MUGM 22933 Liberty Formation,

Pedicle valve interior, showing muscle scars,

MUGM 23037, Arnheim Formation,

Brachial valve exterior. area.

Plaesiomys subquadrata

Brachial valve exterior, inarticulate brachiopod Philhedra laelia on beak,

E.

Butler Co.,

Pedicle valve exterior,

Ohio.

A Sea without Fish

D-F.

B.

Pedicle

Retrorsirostra carleyi

D.

note triangular pedicle opening and flanking inter-

Pedicle valve interior, showing muscle scars,

108

x 2.4.

Preble Co.,

x 2.1.

x 2.

Figure

8.7.

Three

of the

many described species

of Platystrophia.

ste, MUGM 24060, Maysvillian, Campbell Co., Kentucky valve exterior, 1.0.

D.

x

7.5.

C.

A.

robust brachiopod as

the

both x

7.5.

G,

rior.,

H.

Brachial valve exterior,

x

1.3.

x

ponderosa

7.5.

B.

E.

Foer-

Brachial x

Early collectors of Cincinnatian fossils referred

"double-headed Dutchman." E,

(Meek), MUGM 11315, Maysvillian, Cincinnati, Ohio. H

Platystrophia

Pedicle valve interior, showing pedicle opening and deep muscle scar,

Brachial valve interior, showing muscle scars,

to this large,

A-D.

Pedicle valve exterior,

F.

Platystrophia

Brachial valve exterior.

Platystrophia acutilirata (Conrad), MUGM 23360. Whitewater Formation, Preble Co.,

G.

F.

Pedicle valve exte-

Ohio, both x

Brachiopods

laticosta

Pedicle valve exterior,

1.4.

109

Figure 8.8.

The many faces of Rafinesquina:

nati collections, chial

valve

left,

up,

pedicle valve up,

with

encrusting

wedged beneath lower left edge, valve interior,

up,

showing

Boone Co.,

Fairview Formation,

to bedding,

alternata

edrioasteroid

Streptaster

Corryville Formation,

A.

University of Cincin-

Boone Co.,

Kentucky, C.

and

crinoid

Kenton

Fairview Formation,

size about same as in B. Co.,

Kentucky.

Kenton

Co.,

E. Kentucky.

A Sea without Fish

D.

scale in mm.

Rafinesquina pavement, with

top,

bra-

subcrassus

B.

Brachial valve interior,

Shingled Rafinesquina bed,

right,

locrinus

muscle scars and pair of cardinal processes along hinge at

Kentucky,

110

(Conrad).

vorticellatus

CMC IP 51111, showing muscle scars, scale in mm.

Cincinnati collections, Formation,

Rafinesquina

with encrusting edriasteroids and cyclostome bryozoans,

Pedicle

University of Corryville

with pedicle valves valves perpendicular

Figure 8.9. A, Ohio.

A.

x 2.2. C, Ohio. 3.4.

D. C.

E.

life position

B.

Hiscobeccus capax (Conrad),

Brachial valve exterior. Rhynchotrema

B.

Zygospira modesta (Say), to bryozoan

D.

Preble Co.,

Whitewater Formation,

Preble Co.,

Anterior view of articulated valves showing pronounced sulcus,

CMC IP 51112,

Parvohallopora sp.,

Jefferson Co.,

Liberty Formation,

dentatum (Hall), MUGM 25933,

Pedicle valve exterior.

Waynesville Formation,

MUGM 25490,

Posterior view of articulated valves showing small pedicle opening,

Indiana,

x

1.9.

Corryville Formation, F.

Boone Co.,

Catazyga schuchertana

brachial valve exterior,

x 3.4.

Brachiopods in

x

Kentucky, attached in

(Ulrich),

MUGM 7614,

Figure 8.10.

Environmen-

tal distribution chiopods in natian

of bra-

the

Series.

environments lent to

CincinShoreface

are

equiva-

the shallow sub-

tidal (1-2 m or 3-6 ft); transition ments

zone

are

environ-

deeper sub-

tidal (3-6 m or and

offshore

ments

are

with a

10-20 ft),

environ-

deeper

water,

maximum

depth

of about 30 m (100 ft). The heavy lines indicate the

environments

each

genus

where

is most abun-

dant and thin lines indicate

environments

where

a genus is present at lower abundance. Holland (1997), ontological Copyright

Events. 1997

University Press. printed of

From in Pale-

with

Columbia Re-

permission

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 rable m u s c l e scars.

the publisher.

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 (anatomically 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 considerable 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 , usi 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 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 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

Brachiopods

in

Time and Space

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 Ectenocrimis, 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 Platystrophia 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 plications (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 vario 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 prod 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 edrioasteroid 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 feature 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 pavem 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) suggested 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 presentday 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 characteristic, 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 differe 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

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 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 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 .

Figure 9.1.

Gastropod

mollusc,

showing

features.

Drawing by

Kevina

(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 individual 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

Vulinec.

internal

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 extent. 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 representative 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 largest 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 tt h h ee Molluscs Molluscs

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 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 followi 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 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

Class G a s t r o p o d a — Class T T hhee Snails 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 relatives, 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 monoplacophorans bellerophontid pod.

a

oran,

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

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

Archinacella

Wahlman,

40615,

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 a

gastro-

A.

area

and

USNM

monoplacoph-

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 Cyclonema; 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.

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,

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

plate 3,

figures 7, 9, 10).

B. Hel-

cionopsis

Ulrich

striata

and Scofield, 45827,

USNM

a

oran,

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 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

monoplacoph-

Richmondian,

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

Mar-

ion Co., Kentucky, a, dorsal, b, oblique-lateral, c, lateral, all x 1.7. From

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 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 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 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

plate 2,

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

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

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

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 -

Wahlman (1992, ornatus

USNM 265906, ite mold of a cophoran, mation,

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

Corryville ForCincinnati,

Ohio,

a, lateral, b, dorsal, x 1.7. From

Wahlman

(1992,

D.

Cyrtolites orna-

tus,

MUGM

18120,

individuals served by

two

with

bryozoan

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

Cincinna-

tian. x 1.7. cancellatus

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

Leptotrypa

clavacoidea.

E.

Sinuites

(Hall),

CMC

IP 44304, a calcitic specimen

of a

oran.

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 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

opposing each

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.

pre-

somehow

apertures,

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 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, 7).

( 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 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

monoplacoph-

Cincinnatian,

Cincin-

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 Form 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 (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

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 , 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 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,

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

x

1.5. From

Wahlman (1992, plate figures 7, 9). gostoma Ulrich,

F.

richmondensis USNM 45983, an

internal mold of a phontid

bellero-

gastropod.

Whitewater

From Wahlman,

x

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 tional e n v i r o n m e n t . 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 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 and crinoids is o n e of the best-known cases of interaction between species 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-

Formation,

Richmond, Indiana, plate 26,

11,

Salpin-

1.4.

clonema a n d

Naticonema are f o u n d a t t a c h e d

u s u a l l y of Glyptocrinus a n d Pycnocrinus

1992,

(see

to the t e g m e n s of c r i n o i d s , c h a p t e r 12). B e c a u s e , in m o s t

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

figure 2.

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.

varicosum a t t a c h e d to Pycnocrinus are k n o w n

gastropods.

from

the L e x i n g t o n L i m e -

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 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 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 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 -

A.

clonema Hall,

Cyn-

Formation,

Lexing-

ton, Kentucky, x 1.7. Cyclonema

ism, individuals of the associated species gain s o m e advantage w h i l e the

sum, lection,

Steven

ture.

Formation,

of

Ohio, x 1.2.

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 , 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 tion provided by the c r i n o i d .

Felton

showing

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 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

B.

humero-

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 Cornulites

Cy-

varicosum

CMC IP 51118,

thiana

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

(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

Cincinnatian

col-

aper-

Grant Lake Brown C.

clonema Ulrich,

Co., Cy-

humerosum CMC IP 51117,

Maysvillian,

Cincinnati,

Ohio, x 1.6. clonema

D.

bilix

Cy-

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),

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

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,

1.7.

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 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 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

CMC IP 51116, ArnFormation, E.

sublaeve 51114, 2.1.

Ulrich,

Cincinnati, F.

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

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,

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,

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 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-

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 had a s i m i l a r r a n g e of f e e d i n g habits.

striatulum

CMC IP 51115,

1.3. moorei MUGM

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

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 -

Kentucky,

encrusted

tabulate coral Prorichmondensis, Formation, x

Note slit in aperture.

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 resulting 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 attracted 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

123

x

(Safford),

Preble Co., Ohio,

M a n y present-day predatory snails attack their prey by " d r i l l i n g "

Molluscs

internal Formation,

23292,

Whitewater

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 .

Subu-

Ulrich,

Paupospira

taraea

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 fossils.

Kope H.

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

tory, it is possible that the snails of Subulites (Fusispira) w e r e p r e d a t o r s .

G.

(Fusispira)

mold,

Kope

Cincinnati,

Ohio, x 3.2. lites

x

Ul-

CMC IP 51113,

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

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 -

Ohio,

Cyclonema

Kenton Co.,

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

CMC IP

Fairview Forma-

tion,

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-

x

1.5.

Figure 9.4.

Gastropod-

rich beds. itown

A.

Shale,

Fossil Park, Ohio.

Sharonville,

Note shells

geopetal part

MiamTrammel

cavities

filled

with

upper part

filled

calcite crystals). ble Hill Bed, Formation, Kentucky.

with (lower sediment, with B.

Mar-

Waynesville Trimble Co.,

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 similarities 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 interpretat 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 interior 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

9.5.

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

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

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

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 -

Kevina

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 thickness 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

Pelecypod

Vulinec.

internal

Figure 9.6.

Cincinnatian pelecypods.

locality information,

x 0.6.

mation, Butler Co., Ohio, mation,

Clermont Co.,

right valve, MUGM

right valve, 15066,

A.

x

Ambonychia sp.,

C. 1.1.

right valve,

MUGM 29546,

Cincinnatian,

right valve,

Carotidens demissa (Conrad), MUGM 29431, left valve, D.

Anomalodonta gigantea Miller,

Warren Co.,

Whitewater Formation,

internal mold of right valve,

126

MUGM De549A,

Ambonychia cultrata (Ulrich),

x 0.8.

Ohio,

Waynesville Formation,

MUGM 23392, rich,

B.

A Sea without Fish

Ohio,

x 0.8.

E.

Preble Co.,

Ohio,

CMC IP 35898,

no

Waynesville ForCorryville Forinternal mold of

Opisthoptera casei (Meek and Worthen), x 0.9.

Whitewater Formation,

F.

Butler Co.,

Ischyrodonta truncata UlOhio,

x

1.1.

Figure 9.7.

Cincinnatian pelecypods and rostroconchs.

USNM 46230, tion,

"butterflied" specimen

Covington,

(Conrad),

Kentucky,

USNM 46719,

Fairview Formation, rus faberi Miller, right-lateral view,

black

A.

Modiolopsis sp.

organic film

x 0.85. From Pojeta (1971, plate

USNM 07219, 11,

Ohio,

cf.

15,

figure 6). to

B.

figure 3).

Whitewater Formation,

Kope Formation,

D. Butler Co.,

Covington,

figure5).

Kentucky,

C. x 3.

Technophorus milleri Pojeta and Runnegar, Ohio,

x 5.1.

TechnophoFrom Pojeta

MUGM 6848,

From Pojeta and Runnegar (1976,

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 Paupospira

Fairview Forma-

Modiolopsis modiolaris

14, figure 7).

m e n s of three s p e c i e s of g a s t r o p o d s ,

modiolaris (Conrad),

bedding and highly foreshortened,

x 2.4. From Pojeta et al. (1986, plate 17,

left-lateral view,

M.

and remnants of ligament,

internal mold preserved at right angles

Cincinnati,

and Runnegar (1976, plate

with

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

plate

Figure 9.8. pied by

A.

Basal side of stromatoporoid with borings made by pelecypods,

the pelecypod

Brown Co.,

Ohio,

x 0.6.

inarticulate brachiopod, Clinton Co.,

Corallidomus scobina,

Ohio,

B.

OSU 8420,

Corallidomus scobina,

Trematis sp.,

in

life position

x 0.7. From Pojeta (1971, plate

16,

probably

USNM 70458,

among

valve

with

Clermont Co.,

edrioasteroid

Ohio,

x

Isorophus

cincinnatiensis

1.4. From Pojeta (1971, plate 10,

128

A Sea without Fish

bryozoan

figure 5).

left valve of specimen attached to bryozoan colony shown in B, left

from

and

C.

x 2.8. encrusting

the

some of which are occu-

Waynesville

Formation,

bryozoan colony with pelecypods and branches,

Waynesville

Formation,

Corallidomus scobina, USNM 70458, D.

Carotidens sp.,

bryozoans,

Corryville

USNM

162734,

Formation,

figure 15). A and C, courtesy of John Pojeta, Jr.

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

ford, i n

pods.

From Pojeta

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

(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

pelecy(1971,

figure 9).

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 arenot 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 differentiate 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 notu 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 buried 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 pies 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 , siltstones, 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 Modiolopsis 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 substrata 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 hing ( 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 singularly 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 externally. 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 nautiloids

attained

happi-

ness. In the top diagram, the center of gravity,

G,

is inferred to lie in the body

chamber

nautiloid,

of

and

the

the center

of buoyancy, 6, to lie in the

gas-filled

phragmo-

forcing

the animal

cone,

into a vertical,

head-

down position.

In this

position

it could not

swim efficiently.

In the

middle diagram, mation

the for-

of calcareous

cameral deposits in the apical part of the phragmocone the

should

result

in

center of gravity and

center ing

of

buoyancy

toward

the

shift-

midpoint

of the animal's length B ), a

(G

2

permitting stability in

2

horizontal

orientation.

In the lower diagram,

the

nautiloid has grown, additional posits

cameral

centers

(G ,B ) 3

tain

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.

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.

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

Flower (1957,

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

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.

133

See

text for

of the

Society

From

figures

and reprinted

America.

h i g h e r taxon w i t h o u t d a m a g i n g it.

Molluscs

a stable horizontal

permission

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

main-

3

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

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 .

to

so that the

of gravity and

buoyancy

4-6)

de-

formed closer

the head,

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

with

of

by Geo-

Figure 9 . 1 1 .

Cincinnatian

orthoconic A-C

nautiloids.

from Davis and

Mapes

(1996),

of the

Ohio

courtesy Department

of Natural Resources sion

B.

Treptoceras

duseri (Hall and field),

bars = 1 cm. 47422,

of

A.

OSU

phragmocone

and living

with dark lines

which

are remains of what, life,

in

aequabile

nal mold of portion siphuncle,

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.

OSU 47420,

1 cm.

Endoceras

D.

MUGM 29579,

inter-

nal mold of part of phragmocone tion,

and

sip-

scale in mm. byoceras

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 an o r t h o c o n i c

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 " :

c u l a r deposits).

CMC IP 31393, fine

longitudinal lines, water Formation,

Whitex

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

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 -

Gor-

with

cephalopod

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

duncanae

external shell

definitely do not

deposits);

Ohio,

E.

they

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

Liberty Forma-

Butler Co.,

Flower,

tion, for short. S o m e paleontologists and other biologists speak of evolutionary

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 .

scale bar =

huncle,

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-

inter-

Cincinnatian, sp.,

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

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

in-

(Miller),

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

"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

were color bands.

C. Cameroceras

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 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

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-

part of

external shell, longitudinal

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

enjoyable?) that direction m i g h t prove to be. Rather, it a particular direction

chamber.

B. OSU 47417,

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

scale

internal mold of

part

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

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 .

Whit-

Cincinnatian,

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 g r e a t e r or less e x t e n t .

Divi-

of Geological Sur-

vey. A,

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 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

1.4.

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

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 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 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

c u r v e d shells) Manitoulinoceras tenuiseptum a n d M.

nautiloids.

Oncoceras delicatum

(4 p e r c e n t ) , Zittelloceras russelli,

williamsae (8 p e r c e n t ) , and

Z.

williamsae (<1

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) tentatively 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-

(Richmondian)

toceras

A.

Charac-

baeri

Worthen),

(Meek

MUGM

and

29591,

Whitewater

Formation

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

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.

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

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 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 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 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 infilled w i t h c a l c i t e crystals.

These preservational features led Frey to c o n -

Nautilus, B.

CMC IP 24415, mold

of phragmocone

water Formation, Co., Ohio, x 1. estoceras

eos

Whitfield),

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

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

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 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 .

Di-

(Hall and 382,

of partial and

body

Whitewater

Formation,

Preble

Ohio, scale in mm.

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 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 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 (Felton, 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 Cameroceras, 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

Butler C.

MUGM

mold

of

White-

chamber,

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 ,

the top predators.

(Miller), internal

a cyrtoconic form,

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

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 -

Beloito-

amoenum

137

Co.,

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 scrutiny. 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 portions 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 (chitons), 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 scap 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.

richmondensis (Miller),

CMC IP

17551,

A.

Waynesville Formation,

Ohio. Slab showing parallel alignment of shells. Scale in mm. worm,

Protoscolex ornatus

ington,

Kentucky.

vation,

x 7.5.

crinoid

locrinus

142

CMC IP 37990,

C.

Co.,

Annelid

Kope Formation,

Cov-

Tubes of Cornulites sp. attached to the column of the

subcrassus,

Hamilton Co.,

coleus sp.

cf.

University

of Cincinnati collections,

Ohio. Scale in mm.

L. jamesi (Hall and Whitfield),

Corryville Formation,

A Sea without Fish

Clinton B.

This is a rare case in the Cincinnatian of soft-body preser-

Formation, tions,

Ulrich,

Tentaculites

Boone Co.,

D.

Corryville

The machaeridian Lepido-

University of Cincinnati collec-

Kentucky.

Scale in mm.

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 .

...

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

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

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

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 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 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 -

at

we can easily imagine the

ocean

beneath

Cincinnati deposited,

times swarmed with

innumerable

worms,

which have, so far as 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

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

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,

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

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

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 individual 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 softb 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

traces of themexcepting tracks,

their

and pos-

few rude impres-

of their bodies.

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 (Figure 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 Form 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 Paleontology. 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 (Howell 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 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

Conical Fossils

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 striations. 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 , Gornulites 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, unatt 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 att 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

richmondensis,

the

type-Cincinnatian,

both from the

Tentaculites sterlingensis a n d T.

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 , taperi 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 att 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 interpreted 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 , trilobites, 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. Duri 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 , ostrac 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 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

Figure 11.1. The Ordovician

trilobite

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.

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

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 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 ind 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.

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 exoskeleton 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

Drawings by

Vulinec.

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

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 -

Triarthrus.

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 furrows 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 tessellatus 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 segm 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 hyp 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 exceptional 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 pygidi 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 stratig 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 bites,

Ordovician

showing

structions ages.

of

the

mene

A. B.

Isotelus

(Con-

Composite of

maximus

latus,

ted because

tessellatus Raymond

figures 9,

16, 20) and by

permission

Connecticut Acad-

emy of Arts and

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 distributed 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 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 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

Green. (1920,

reprinted

Sciences.

149

un-

Cryptoli-

thus

the

I.

omit-

they are C.

From

of

trilobites with

and

(exopodites

known).

top

Flexicaly-

senaria

rad).

Waynesville

append-

Anterior at the

in each.

Cincinnatian

trilo-

recon-

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 Foerste's C. meeki-retrorsa. Ross c o n s i d e r e d F. retrorsa to be a valid species alt h o u g h it has little to d i s t i n g u i s h it

750

A Sea without Fish

from

meeki except the size, s h a p e , and

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 that only a single species, F. meeki, is valid, a n d that b o t h F. granulosa and 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. 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 of F. meeki, F. retrorsa, and F. granulosa as valid s p e c i e s .

(Foerste),

collections,

Maysvillian,

Corryville

Formation, Ohio, C.

Hamilton

enrolled

cephalic

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-

University of

Cincinnati

width

rorsa (Foerste),

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

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 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 butter 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

mation,

Arnheim

Mark Peter col-

Formation,

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,

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 shaleb 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 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 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 features 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 possiblebehaviors 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 discoveries 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

For-

Ohio, x 1.4. D. Flexicalymene granulosa (Foerste),

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

1980,

Rich-

Highland Co.,

lection,

m o v e m e n t s of hue-grained sediments (Brandt

ret-

CMC IP,

Collection,

mondian,

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 -

furth

28 mm.

Flexicalymene

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

Co.,

specimen,

151

Edenian, Brown

x 2.5.

Kope Co.,

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 Rusophycus 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.

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,

small, infaunal o r g a n i s m s (Fortey a n d O w e n s 1999). 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 Flexicalymene, 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

2250,

Riehmondian,

heim

Formation,

land Co., Ohio, B.

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, lian,

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

lymene.

Robert

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 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 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 the closely related F. senaria ( f i g u r e 1 1 . 2 A ; R a y m o n d 1920).

x

1.3.

Maysvil-

Clermont Co.,

7.

Ohio,

CMC IP 51,

C.

Nestor

Maysvillian,

collection,

Corryville

Formation,

Clermont

Co.,

Ohio, x 1.8.

D.

pendages

ventral side

of a

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 -

Arn-

High-

Large hypostome,

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 The study by H u g h e s and C o o p e r revealed pyritized material c o n -

En-

A.

CMC IP

on

complete

USNM

Ap-

specimen,

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

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 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 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

leborough courtesy 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 duri 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. gigas 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 , flattened 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 narrower 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

was

orig-

inally illustrated by Mick-

153

(1883). of Loren

Photo

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, riv a l e d o n l y by t h e less c o m m o n

Megulograptus and a l o n e , Isotelus m i g h t b e

euryptcrid

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

endocerid 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.

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

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 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 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 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 the

Isotelus

exoskeleton

and

is o f t e n

found

as a n

isolated c o m p o n e n t

( F i g u r e 11.4B).

(Hall),

A.

CMC IP 46429,

Edenian

(figured in Davis

[1992, plate 2, as

Proetus

B.

eatoni

collector,

Steve

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 m e n s often show

Ohio, x 4.6.

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

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 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 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

C.

tessellatus

Green,

University of Cin-

cinnati nian,

collections,

Hamilton Co.,

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 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 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 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 curTrace 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). Different 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 varyi 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 ) ,

155

Ede-

Kope Formation,

3.7.

Arthropods

Co.,

Cryp-

tolithus

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

long, comb-like filaments (Figure 11.2C).

Kope

Hamilton

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

the a p p e n d a g e s of Cryptolithus.

Rush

Edenian,

Formation,

rent.

Triar-

(Hall),

Brown collection, J.

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

Isotelus

figure 23] parviuscu-

lus), x 7.7. thrus

Deco-

parviusculus

Ohio,

x

Figure

11.6.

B,

D.

Ohio,

x 4.0.

A. D,

Steve Brown collection, J.

Undetermined.

B.

Steve Brown collection, J. Rush collector,

of Cincinnati collections, bryozoan

Acidaspis cincinnatiensis Meek,

Odontopleurid, gen. and sp. Kope Formation,

Peronopora sp.,

Hamilton Co.,

MUGM 4001-A,

x

2.8,

A Sea without Fish

C.

x 6.

F.

Rush collector,

x 2.6.

Richmondian, Oxford, Butler Co.,

Primaspis crosotus (Locke), E.

from Shrake (1989,

ventral side of trilobite showing hypostome in place, 156

x 2.6.

Ohio,

lata bioclaustrations in upper right part of bryozoan. 4010,

MUGM 29056,

University

Primaspis crosotus (Locke) on

plate 3C).

Note

Catellocaula

Primaspis crosotus (Locke) on bryozoan, x 5.2,

from Shrake (1989,

plate 4B).

valMUGM

Figure

11.7.

mondian, 29057,

Riehmondian,

and Gurley, D.

Cincinnatian phacopid trilobites.

Waynesville Formation,

Waynesville Formation,

CMC IP 5199,

A.

Montgomery Co.,

Platycoryphe christyi (Hall), Ohio,

Franklin Co.,

x 2.6. Indiana,

B.

Mark Peter collection,

Tricopelta breviceps (Hall),

x 5.5.

C.

Ceraurus milleranus Miller

enrolled, no unit or locality (figured in Davis [1992, plate 3,

Ceraurinus icarus (Billings),

Steve Brown collection, J.

Rush collector,

Riehmondian,

Arthropods

RiehMUGM

157

figure 28]), x 2.

x 2.5.

Figure 11.8. A,

C

printed by permission sis

from Caster and Kjellesvig-Waering (1964, plate 45, of the

Caster and Kjellesvig-Waering,

abdomen

of Megalograptus ohioensis. strabops

martini

Ohio,

Society

Richmondian,

and last (sixth) pair of legs

to right near top), holotype,

Co.,

Paleontological Research

for

x

Elkhorn

(darker segments),

CMC IP 24119A, C.

x 0.14.

holotype,

158

Geology.

A Sea without Fish

Adams

Co.,

Ohio;

Professor Kenneth E. CMC IP 24117A,

109,

Maysvillian,

figure 2),

figure 4) and re-

Eurypterid Megalograptus

dorsal imprint of prosoma B.

CMC IP 25569,

1.4. From Caster and Macke (1952, plate

figure 1, plate 46, A.

Formation,

First walking leg, paratype,

Caster and Macke,

Sedimentary

Institution.

ohioen-

ventral side of post-

(grey segments

turned

Caster with type specimens

x 0.9.

D. Aglaspid Neo-

Corryville Formation,

Clermont

and reprinted by permission of the

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 Hirnantian. 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 osculum 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 stromatoporoids. C. Scleractinian coral, Tubastrea coccinea Lesson, showing tentaclebearing 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, unidentified 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 annelid 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 coloration. 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,

Plate 8.

A. Flexicalymene retrorsa on internal mold

CMC IP 50168, Richmondian,

of nautiloid, CMC IP 50844, Arnheim Formation,

Adams Co., Ohio. This exceptional

Thomas Weaver Collection. Trilobite is located

complete specimen is 37.5 cm

on phragmocone (chambered) section of the

in length and w a s collected

nautiloid mold, with head toward body chamber.

and prepared by Thomas T.

Scale in mm. B. Parallel alignment of orthoconic

Johnson. Isotelus is the State

nautiloids, C M C IP, Cincinnatian. Scale in mm. Note

Invertebrate Fossil of Ohio.

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. Carpenter), 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, Washington, 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 location 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 Kenneth 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.

Kjellesvig-Waering (1964, plate 49,

figure 2, plate 48,

adult female (from Caster and Kjellesvig-Waering [1964,

figure

1).

plate 43]),

A. x 0.14.

surface of adult female (from Caster and Kjellesvig-Waering [1964, plate 44]), second walking leg, 0.7. x

D. 0.4.

note bulbous expansion

Third paired appendage, A-D

C, B.

(sensory?) at end of longest spine,

of the Paleontological Research

From Caster and

Reconstruction of ventral

x 0.14.

C.

paratype,

showing large coxa and part of first four joints,

reprinted by permission

D.

Reconstruction of dorsal surface of

paratype,

Institution.

Arthropods

Distal end of CMC IP 24115,

159

x

CMC IP 24103A.

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 signific 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 app 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 suggests 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 segm 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 11.10. Cincinnatian

Figure

ostracodes. atopsis

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) 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-

CMC IP 40171,

Formation,

Cincin-

nati, Ohio. From Warshauer (1972, plate 5), 30.

B.

33085,

Cincinnati, 33. regularis

CMC IP

Kope C.

Formation,

Ohio,

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 association w i t h a cluster of Flexicalymene ( H u g h e s a n d C o o p e r 1999).

x

Ctenobolbina

alata Ulrich,

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 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

Cer-

chambersi

(Miller), Kope

A.

x

Quadrijugator

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 debatea 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 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 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 posit 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 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

(Emmons),

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 (Figures 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 exoskeleton 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. ohioensis, 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 considered 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 KjellesvigW 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 ohioensis 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 reg 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, Megalograptus 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 having 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 evolutionary 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 disting 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 classification, 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) reported 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 dissolution 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 Arnh 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. a

The arm of

crinoid is composed of

a series of these elements,

connected

muscles

and

Comactinia bean.

sp.,

Carib-

Scanning

micrograph,

by

ligaments.

x

electron

79.

Figure 1 2 . 2 . Arm of modern with

crinoid

pinnules

branches) tube

bearing

feet in

ture.

Pinnule

about

1

photo,

(dark) (light fine

feeding poslength

cm. Aquarium comasterid

noid,

Curacao,

lands

Antilles

cri-

Nether-

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 .

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

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

puzzle

the

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

. here salute the echinoderms to

zoologist.

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 diversified 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 ) existed 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 fossils. 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 invertebrates 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 intricate 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). Skele 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 nurturi 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 respiratory 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 , tubefeet 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 skele 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 pres 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 sign 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 struct 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 persisted 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 C o m p l e t e fossil

echinoderms

are

1996).

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 requires 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 preservation 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 holdfast 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

locrinus

crinoids.

subcrassus

(Meek

and

Worthen).

A. Articulated

individual

with

partially disarticu-

lated

sections

and matrix lated

of stalk,

of disarticu-

skeletal

nents.

compo-

Upper

Ordovician,

Corryville Formation, cinnati, of

Cin-

Ohio.

University

Cincinnati

collections.

B. Two

articulated

crowns,

detached

stalk, but in

opposite

preserved on bed.

from

oriented parallel direction,

base

of

Upper Ordovician,

Corryville

Formation,

Clermont Co.,

Ohio.

CMC

IP 44362. Scale in mm. B in

upper right denotes

basal plate; radial

R denotes

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 beautiful noidea,

forms

cyclic) of basals or t w o circlets (dicyclic) of basals a n d infrabasals o c c u r r i n g

of Cri-

or stone-lilies .

.

C h a r l e s Lyell 1845, 50

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 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 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 situated 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 feedi 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 tion

12.4. of

Reconstruc-

Glyptocrinus

decadactylus

in

life

po-

sition. Calyx is in a horizontal position,

with

the

arms splayed into a filtration fan.

By analogy with

living crinoids,

current

flow was from left to right. Agnew.

Echinoderms

171

Drawing by John

Figure 12.5. Cincinnatian crinoid columnals and holdfasts. A,

H.

naticrinus

Cincin-

varibrachialus

Warn and Strimple;

A.

Articular surface x 9.6; H. Lateral view of mature section x 5.9. B, enocrinus

I. Ect-

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, Glyptocrinus lus Hall; D, surfaces

K. decadacty-

G. Articular

of internodal,

nodal respectively, 4.4;

x

K. Lateral view x 6;

note three cycles ofinternodals.

E.

Merocrinus

curtus Ulrich, articular surface x 5.2. alocrinus

F.

Anom-

incurvus

(Meek

and Worthen), lateral view of

compressed

with

section

fracture separating

meres, x 1. A - K , from Meyer et al. (2002, figure 2) and reprinted by permission of The Paleontological Society. nus

I,

M. Lichenocri-

crateriformis

Hall,

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

FMNH8810. L.

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. From Faber (1929, plate 32, figures 5, 6) and reprinted by permission of the American uralist.

Midland N.

milleri Faber,

Nat-

Lichenocrinus

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 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). 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 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 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 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 -

CMC IP

10047, Elkhorn Formation,

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

cross-section

that are less c o m m o n .

showing

floor

plate with central node,

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.

crater plates, and base of column, y.3.7, from Faber (1929, plate 33, figure 9).

172

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 A Sea without Fish

Figure A,

12.6. B.

(Ulrich),

Cincinnatian disparid crinoids.

YPM 24801,

Fairview Formation,

showing regeneration of arms, crinus geniculatus (Ulrich), E. simplex (Hall) plate 3, the

A,

Holotype, CMC IP 3871, Kope Formation,

figures

F. CMC IP, x 7.5. 1, 2, plate 6,

Paleontological

Research

Cincinnati,

USNM 93223,

CMC IP 36313, figure

B.

Ohio,

Ohio, x

Fairview Formation,

Kope Formation, G.

Cincinnaticrinus varibrachialus

Cincinnati,

x

1.3.

1.0 and x 3.1. D. Ohio,

C. pentagonus

C.

Dystactocrinus constrictus (Hall),

Hamilton Co.,

Cincinnati,

Warn andStrimple.

x

Ohio, 7.4.

x 2.0. F,

E.

Ecteno-

G.

CMC IP 42679, x 0.3. A - C , E, G from Warn and Strimple (1977,

1, plate 13,

figure 3, plate!4,

figure 8) and reprinted by permission of

Institution.

Echinoderms

173

Figure

12.7.

Cincinnatian disparid crinoids.

horizon and locality unknown, Formation,

Cold Spring,

logued, Cincinnati, OH,

x

1.4.

Kentucky, x

174

1.4.

x D.

B-D. 1.8.

A.

locrinus subcrassus (Meek and Worthen),

Anomalocrinus. C.

B.

A. sp,

Arms of A. incurvus Meek and Worthen,

A. sp, CMC IP 7341, Cincinnati, Ohio.

A Sea without Fish

CMC IP 170,

MUGM 28048,

Bellevue-Corryville CMC IP,

uncata-

Figure 12.8. monobathrid

camerate

crinoids. nus

Cincinnatian

A.

Glyptocri-

decadactylus

MUGM

28046,

Formation

(?),

locality

unknown, x 1.5. Pycnocrinus (Meek), Corryville Morrow,

175

B.

dyeri

USNM 40762, Formation, Warren

Ohio, x 1.2.

Echinoderms

Hall, Fairview

Co.,

Figure 28121,

12.9.

Cincinnatian

Liberty Formation,

Cyclonema sp.,

monobathrid camerate crinoids. Butler Co.,

MUGM uncatalogued,

Wayne State University Collection, eter.

Note

Figure tion,

12.10.

Formation, S9I,

A.

A.

B.

Arnheim Formation,

Dent,

Dent,

of articulated crowns,

x

7.2.

C.

east of Lebanon,

Warren Co.,

A Sea without Fish

Ohio, Ohio,

No. 366,

MUGM

Ohio,

x

1.3.

x 2.0.

C.

P.

dyeri,

lens cap 55 mm diamdown.

Bruce and Charlotte Gibson Collec-

Xenocrinus baeri (Meek),

Diplobathrid camerate crinoid,

figure 3a).

176

Co.,

some splayed oralside

Merocrinus curtus Ulrich, B.

Hamilton Co.,

Hamilton

Ohio,

x 4.2.

Glyptocrinus fornshelli Miller,

Pycnocrinus dyeri (Meek) with attached gastropod,

Cladid crinoid,

southwestern Ohio, plate 2,

7.7.

Hamilton Co.,

Waynesville Formation,

Meek (1873,

x

Arnheim Formation,

remarkable preservation

Kope Formation,

Ohio,

MUGM 28347,

Liberty

Gaurocrinus nealli (Hall) USNM

This specimen was illustrated by

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.

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

A, B.

padocystis

Le-

moorei

(Meek),

CMC IP 24680,

Elkhorn

Formation,

ble Co.,

Ohio,

C.

fulton-

Sumrall and Schu-

macher,

holotype,

IP 50402, tion,

x 4.3.

Cheirocystis

ensis

Pre-

CMC

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 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 Cincinnatian

c r i n o i d s , Cincinnaticrinus,

Ectenocrimts,

and

locrinus are dis-

parids ( F i g u r e s 12.6,12.7; Plate 10). Cincinnaticrinus a n d probably Ectenocrimis 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 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

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

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. struction

D.

Recon-

of late

Riehmondian

sea

southeastern

Indiana

southwestern showing moorei

(A)

attached

(B),

to

brachiomod-

and an edrioast-

Carneyella sp.

(D).

A, B, D from Kesling and Mintz (1961, plate 6, ures 8, reprinted

of the Museum ontology,

fig-

10, plate 7) and by

served fossils are f o u n d that reveal the entire a n i m a l . Cincinnaticrinus

Ohio,

Zygospira

esta (C), eroid,

and

Lepadocystis

bryozoans pods,

floor of

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 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 preand

Ectenocrinus

are

frequently

found

together

in

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 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 tog 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 , 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 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. 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 is larger a n d m o r e robust in s t r u c t u r e than (lincinnaticrinus and Ectenocrinus ( 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-

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 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

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). least 10 cm h i g h a n d had up to 500 b r a n c h e s .

The c r o w n was at

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. Edrioasteroid

Isorophus

tiensis

cincinna-

(Roemer),

recon-

structed as in life, food grooves feeding.

with

open

for

Drawing by John

Agnew.

Figure

12.13. Cincinnatian

edrioasteroids

A,

B.

cincinna-

Isorophus

tiensis A.

(Roemer).

Corryville Formation,

Hamilton Co., versity

Ohio,

lection, x 2.2. individual against

B.

smaller

individual,

of

Carneyella

C-E.

E. At-

crinoid column,

CMC IP 26324, F.

Streptaster

vorticel-

CMC IP

24700, x3.2.

G.

stellatus

CMC IP 40481, Photos C - G

(Hall),

13,

10,

figure

figure14, plate

figure 7) and re-

printed by permission Albany,

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 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). 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 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

of

the New York State Museum,

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 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

10, plate 17, figures 1, 3, plate

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 ,

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

x 6.2. 16,

( 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

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 Cys-

from Bell

(1976a, plate

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 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

x 2.1.

latus (Hall), taster

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-

C. CMC IP to

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 Form 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

pilea

34537, x 3.2. D, tached

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

Cincinnati

collection, x 3.

pattern of ridges on t h e c a l y x plates. Glyptocrinus decadactylus o c c u r s in the 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 -

Kentucky,

University

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 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

Formation,

Co.,

(Hall).

Large

crowding

Corryville Boone

Uni-

of Cincinnati col-

N.Y.,

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 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 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

asteroids. A,

B.

mopalaeaster

Prospecio-

sus (Meek).

A.

Associations of Crinoids with O t h e r Species 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 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 -

Oral

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,

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 -

side.

B.

108059, x 2. mopalaeaster e s (Miller) side. holotype,

magnifiC.

D.

MCI

C, D. ProOral

Richmondian,

40883,

Waynes-

ville, Warren Co., Ohio, x 0.8.

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 earliestk 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,

Aboral side, USNM

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

Photos courtesy of

Jon W. Branstrator.

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 subs 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 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). 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. 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 discoveries 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.

side.

B.

Cincinnatian asteroids.

Oral side,

(Miller and Dyer),

holotype,

(Meek),

MUGM 29664,

mation,

Brookville,

Waynesville,

A,

B.

Lanthanaster intermedius (Schuchert).

holotype, FMNH 9575, Maysvillian, MCZ 108063,

with arms

Franklin

Warrern Co.,

184

Co.,

Ohio,

Maysvillian,

Cincinnati,

Preble Co.,

wrapped around bivalve

Indiana.

E.

x 3. Photos A,

A Sea without Fish

Ohio, Ohio,

A.

Aboral

x 3.

C.

Petraster speciosus

7.

D.

Promopalaeaster dyeri

x

in presumed feeding posture,

Salteraster grandis (Meek),

USNM 40885,

B courtesy of Jon W. Branstrator.

Waynesville ForRichmondian,

Figure

12.16.

Waynesville,

A.

Asteroid,

Warren Co.,

Ohio,

cian, southern Pennsylvania, Boardman

et

Publishing;

C,

al.

(1987,

x

figure

Hudsonaster simplex (Miller and Dyer), x 3.1. 1.4.

B. C.

18.45B),

USNM 40882,

Taeniaster spinosus (Billings),

Cyclocystoid,

USNM,

Richmondian,

Zygocycloides magnus (Miller and Dyer).

courtesy of James Sprinkle,

near

Middle or Upper OrdoviB,

from

and reprinted by permission of Blackwell

courtesy of Colin Sumrall. Echinoderms

185

Figure

12.17.

stylophoran A.

1-6.

1 -3.

sal view.

3.

view, x 2.3. type,

2.

CMC IP 25257,

ryville Formation, ton Co., Ohio. view. 6. 1-6

5.

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 . 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

Lateral 4-6.

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 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

Ohio.

Ventral view.

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) 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).

Formation,

Clermont Co., 1.

Holo-

CMC IP 25993,

Corryville

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 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

Enoploura p o -

p e / Caster. type,

Cincinnatian carpoids.

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

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

Hamil4. Dorsal

Lateral view.

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 , alt h o u g h the identity of the culprits remains uncertain.

Ventral view, x 2.3. from

plate 1).

Caster (1952, B.

tion of E. popei, Parsley

Rhombiferan "cystoids"

Reconstruc-

(1991,

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 -

from

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

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

Paleonto-

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 . Cheirocystis 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 fossils 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 intera 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 app 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 (Figu 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 lowprofile, 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 disa 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 species 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 brac 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 bestk n o w n C i n c i n n a t i a n sea star, Promopalaeaster, stratigraphic

section.

An

extraordinary

is f o u n d t h r o u g h o u t t h e

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 sedim 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 Taeniaster, 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 interradial 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 circ 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 favored 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 M a v s v i l l i a n ( F i g u r e 12.17)

a n c

'

t w o s p e c i e s , E. popei C a s t e r in

the

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 attached 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 interp 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 wellp 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 stvl 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 preserved 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 skeleton ( 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.

o r g a n i s m : narrow

lite

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

A.

morphology.

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.

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.

typicalis (Hall).

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

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 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 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 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

Sin-

courtesy

Goldman.

Cluster of partly paral-

lel-oriented MUGM 29469,

rhabdosomes, Cincinna-

tian, Butler Co., Ohio, 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 biostratig 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 .

B.

rhabdosome,

of Daniel C.

B,

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 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

GraptoDrawing

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

conodonts.

Those

are among common from

the most

forms.

the

dian

shown

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-

All are

lower Riehmon-

Stage near

Brookville,

Franklin

Indiana.

Co.,

A, G, H.

todina

tenuis

\ 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 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

Plec-

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 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 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 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

(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.

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

G.

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

(Branson,

Mehl,

Branson),

Pb

C,

D.

thus

and

Amorphogna-

son and Mehl. element.

BranC.

D.

ment. dus

E.

and Mehl). modus

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 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

(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 -

Phrag-

undatus

Branson

S element.

I

Rhodesognathus

egans

(Rhodes),

ment.

J.

Pb

tron

Scanning

micrographs

tesy of Stig M. strom, 130.

apparatus, S

M

elements

series

are

elements,

ing position in

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).

pterobranchs that interconnects zooids within the colony.

The identifica-

tion of the protein c o l l a g e n in the tubes of both graptolites and pterox

the

elements

medial elements,

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-

elec-

a and b in

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 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

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-

Berg-

denoting position

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

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

cour-

approximately

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 -

Pb

P elements are prin-

cipal elements,

are

ele-

Branson,

Mehl and Branson, element.

el-

Aphelogna-

grandis

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 ans, and hemichordates, or were considered to be unrelated to any living

DrepanoistoF.

and Mehl,

Sc

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

Pa ele-

suberectus

thus

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

element.

ordovicicus

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 .

and

symmetry c denot-

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 chord 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. 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 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 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 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 purposes of suspension feeding and deposition of the outer cortical band a g e s o f the t h e c a l w a l l .

the

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 elem 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 possible 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 apparatus 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 bilateral 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, rem 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 distribution 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 tion

13.3.

of the

ReconstrucOrdovician

jawless fish, As t rasp is desiderata from stone

Walcott,

the Harding Sandof

Colorado.

Length about

13 cm.

From Cowen (2005, figure

7.4).

86,

Reprinted

with

permission

well

Publishing.

of Black-

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)

conodont species representing twenty genera (Figure

r e c o r d e d thirtv-live

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 , variations 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 prefere 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.

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

diverse in C i n c i n n a t i a n strata. 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 preserved 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 , 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

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-

A.

Formation,

Cincinnati,

Ohio, x 7.

B.

Repichnia

trilobite

Cryptoli-

of the thus,

similar to

ana,

CMC IP 37622,

zon

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. bite

Lebensspuren) has b e c o m e a s p e c i a l -

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 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 . 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 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

trail,

lian,

Rusophycus Maysvil-

Clermont Co.,

Ohio

(from

Osgood [1970,

plate

66,

figure 3[), x 0.8. D.

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,

'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 -

Trilo-

Corryville Formation,

ictyon,

m e n t of m o d e r n i e h n o l o g v .

C.

Cruziana,

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 .

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 -

hori-

intermediate

between and

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

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

Cruzi-

and locality

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 fossils (also k n o w n as i c b n o f o s s i l s or

Repich-

Paschichnia, CMC IP

?Paleod17431,

Kope Forma-

tion, Cincinnati, Ohio, 3.

E.

x

Fodinichnia or

domichnia, track,"

the

"turkey

Trichophycus

venosum

Miller,

CMC IP

struct the life of the past i( )sgood 1975). In rare eases a trace fossil c a n be

37575,

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,

( 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,

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

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 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 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

Campbell

Co.,

x 0.4. From plate

60,

by permission of the Paleontological Institution.

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

Research

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 patt 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 structures 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 interp 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

a n d Allocotkhnus trace

of t h e

t r a c e s . Asaphoidichnus

dyeri

trilobite

trifidum

M i l l e r r e p r e s e n t different lsotelus

(Osgood

1970).

(Miller) forms

(Figure

of the

Trachomatichnus

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

14.1A)

crawling numerosum 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 epireliefs. (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 des 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 . ies of the

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-

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 carb 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 . Chondrites, 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 overlapp 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 resistant 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 . tracks" (width 2.5-3.5

c

m

The large size of the "turkey

) 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 / ) . cincinnatiensis ( 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 organism 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 feeding 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 . nia

A.

and sderite

sions

of

trilobite

Rusophycus James),

Isotelus,

carleyi (J.

F.

CMC IP 46411,

Maysvillian,

Corryville

Formation,

Clermont

Ohio,

x 0.6.

sions

of cephalic

margin,

pleurae,

pygidial margin, coxae

Co.,

Note impres-

genal spines, as

Cubich-

impres-

as well

(medial paired

lobes,

flanked by cres-

centic

impressions

made

by legs. Also, at top, note Paleophycus terminating mate

burrow at

approxi-

position

of trilobite

mouth,

with

posed

scratchmarks.

superim-

B.

Flexicalymene

meeki

(Foerste),

37574,

CMC IP

Maysvillian,

ryville Formation, mont Co., 1.4.

Ohio,

Right:

x

Reverse side

of specimen shown cubichnia

CorCler-

in

pudicum

Hall,

convex

hyporelief,

identifying

trilobite

tracemaker.

This

as

exceptional

speci-

men is one of the very few known

with

trace

tracemaker

and

preserved

Type-Cinannatian Trace Fossils

207

B,

Rusophycus

both

together.

Figure

14.3.

Diplocraterion

A.

Kope Formation,

Osgood;

Cincinnati, Ohio,

Corryville Formation, mondian,

Vertical joint surface showing two commonly associated trace fossils: at left,

cincinnatiensis

x

at right, 1.3.

Clermont Co., Ohio,

Whitewater Formation,

forme (Miller and Dyer),

B. x 0.9.

Wayne Co.,

CMC IP uncatalogued,

208

fodinichnia, Fodinichnia, C.

Indiana,

1.2.

type-A,

Chondrites, type-B,

Fodinichnia, x

Maysvillian,

A Sea without Fish

Chondrites,

D.

domichnia,

CMC IP 37607,

Edenian,

CMC IP 37623, Maysvillian,

Chondrites, type-C,

CMC IP 37678, Rich-

Cubichnia of sea star, Asteriacites stelli-

PCorryville Formation,

Cincinnati,

Ohio,

x

7.

Figure

14.4.

A.

uncatalogued, rion

Vertical joint surface showing domichnia,

Edenian,

biclavata

Kope Formation,

(Miller),

showing

Campbell Co.,

U-tube and spreiten

Diplocraterion

Kentucky,

x 0.8.

constructed in

from Osgood (1970). IP 37657, Maysvillian, 0.4.

D.

C.

the upper surface of the calcisiltite,

Domichnia Diplocraterion biclavata

Corryville Formation, Clermont Co., Ohio,

bioimmuration structure in bryozoan,

Catellocaula vallata Palmer and Wilson, x.2.3.

E.

cypods, Lockeia siliquaria U. P. James, CMC IP 37597, Edenian, Kenton Co., Kentucky, holotype,

originally interpreted as (1970) to be either a tube,

x 0.6.

B,

CMC IP 24079, the impression

Maysvillian,

Corryville Formation,

of a porpitid jellyfish,

this impression

base a

Modified

"dumbbell" on upper surface,

(from Osgood [1970, plate 61,

nati collection, Edenian, Point Pleasant Formation, Bracken Co., Kentucky, floweri Caster,

with

When shale is eroded away,

as shown in lower diagram.

(Miller),

CMC IP,

Reconstruction of Diplocrate-

upper shale (dashed pattern)

of U and paired lateral extensions in underlying calcisiltite (stippled pattern). "dumbbell" impression remains on

cincinnatiensis Osgood, B.

figure 3]),

CMC x

University of Cincin-

Cubichnia of pelex 0.5.

Clermont Co.,

F.

Palaeoscia

Ohio. Although

was considered by Osgood

feeding trace or of inorganic origin as a result of rotatory sweeping of an agglutinated

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 exc 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 , intersection 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 stelliforme ( 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 m e n t , c h i e f l y in relation to d e p t h .

Interpretation

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, nearshore 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 increases, 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 , creating 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 expanded 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 Domichnia Repichnia

5 6 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 fod 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 , "Blastophycus," 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 , perfectly 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 currents 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 Geologists tionally

strata.

have

tradi-

defined

sedimen-

tary rocks on the basis of time,

usually inferred

from

fossil

and on type.

assemblages,

the basis of rock The

has

Cincinnatian

been

traditionally

divided into

three stages,

shown at the far left,

and

there is currently disagreement tive

over

the

durations

three stages. stage,

rela-

of these A

fourth

indicated by

and called the chian Stage, ent in

"G"

Gama-

is not pres-

the Cincinnati area.

The divisions based on rock type are shown at the right.

Most of those

in modern shown, different have

usage are

but dozens of named

been

divisions

proposed

over

the years and are not shown.

The names cur-

rently used by the and

Kentucky

surveys differ, dashed cate

Ohio

geological and

vertical lines

these

indi-

"stateline

stratigraphic

divisions."

The

Geological

Indiana

Survey

recognizes

Kope,

Whitewater,

Saluda

Formations,

assigns

all strata

the

Kope and

the and and between

Whitewa-

ter to the Bull Fork Formation

(not shown).

Type-Cincinnatian have also

been

strata divided

into six units that reflect cycles

of

sedimentation

produced by

the rise and

fall of sea level. depositional numbered bounded

These sequences,

C1-C6, by

reflect falls in sea level that drained the seas from the Cincinnati area, ing in

no deposition

ties is poorly known.

are

unconformi-

ties shown in gray that

214

A Sea without Fish

of sediments.

The actual duration

of these

result-

unconformi-

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 situated 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 cove 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 today'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-advanci 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 levels 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 of

15.2.

benthic

by

Destruction communities

storm-generated

waves and currents.

Dur-

ing calm,

con-

pre-storm

ditions,

benthic

nities

commu-

of organisms

develop on Under

the sea

storm

high

winds

floor.

conditions, generate

large waves that stir up fine-grained iments

bottom

into

sed-

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

suspension.

Stronger wave and current forces can benthic

suspended clog

displace

organisms,

and

sediment

feeding

can

and respira-

tory mechanisms

of or-

ganisms,

and even

smother

entire

benthic

communities.

Mobile

ganisms can

escape if

or-

burial is not too severe, but storms can be lethal for many thic

a storm, can until

immobile

organisms. cover

the

bottom communi-

develop

larval settlement migration. Hinterlong

S T A G E II. S T O R M C O N D I T I O N S

barren sediment

benthic

ties again

ben-

Following

from

or im-

Modified (1981).

tesy of Wayne D.

after

CourMartin.

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 carried. 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 3—5 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 sedim 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 lardg 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 interpretation 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), Hiscobeccus,

several

brachiopods

Lepidocyclus,

(Glyptorthis,

Holtedahlina,

Plaesiomys,

Rhynchotrema,

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 nutrientr 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 susp 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 sedim 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 Cincinnati Arch

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 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 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

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

Figure 1 5 . 3 . The four principal

sedimentary

environments

of

the

type-Cincinnatian.

and t e m p e r a t u r e on a d a i l y basis. 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 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 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

cinnatian

seas

deepened from

northward

shallow

tucky

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

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 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

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

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 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. 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

water envi-

ronments in

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

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

Cingenerally

to

central Ken-

deeper in

water Ohio

The bound-

between

these

four

correspond

to sea level,

fairweather

wave base,

and storm

wave base.

Wave base

reflects

the depth at

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

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

the sea

depth

floor

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

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,

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

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-bodied 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 extensiveb 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 duri 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

the height and peincrease

changed

over

both of during

thousands

to millions of years, environments

with

shifting

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

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

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 , 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 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 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

rising sea level. Falls in sea level resulted in

and

Paleoenvironment

the

draining of the seas from the

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

southward

times of rapidly

221

Cincinnati area.

Figure 1 5 . 4 . crop

A. Out-

photograph

laminated

of finely

dolomite

de-

posited in a tidal flat environment.

B.

photograph

of

weathering,

Outcrop rubbly

nodular

limestone

and

deposited in

mudstone

a

shallow

subtidal

environ-

ment.

C.

Outcrop pho-

tograph

of

limestone

and

interbedded mudstone

deposited in a

deep sub-

tidal environment, the

limestone

cording

re-

deposition

ing hurricanes. crop

with

beds D.

photograph

mudstone

with

of limestone stone,

durOut-

of thin

beds

and silt-

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 every

the

history of

sedimentary

there lurks an tem,

rock

ecosys-

of

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 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

but what one

first sees is an environment

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

by soft-bodied organisms such as w o r m s and arthropods. Shallow subtidal deposits are f o u n d in the

deposition.

type-Cincinnatian

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

E d w a r d S. D e e v e y

a n d are

m a r i n e deposits that

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 subtidal 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 .

222

A Sea without Fish

This distinctive b e d d i n g results from the per-

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 , sheetlike, 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

carnivorous

attached pelecypods

cephalopod

Treptoceras,

Ambonychia

a n d Caritodens, t h e

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 duri 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 rippled 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 subtidal 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 , sugg 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 Leptaena,

m a y be p r e s e n t ,

Hiscobeccus,

Platystrophia,

i n c l u d i n g Rafinesquina, Plectorthis,

Glyptorthis,

Strophomena, 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 indiv 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 Trichophycus

i n t h e siltstone b e d s . Chondrites a n d

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 mudstone 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

Dalmanella a n d Sowerbyella, Pseudolingula a n d Leptobolus, w h i c h

gregarious

ticulates

and are

the

burrowing

sometimes

found

inarpre-

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

Ambonychia Deceptrix and Rhytimya, t h e

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

olopsis,

the b u r r o w i n g p e l e c y p o d s

pelecypods

a n d Modiscavenging

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 c e p h a l o p o d s 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, Gravicalymene, 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 gastropods

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 crinoid 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, Trichophycus a n d Paleophycus are all

Diplocraterion,

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

226

A Sea without Fish

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 .

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

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

Ecological

Theater

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 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 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 particular their m o d e s of life and f e e d i n g habits. 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 organisms of a single species with the e n v i r o n m e n t , assessing the influence 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 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 "ancient" and enter the a n c i e n t realm of paleoautecology. 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 ,

Figure 16.1. toceras

duseri

Whitfield), (Meek) body

Trepcrinoid

baeri

preserved

Arnold

within Richard

Davis

Waynesville

collection, Formation,

Adams Co.,

Ohio,

col-

lected by Thomas Johnson. Taeniaster

B.

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),

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

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

with

chamber.

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-

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 ,

Internal

(Hall and

Xenocrinus

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 -

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

A.

mold of nautiloid,

mold

T.

Ophiuroid, spinosus

MUGM 28187, on

internal

of nautiloid,

Waynesville

Formation,

Butler Co.,

Ohio, scale in

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

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,

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 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 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 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 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 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 ? 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

CMC IP 2257, tian, bites, meeki

(Foerste),

served on

Trilopre-

internal mold

of nautiloid,

?Trepto-

ceras sp.,

OSU 50329,

Cincinnatian, Cincinnati,

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

D.

Flexicalymene

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 -

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

sp.,

Cincinna-

vicinity of Cincinnati,

Ohio, x 0.9.

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 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

of nau-

?Treptoceras

vicinity of

Ohio,

x 2.6.

C, D from Davis et al. (2001,

figures 2, 5), and

reprinted of

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

by

Blackwell

permission Publishing.

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 assemb 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 assemb 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 biostratig 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 rem 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 nus,

are

the slender crinoids

the small, thin-shelled

Cryptolithus a n d

Acidaspis

larger b r a c h i o p o d

Ectenocrinus and

b r a c h i o p o d Sowerhyella, a n d

Cincinnaticri-

t h e trilobites

(see F i g u r e s 11.5,11.6). Higher in the K o p e , the

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 abundant

brachiopods

brachiopods,

Rafinesquina a n d Strophomena are t h e m o s t

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 overlyi 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 faunal 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 gradient 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 ,

limited to particular fa-

brachiopods,

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 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 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 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 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 s u c h as the

brachiopods

Platystrophia a n d Strophomena

(Holland

1997).

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 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 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). 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 assemb l a g e s , s u c h as Hebertella a n d

Platystrophia,

are p r e s e n t in

cies, or

trophic groups, life-habit

rather

the

groups; Riehmondian

Invasion was a major ecological affecting the

revolution all aspects

Cincinnatian

of

seas.

Steven M. Holland 1997, 320

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,

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

Platystrophia, a n d b r a n c h i n g

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 present-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 understand 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 ecosyst 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 determ 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 prese 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 predatorprey 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 asma (Elias 1984).

Grewingkia

and

Streptel-

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 responsible. 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 original 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 parasitic, 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 ass 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 either 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 associated 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 gastrop 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-

tion of life on the Cincinnatian sea floor. by Anneliese Elizabeth A.

Drawing

Caster and Dalve.

From

von Engeln and Caster, Geology, 347,

1952,

of

the

by

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-

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 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 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).

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-

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.

McGraw-Hill,

reprinted

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

permission

McGraw-Hill

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 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, actively 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 diversify 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 presentdav 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 preserved 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 productivity 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 available.

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-environment diagram

for the

Cincinnatian Series.

The

vertical axis shows

the

timescale and six major shallowing-upward quences

of the

seCincinna-

tian (see chapters 4 and 15).

G = Gamachian Stage

(not preserved in nati region).

Cincin-

The horizon-

tal axis shows the major environments

of the

Cin-

cinnatian (see chapter Offshore

15).

environments

are located

towards

the

present north in Ohio and westward in Indiana, shoal and lagoon ments are

and

environ-

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 each

found

sequence

ronment,

in

and

envi-

defined as

fos-

sils occurring in >20% of samples ting.

from each set-

Letter-codes as fol-

lows: corals: Gr = Gewingkia,

Sp =

Streptelasma, radium,

Te =

Pr -

Tet-

Protaraea;

bryozoans: bf = bifoliate forms,

en = encrusting

forms, nr = thin ramose forms, ra = ramose forms; brachiopods: Da = Dalmanella, tella.

He

Hi =

=

Heber-

Hiscobeccus,

Ht = Holtedahlina,

Lp =

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 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 .

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,

= Plectorthis,

Ra

=

Rafinesquina,

Re

=

rorsirostra, chotrema, byella,

Pt Ret-

Rn = RhynSo

=

Sower-

St =

The causes of

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 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 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 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 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-

240

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

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,

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-

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 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 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.

Ct =

Cryptoli-

c o d e s = os; g a s t r o p o d s = ga; p e l e c y p o d s ; Am = Ambonychia,

by

= inde-

terminate pelecypod,

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,

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;

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.

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,

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,

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 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 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 -

cri-

Ci = CincinnaticriEc

=

Xenocrinus; gra.

Ca

mo = modi-

Ectenocrinus, Xe =

graptolites

those

invading

during

the

region

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.

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);

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

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 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

=

Italicized fossils are

bution

Information

desee

for distri-

of additional

fossils.

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 . Predator

Prey

References

ostracodes?

Elias 1 9 8 4

Nautiloids

brachiopods

Alexander 1 9 8 6

trilobites

Babcock 2003

crinoids

Prey among Taxa

Rugose corals

Unspecified

Table 2. Predators and

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

Cincinnatian

Table among

3.

Associations

Individuals

cinnatian

ARRANGED BY "HOST"

of Cin-

Taxa

Host

Associate

Type of Association

References /

bioimmuration; epizoism

Wilson, Palmer,

pelecypods

boring

Pojeta and Palmer (1976); Wilson and Palmer (1988)

bryozoan

encrustation (post-

Wilson, Palmer,

mortem);

and Taylor (1994)

[Annotations]

PROTISTA foraminifers

bryozoan

and Taylor (1994)

PORIFERA stromatoporoids

CNIDARIA hydrozoan

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)

algae

boring

Kobluk and Risk (1977)

corals

epizoism

Elias (1983)

post-mortem

Shrake(1989)

BRACHIOPODA brachiopods

brachiopods

attachment

bryozoans

encrustation; epizoism

Nicholson (1875)

bryozoans

bioimmuration; epizoism

Wilson, Palmer, and Taylor (1994)

"boring";

Anstey and Wil-

presumably post-

son (1996)

bryozoans

mortem bryozoans

"boring"

Pohowsky (1974, 1978)

bryozoans

post-mortem attachment

242

A Sea without Fish

Shrake(1989)

ARRANGED BY "HOST" Host

Associate

Type of Association

References /

bryozoans

encrustation; ? epizoism

Anstey and Wilson (1996) [Corynotrypa on interior of brachiopod shell]

bryozoans

encrustation; ?

Anstey and Wilson (1996) [Cuffeyella on interior of brachiopod shell]

[Annotations]

post-mortem

bryozoans

encrustation

Ulrich (1879)

bryozoans

encrustation

Ulrich (1883)

brachiopods

epizoism

Alexander and

Cornulites

? commensalism

Morris and Rollins (1971)

post-mortem

Shrake (1989)

Scharpf (1990)

crinoids

attachment edrioasteroids

epizoism

Richards (1972)

epizoism or

Meyer (1990)

commensalism gastropods

Sphenothallus

borings

Fenton and Fenton (1931)

borings

Bucher (1938)

epizoism

Neal and Hannibal (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; epizoism

Nicholson (1875)

bryozoans

encrustation

Ulrich (1879a)

bryozoans

encrustation

Ulrich (1883)

bryozoans

aegism; epizoism

Shrake (1989)

bryozoans

bioimmuration;

Wilson, Palmer,

epizoism

and Taylor (1994)

bioclaustration; parasitism;

son (1988)

Catellocaula

Palmer and Wil-

epizoism

Life in the Cincinnatian Sea

243

A R R A N G E D BY "HOST Host

Associate

Type of Association

Cornulites

References / [Annotations]

epizoism; ?

Morris and Rollins

commensalism

(1971)

encrustation and

Baird, Brett, and

intergrowth

Frey(1989)

pelecypods

boring

Wilson and

Sanctum

boring

Erickson and

Sphenothallus

epizoism

cornulitids

Palmer (1988) Bouchard (2003) Bodenbender et al. (1989) trilobites "worms"

aegism —

borings

Trypanites

Shrake(1989) Palmer and Wilson (1988)

CORNUUTIDS cornulitids

bryozoans

encrustation and intergrowth

cornulitids

bryozoans

bioimmuration; epizoism

Baird, Brett, and Frey (1989) Wilson, Palmer, and Taylor (1994)

MOLLUSCA Mollusca —

brachiopods

epizoism

gastropods Mollusca —

Cornulites

gastropods Mollusca —

Morris and Felton (1993)

Cornulites

encrustation;

S. A. Miller

? commensalism

(1874c)

commensalism

Morris and Rollins (1971);

gastropods

Morris and Felton (1993) Mollusca —

Cornulites

gastropods Mollusca —

? epizoism: ?

Richards (1974)

commensalism trilobites

? endozoism ?

Brandt (1993)

bryozoans

encrustation

Ulrich (1883)

Cornulites

epizoism or

Morris and Rollins (1971)

gastropods Mollusca — monoplacophorans Mollusca — nautiloids

brachiopods — (inartic.)

encrustation

Mollusca —

bryozoans

encrustation;

nautiloids

244

commensalism

A Sea without Fish

Davis and M a p e s (1996)

epizoism

Nicholson (1875)

ARRANGED BY "HOST" Host

Mollusca

Associate

bryozoans

"nautiloids" — actinoceroids endoceroids

bryozoans

Type of Association

References/ [Annotations]

epizoism; ?

Frey (1988, 1989);

commensalism

Baird et al. (1989)

encrustation

Davis and M a p e s (1996)

presumably epizoism;

nautiloids

? commensalism bryozoans

encrustation

bryozoans

encrustation

Ulrich (1879a) U. P. James (1884b)

bryozoans

encrustation

Ulrich (1883); Bassler (1953)

bryozoans

encrustation (post-

Wilson, Palmer, and Taylor (1994)

mortem); dwelling inside empty shell Cornulites

? epizoism ? commensalism

Richards (1974)

hydrozoan

encrustation (postmortem); dwelling inside empty shell

Wilson, Palmer, and Taylor (1994)

stromatoporoid

encrustation

J. F. James (1886)

stromatoporoid

encrustation

Baird, Brett, and

trilobites

? incolenism

Davis et al. (2001)

epizoism or

Morris and Rollins (1971);

Frey (1989)

Mollusca —

Cornulites

pelecypods

commensalism

Morris and Felton (2003) ARTHROPODA trilobites

bryozoans

epizoism

Brandt (1996)

trilobites

Cornulites

epizoism

Morris and Rollins (1971); Brandt (1996)

Echinodermata — crinoids

brachiopods

aegism; epizoism

Shrake (1989)

Echinodermata —

brachiopods —

commensal

Sandy (1996)

bryozoans

encrustation

Ulrich (1883)

bryozoans

encrustation: epizoism

Bassler (1953)

ECHINODERMATA

crinoids Echinodermata —

Zygospira

crinoids Echinodermata — crinoids

Life in the Cincinnatian Sea

245

ARRANGED BY "HOST" Host

Echinodermata

—

Associate

Type of Association

References/ [Annotations]

byroniids

parasitism

Warn (1974); Welch (1976);

crinoids

Malinky et al. (2004) Echinodermata

—

Cornulites

commensalism

crinoids

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

Echinodermata —

gastropods

commensalism

gastropods

? parasitism ?

crinoids Echinodermata crinoids "WORMS" (see also: cornulitids)

246

A Sea without Fish

—

Bowsher (1955); Morris and Felton (1993) Baumiller and Gahn (2002)

SUSPENSION

HERBIVORE

chitinozoans (?) conodonts graptolites trilobites (pt)

SUSPENSION trilobites (pt)

EPIFAUNA

MOBILE

ATTACHED LOW

ATTACHED ERECT

RECLINING

stromatoporoids (pt) tab., rugose corals (pt) bryozoans craniate brachs rhynch. brachs bivalves cornulitids edrioasteroids cyclocystoids sponges conulariids stromatoporoids (pt) tabulate corals (pt) bryozoans rhombiferans crinoids

rugose corals (pt) strophomenate brachs tentaculitids hyolithids stylophorans

DEPOSIT

PRIMARY PRODUCERS

CARNIVORE cephalopods eurypterids

acritarchs

HERBIVORE

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

CARNIVORE trilobites (pt) asteroids

SUSPENSION

DEPOSIT

CARNIVORE

rostroconchs SHALLOW PASSIVE

SHALLOW ACTIVE

Ungulate brachs bivalves Diplocraterion

bivalves polychaetes

polychaetes

DEEP PASSIVE

DEEP ACTIVE

Table 4 A . tian

Marine

Modified and

Life in the Cincinnatian Sea

247

Type-CincinnaGuilds

after

Sheehan

Droser (1997).

PELAGIC

SUSPENSION gastropods malacostracans mammals

SUSPENSION

ATTACHED LOW

ATTACHED ERECT

RECLINING

bony fish mammals

DEPOSIT

cephalopods dinoflagellates chondrichthyans coccolithophores bony fish diatoms reptiles mammals

chitons gastropods ostracods malacostracans echinoids

sponges corals bryozoans brachiopods polychaetes bivalves barnacles sponges corals bryozoans crinoids corals (pt) bivalves

SHALLOW PASSIVE

<

Mesozoic and

Cenozoic

Marine

Modeled

after

Guilds Bambach

(1983).

248

CARNIVORE gastropods cephalopods malacostracans asteroids

SUSPENSION

SHALLOW ACTIVE

DEEP PASSIVE

DEEP ACTIVE

Table 4B.

PRIMARY PRODUCERS

CARNIVORE

HERBIVORE

gastropods ostracods malacostracans ophiuroids asteroids (pt) echinoids holothuroids

INFAUNA

EPIFAUNA

MOBILE

bivalves crinoids ophiuroids (pt) asteroids (pt) holothuroids

HERBIVORE

A Sea without Fish

gastropods bivalves echinoids lingulate brachs bivalves polychaetes echinoids

DEPOSIT bivalves

bivalves polychaetes echinoids holothuroids

CARNIVORE bivalves

gastropods malacostracans polychaetes

bivalves

bivalves bivalves polychaetes polychaetes malacostracans

polychaetes

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 foss 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 recreated 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 65°F, 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 separated 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 percent. 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 app 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 bott 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 ins 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. Flexicalymene 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 current; 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 allowable 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 appare 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)

Publications of

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

(Feldmann

Exploring

Geology

the

of

Geological Surveys

a n d H a c k a t h o r n 1996) 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, 1961)

books

Encyclopedic w o r k s

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

R. A.

Locally published

on

Invertebrate

Paleontology

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 University 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 (acc 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 / (acc 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 (acc 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 roadlogs 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 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

Scientific societies 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/ (acc 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 organiz 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 .

Albers, H. E.

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).

Publications by T r u m a n H e m i n w a y Aldrich on

Tertiary a n d present-day

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

Aldrich, T. H. (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 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

A n t h o n y , John 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. (1878-1961)

( S e e c h a p t e r 2)

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 (1889-1971)

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 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 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.

Bridge, Josiah (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 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

Byrnes, Richard M. (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, origin 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 alm 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 M u s e u m of Natural History

( 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)

T h e Cincinnati Normal School

" N o r m a l s c h o o l s " were institutions that trained teachers and would-be 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 Society for t h e Promotion of Useful K n o w l e d g e

(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)

The Cincinnati Society of Natural History

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 . 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 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

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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 addition 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 fossils 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 University, 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 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

Deiss, Charles Frederick (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 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 -

Drake, Daniel (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 (1806-1883)

( S e e c h a p t e r 2)

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 (died 1930)

( S e e c h a p t e r 2)

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 F. (1862-1936)

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 , 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 Washi 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 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

Grinnell, 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 bryozoans

now

known

as

Parvohallopora

rugosa

and

Dekayia

Haime, Jules

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 society 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 W. (1832-1918)

( S e e c h a p t e r 2)

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 speciosa 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 subcylindrica 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 . ) .

( S e e c h a p t e r 2)

James, Joseph Francis (1857-1897)

( S e e c h a p t e r 2)

James, Uriah 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 (1806-1889)

( 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 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 1818—just 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 (1844-1912)

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 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 (1792-1856)

Dr. L o c k e was a c h e m i s t , educator, geologist, paleontologist, inventor of 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 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.),

Lyon, Sidney S. (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 (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 -

Mather, William 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. (1817-1876)

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 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 A l m o n d (1837-1897)

Milne-Edwards, Henri

( S e e c h a p t e r 2)

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 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 B. (1815-1885)

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 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. (1822-1892)

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 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 (1844-1899) 270

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 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

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 bryozoans

Monticulipora

mammulata,

Parvohallopora

ramosa,

and

d'Orbigny, Alcide

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 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

O r t o n , Edward (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 (1807-1860)

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 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 (1820-1902)

Henry Probasco was in the hardware business in C i n c i n n a t i . He was an 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 Proje 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. S. (1783-1840)

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 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 selfp 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 Glyptocrinus 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 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

Schlemmer, Charh (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 Berryman (1858-1947)

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 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 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 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

Appendix 2

Shaler, Nathaniel Southgate (1841-1906) 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. (1886-1958)

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 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 fossils 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 starfish 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 Henry (1875-1957)

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 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 Norm 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 B. (died 1933)

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 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, G e o r g e L.

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 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 , Enoploura, 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 . 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 -

Vaupel, Ernst H. (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 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

Warder, John 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 Academy of Natural Sciences (1835-1871)

T h e Western A c a d e m y of Natural Sciences was founded by Dr. Daniel 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 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 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 previo 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 Museum (1820-1867)

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 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 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 Gallatin (1833-1902)

Whitfield, R. P.

( S e e c h a p t e r 2)

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).

Willet

(See Letton's M u s e u m )

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 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

W o r t h e n , A. H.

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 acquisition. . . " ( 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 classification; 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 anat 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 bilaterally 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 Catellocaula 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 , litho 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 biostrome

(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 .

(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 biv 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 coelom

(see: t a x o n o m y ) .

(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 arthropods 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

282

Glossary

(adj.) f e e d i n g u p o n d u n g .

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 yourself (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 term

(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

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 "dorsal"; cf: anterior; distal; lateral; m e d i a l ; posterior; p r o x i m a l ; ventral.) durophagous

E

(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 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 "epiz 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 volcanica 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 fucoid

see b i o z o n e ) .

(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 presentday 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 org 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 Ord 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 paleontology

(n.) d e p t h o f a n a n c i e n t b o d y o f water.

(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 movem 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 , pres 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 ; rostroc 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 carbon-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

scientific n a m e

(see l i t h o s t r a t i g r a p h i c u n i t ) .

s

(see: s p e c i e s n a m e ) .

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 effect, 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 scie 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 specific n a m e

(see: s p e c i f i c 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 disturb 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 calcareo 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 involved, 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 system

(see: c o t y p e s ) .

(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, burrows, 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

unconformity

(see: s p e c i f i c n a m e ) .

(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 (Int 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. I .ike most indexes, this one is nowhere near as thorough as it might have been; the tremendous number of items that might have been included would have made the index completely unwieldy. Certain words and concepts are not separately listed in the index, because they appear far too many times in the book. Examples include Cincinnatian, Ordovician, and Ohio River. Although places in the states of Indiana, Kentucky, and Ohio are included in the index, every appearance of the name of the whole state is not. This volume is not intended as a comprehensive taxonomic work. Hence, we have not endeavored to put every genus, species, and so on, in its complete Linnaean Hierarchy in the index. Rather, we have used the names of upper-level taxa that appear in the text. However, we have departed from this policy in some cases, with the hope of greater clarity and usefulness for the reader. Although so-called "common names"—things like "brittle star" and "spiny oyster"— probably should he sandwiched between quotation marks, we chose not to do this so as not to clutter the index more than absolutely necessary.

Chlorophyta ("green algae"), 67, 68 Dasyeladaceae, 67, 68

Anomaloides: A. reticulatus, 68 Cyclocrinites: C. darwini, 66, 6 7 - 6 8 Ischadites: I. circularis, 68 Lepidolites: L. dickhauti, 66, 67, 68 Cyanophyta (called "blue-green algae," but not actually algae). See Cyanobacteria

Cylindrocoelia: C. covingtonensis, 68 Faberia: F. anomala, 68 Girvanella. See Cyanobacteria oncolites, 67 Phaeophyta ("brown algae"), 67 Pyrrophyta: dinoflagellates, 69, 239, 248, 283 Rhodophyta ("red algae"), 67, 68

Solenopora: S. richmondensis, 68 stromatolites. See Cyanobacteria a l i g n m e n t of fossils. See taphonomy a l l o c h e m , 49, 50

Allocotichnus: A. dyeri. See taphonomy: trace fossils 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 absolute age, absolute dating. See time Acadian Orogeny, M o u n t a i n s , 8, 62 Acanthochaetetes. See Porifera: sclerosponges, coralline sponges acetate peels. See techniques acid dissolution, acid etching. See techniques Acidaspis. See Arthropoda: Trilobita: Odontopleurida acme zone. See biostratigraphic units: zone: epibole acritarchs, 5 2 , 6 7 , 6 8 - 6 9 , 239, 247, 279, 309

Multiplicisphaeridium: M. micraulaxum, 6 6 , 67 Ordovicium: O. gracile, 66, 67 Veryhachium: V. edenense, 66, 67 Actinoceratoidea. See Mollusca: C e p h a l o p o d a

acutilirata, Platvstrophia. See Brachiopoda Adams County, Ohio, 75, 158, 162, 163, 216, 229, pl. 7 aegism. See ecologic associations Agassiz, Louis, 21, 260, 273 age, dating. See time Ager, Derek V., 7 Aglaspida. See Arthropoda Agnew, ]ohn. 7 1 , 72, 171, 180, pl. 14 alata, Ctenobolbina. See Arthropoda: Crustacea: Ostracoda alata, Nereigenys. See Annelida: scolecodonts Albers, H. E., 259 Aldrich, T. H., 259 Algae, 49, 5 2 , 6 7 - 6 9 , 123, 139, 212, 219, 234, 239, 241, 242,

279, 292. See also acritarchs; chitinozoans

Eurypterida

Amhonychia. See Mollusca: Pelecypoda American Midland Naturalist, 172 American Midcontinent Province. See faunal provinces American Upper Ordovician Standard, 51 amoenum, Beloitoceras. See Mollusca: C e p h a l o p o d a

Amorphognathus. See conodonts

Amphilichas.

See Arthropoda: Trilobita: Lichida Amsden, T h o m a s W., 54

anacanthus, Arnheimograptus. See

Hemichordata:

Graptoloidea Anderson, D. R„ 259

andersoni, Pontiometra. See Echinodermata: Crinoidea a n e m o n e s . See C n i d a r i a : Anthozoa Angseesing, J. P. A., 212, 297 angularis, Glyptocrinus. See Echinodermata: C r i n o i d e a : Camerata An n elida, 24, 31, 117, 142-145, 147, 182, 265, 306, 309, 316,

319. See also worms

Cornulites, 93, 123, 142, 144, 145, 182, 243, 244, 245, 246, 247, 314, 316 myzostomid annelid worms, 182 Polychaeta, 143, 144, 204, 205, 212, 221, 224, 238, 247, 248 serpulids, 144 spirorbids, 144

Protoscolex, 144 P. omatus, 142 scolecodonts, 143, 265

323

Nereigenys: N. alata, PL 5 Annual Reviews, 4 anomala, Faberia. See Algae Anomalocrinus. See Echinodermata: Crinoidea: Anomalodonta. See Mollusca: Pelecypoda

Eoleperditia, 163 Disparida

Anomaloides. See Algae: Chlorophyta: Dasycladaceae Anthony, John G o u l d , 2 5 9 - 2 6 0 , 261, 269, 276 Anthozoa, anthozoans. See C n i d a r i a Anticosti Island, C a n a d a , 7 2 - 7 3

Aphelognathus. See conodonts Aplaeopliora. See Mollusca Appalachian M o u n t a i n s , basin, 3, 5, 8, 10, 53, 6 1 , 216, 220, 231, 233, pl. 12 approximate, Palaeasterina. See Echinodermata: Asteroidea Arachnida. See Arthropoda: Chelicerata aragonite, aragonitic. See c a l c i u m carbonate area, Archinacetla. See Mollusca: Monoplacophora Archaeogastropoda. See Mollusca: Gastropoda Archinacella. See Mollusca: Monoplacophora A r n h e i m Formation, 44, 4 8 , 5 3 , 6 8 , 105, 107, 108, 123, 151, 153, 164, 176, 180, 196, pl. 8 Oregonia M e m b e r , 44, 53, 56, 63, 223. See also Oregonia Formarion Sunset M e m b e r , 4 4 , 53, 56, 63, 164, 224. See also Sunset Formation Arnheimograptus. See Hemichordata: Graptoloidea Arthropoda, 6, 92, 117, 129, 146-164, 204, 205, 221, 222,

224, 245, 253, 280, 282. See also Chelicerata: Arachnida Aglaspida, aglaspids, 158, 163, 280, 300

Neostrabops, 158, 163; N. martini, 158, 163 C h e l i c e r a t a , 27, 161 Arachnida, 147, 161 mites, 161 scorpions, 161, 162 spiders, 161, 162 ticks, 161

Quadrijugator: Q. regularis, 160, 161 Diplopoda: millipedes, 147 Insecta, 105, 119, 147, 162, 282 scorpions. See Chelicerata: Arachnida spiders. See Chelicerata ticks. See Chelicerata: Arachnida Trilobita, 6, 7, 27, 3 1 , 4 0 , 56, 6 0 , 9 2 , 9 3 , 117, 137, 144,145, 147-157, 1 6 0 - 1 6 1 , 1 6 2 , 1 6 3 , 2 0 2 , 203, 204, 205, 207, 210, 212, 219, 220, 223, 224, 225, 2 2 6 , 2 2 8 ,

229, 231, 232, 233, 235, 238,240, 241, 244, 245, 247, 251, 252, 253, 260, 268, 275, 286

Acidaspis. See Odontoplcurida

Calymene, 149, 150, 296, 305 C. meeki. See Flexicalymene meeki C. meeki-retrorsa. See Flexicalymene retrorsa C. senaria. See Flexicalymene senaria C a l y m e n i d a , c a l y m e n i d s , 149, 151, 226, 241

Flexicalymene, 40, 148, 150, 151, 152, 153, 154, 160, 207, 210, 224, 225, 226, 229, 232, 251, 253, 264, 298, 317 F. granulosa, 149, 150, 151,309 F. meeki, 148, 149,150, 151,207, 228, 229, 264, 298, 317 F. retrorsa, 149, 150, 151, pl. 8

F. senaria, 149, 153 Gravicalymene, 225, 226 Cryptolithus, 113, 155, 203, 204, 210, 225, 226, 232, 240, 241, 253

C. tessellatus, 148, 149, 154, 155 Decoroproetus: D. parviusculus, 154, 155 Flexicalymene. See C a l y m e n i d a Gravicalymene. See C a l y m e n i d a 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, 247, 253, 280, 300, 305

Eocarcinosoma: E. batrachophthalmus, 162 Megalograptus, 154, 158, 159, 161, 162, 253, 305, 312 M. aheolatus, 162 M. ohioensis, 158, 159, 162 M. shideleri, 162 M. welchi, 162 M. williamsae, 162 Xiphosurida: horseshoe crabs, Limulus, 147, 151, 161, 163, 280, pl. 3 C h i l o p o d a : centipedes, 147 C r u s t a c e a , 26, 30, 147, 148, 151, 163, 210, 237, 238, 241 barnacles, 145, 188, 248, 250 Isopoda, 148 pillbug, 148 Malacostraca, 248 crabs, 9 6 , 148, 163, 235 lobsters, 148, 163, 253 shrimps, 163 Ostracoda, 30, 31, 35, 129, 147, 160, 163-164, 221, 235, 241, 247, 248, 297, 320

Ceratopsis: C. chambers!, 160, 16J Ctenobolhina: C. alata, 160, 161 324

Lepcrditicopida, 164 Palaeocopida, 163 Podocopida, 163

Index

210, 224, 225, 226, 232, 235,240, 241, 252, 299, 305, 316, 317

J. brachycephalus, 153 /. gigas, 153 /. latus, 149 1. maximus, 148, 149, 152, 153, pl. 7 1. rex, 153, 316 I j c h i d a , lichids

Mlolichas: A. halli, pl. 6 Amphilichas: A. halli. See Allolichas halli Odontoplcurida, odontopleurids, 156, 160,251

Acidaspis, 228, 229, 232, 251 A. cincinnatiensis, 156, 160 A. onealli, 271 Primaspis, 92, 160 P. crosotus, 156, 160 Olenida, olenids

Triarthrus, 146, 147, 160, 226, 253 T. eatoni, 154, 155, 160 T. spinosus, 160 Phacopida, phacopids

Ceraurinus, 220 C. icarus, 157 Ceraurus: C. milleranus, 157, 269

Platycoryphe: P. christyi, 157 Tricopelta: T. breviceps, 157

Bellevue Limestone, J l , 12, 53, 55, 56, 63, 68, 85, 89, 95, 99,

Platycoryphe. See Phacopida Primaspis. See Odontopleurida

Eormation Beloitoceras. See Mollusca: C e p h a l o p o d a benthic, benthonic, benthos, 3, 4, 7, 50, 102, 145, 163, 195, 204, 205, 218, 230, 234, 237, 239, 280, 288, 290 bentonite. See K-bentonite Berdan, Jean, 163, 164, 320 B e r g m a n , C l a e s F., 143, 304 Bergstrom, Jan, 153, 318 Bergstrom, Stig M., 2 , 5 1 , 62, 196, 306, 308, 319, 321, 331 Best, Robert, 260, 276 bibliographies, compilation of, 15, 33, 35 biclavata, Diplocraterion. See taphonomy: trace fossils bilix lata, Cyclonema. See Mollusca: Gastropoda bilix, Cyclonema. See Mollusca: Gastropoda binomen. See taxonomy: nomenclature: species n a m e Binomial System of Nomenclature. See taxonomy: nomenclature

125, 174,214, 223, 232, 251. See also M c M i l l a n

Proetus parviusculus. See Decoroproetus parviusculus trace fossils, 151-153. See also taphonomy: trace fossils

Rusophycus. See taphonomy: Triarthrus. See Olenida

trace fossils

Articulata. See Brachiopoda taphonomy: trace fossils Ascocerida. See Mollusca: C e p h a l o p o d a : Nautiloidea Ashermann, George (?), 273, 275 Ashgillian Stage, pl. 2 associations. See chapters on individual groups of a n i m a l s ; ecologic associations Asteriacites. See taphonomy: trace fossils Asteroidea. See Echinodermata Astraspis. See Chordata: Vertebrata: Agnatha

Asaphoidichnus. See

auhurnensis, Platystrophia ponderosa. See Brachiopoda: Platystrophia ponderosa Audubon, John James, 16, 277

Aulacera. See Porifera: Stromatoporoidea Ausich, W i l l i a m I„ 186 Austin, George M., 260 Austinella. See Brachiopoda: Articulata Australia, 199, 215, 241, 251. See also Great Barrier Reef auteeology. See ecology Babcock, Loren E. 153

bacteria, 123, 239. See also cyanobacteria

baeri, Charactoceras. See Mollusca: C e p h a l o p o d a baeri, Xenocrinus. See Echinodermata: Crinoidea:

Camerata

Bahama Platform, ix, 220

balanoides, Enoploura. See

Echinodermata: Stylophora Baldwin-Wallace College, 29, 30 ball-and-pillow structures, 60, 280 Baltica. See paleogeography Bambach, Richard K„ 237, 238, 239, 300, 315 Bardstown Eormation, 2H barnacles. See Arthropoda: Crustacea Barnhart, W . C , 260, 266 Bassler, Ray S., 18, J9, 20, 21, 25, 30, 33, 3 4 - 3 6 , 48 Bassler, Simon Stein, 34 "Bassleroscope," 35

batrachophthalmus, Eocarcinosoma. See

Arthropoda: C h e l i c -

erata: Eurypterida Baumiller, Tomasz K., 124, 125, 183, 186 Bean, W. H„ 260 beani, Ceramopora. See Ectoprocta: Cystoporata Bear Creek K-bentonite "zone." See K-bentonite, K-bentonite beds, K-bentonite "zones" Bear Creek Quarry. See Ohio: Clermont C o u n t y bedding index, 49 Bedford, Indiana, 10 behavior (of animals). See chapters on individual groups of a n i m a l s ; taphonomy: trace fossils Bell, Bruce M., 180, 189,251 Bell, C M . , 212 bellerophontids. See Mollusca: Gastropoda Bellevue Hill Park, C i n c i n n a t i , Ohio, 21 Bellevue House, C i n c i n n a t i , Ohio, 21

bioclaustration, 156, 210, 243, 281. See also taphonomy: bioimmuration biocoenose, biocoenosis. See life assemblage biofacies. See facies bioherm, 7 1 , 72, 7 8 - 8 1 , 85, 9 1 , 131, 242, 281 "bryoherm," 91 Great Barrier Reef, Australia, 9 1 , 241 patch reef, 80, 81 bioimmuration. See taphonomy: bioimmuration biostratigraphic units, 4 6 , 281 zone, 46 b i o z o n e , 4 6 , 195, 2 8 1 , 2 8 4 epibole, a c m e zone, 60, 281, 284 biostratigraphy, 52 biostrome, 81, 281 biota! succession, 4 7 - 4 9 , 6 1 , 281, 292 biozone. See biostratigraphic units: zone Bischoff, G. C. O., 182 bivalved a n i m a l s , 281, 290. See also Arthropoda: C r u s t a c e a : Ostracoda; Brachiopoda: Mollusca: Pelecypoda Bivalvia. See Mollusca: Pelecypoda Blackwell Publishing, 185, 199, 229, Blanchester Member. See Waynesville Formation

Blastophycus, 212 Blue Limestone, 47 Blue M i a m i Limestone, 47 B M H N . See Natural History M u s e u m , London Boardman, Richard S., 185 body fossils, 6, 143, 203, 204, 212, 281, 293. See also taphonomy, preservation Bolivia, 200 bony fish. See Chordata: Vertebrata: Osteichthyes Boone County, Kentucky, 87, 110, 111, 142, 180 boring, borings, 74, 7 5 , 7 7 , 9 0 , 9 3 , 119, 123-125, 130, 182, 203, 206, 210, 211, 236, 242, 243, 244, 293. See also taphonomy: trace fossils Bouchard, Timothy D., 206,'304 bowdeni, Loxoplocus. See Mollusca: Gastropoda: Paupospira bowdeni, Paupospira. See Mollusca: Gastropoda Bowsher, Arthur L., 121 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, 129, 130, 137, 144, 162, 172, 178, 179, 182, 188, 189, 206, 219, 220, 221, 223, 224, 225, 231, 232, 233, 235, 237, 240, 241, 242, 243, 244, 245, 248, 250, 251, 252, 253, 260, 261, 272, 274, 280, 281, 283, 285, 287, 291, 293 Articulata, 40, 93, 100, 101, 102, 103, 105, 112, 130, 261, 283,291,293

Austinella, 112 A. scovillei, 273

Ungulate brachiopods, lingulides, 103, 105, 247, 248

Petrocrania: P. scahiosa, 40, 104, 105 Philhedra: P. laelia, 104, 105, 108 Pseudolingula, 104, 105, 225. See also Lingula Schizomania: S. filosa, 104, 105 Trematis, 128 T. millepunctata, 4 0 , 104, 105 Brachiospongia. See Porifera brachycephalus, lsotelus. See Arthropoda: Trilobita

Catazyga, 112

C. schuchertana, 111 Dalmanella, 57, 113, 114, 225, 232, 233, 240, 253 D. emacerata, 106 Glyptorthis, 112, 220, 224, 233 G. insculpta, 107 Hebertella, 100, 104, 105, 113, 223, 233, 240, 251 H. occidentalis, 107 Hiscobeccus, 112, 220, 224, 240 H. capax, 111 Holtedahlina, 1 1 2 , 2 2 0 , 2 4 0 Lepidocyclus, 75, 220 Leptaena, 40, 112, 220, 224, 233, 240 L. richmondensis, 40 Onniella, 112, 124 O. meeki, 60 Orthis: O. scovillei (See Austinella scovillei) Orthorhynchula, 112 Plaesiomys, 108, 112, 220, 224, 233 P. subquadrata, 108 Platystrophia, 109, 112, 113, 223, 224, 232, 233,240, 250,251,252

P. acutilirata, 109 P. laticosta, 109 P. ponderosa, 39, 59, 98. 99, 109, 240, 250 P. ponderosa auburnensis, 39 P. ponderosa ponderosa, 39 Plectorthis, 1 1 2 , 2 2 4 , 2 4 0

P. neglecta, 106 Rafinesquina, 4 0 , 59, 75, 8 7 , 8 8 , 8 9 , 9 9 , 104, 105,110, 112, 113, 114, 189, 223, 224, 232, 233, 235, 240, 251,252,272

R. alternata, 110 "shingled Rafinesquina beds," 59, 9 9 , 110, 114, 251

Retrorsirostra, 112, 240 R. carleyi, 108, 261 Rhynchotrema, 112, 220 R. dentatum, 111 Sowerbyella, 112, 113, 225, 232, 240, 253 S. rugosa, 106 Strophomena, 112, 113, 114, 224, 232, 233, 240, 247, 252, 261

Tetraphalerella, 112 Thaerodonta, 112 7'. rugosa, 40 Thecidellina, pl. 3 Zygospira, 41, 112, 232, 233, 245, 251, 252 Z. modesta, 111, 178, 179 I n a r t i c u l a t a , 4 0 , 9 3 , 101, 102, 103, 108, 115, 128, 225

Leptoholus, 225 Lingula, 5, 103. See also Pseudolingula 326

Index

Bracken County, Kentucky, 178, 209 Bradshaw, Lael E., 22, 24, 25 Brandt, Danita S., 151, 153 Branson, C . C , 196 Branson, E. B., 196, pl. 5 Branstrator, Jon W., 182, 184, 189 Braun, E. Lucy, 260 Braun, Frederick, 2 6 0 - 2 6 1 , 270 Bretsky, Peter, 231 Brett, Carlton E., 60, 6 1 , 236, 295, 297, 310, 311, 318 Trilobita: Phacopida Bridge, Josiah, 261

breviceps, Tricopelta. See Arthropoda:

British M u s e u m (Natural History). See Natural History Muse um, London brittle stars. See Echinodermata: Ophiuroidea Broaddus, Billie, 29 Brongniart, Alexandre, 61 Brookville Formation, 44 Excello M e m b e r , 44 Liberty Member. See Liberty Formation, M e m b e r Station Hollow M e m b e r , 44 Waynesville M e m b e r . See Waynesville Formation Brookville, Franklin County, Indiana, 184,196 Brown County, Ohio, 123, 128, 151 Brown, Roland Wilbur, 40 Brown, Steve, 155, 156, 157 "bryoherm." See bioherm; Ectoprocta bryozoan. See Ectoprocta Bucher, Walter H., 24, 4 8 , 123 Bull Fork Formation, 44, 214 Burgess S h a l e , 6, 249 burrows, burrowing. See taphonomy: trace fossils, ichnofossils, Lebensspuren Butcher, W i l l i a m , 320 Butler County, Ohio, 6 8 , 75, 80, 87, 89,107, 108, 126, 127, 134, 137, 153, 156, 160, 176, 195,229 Byrnes, Richard M., 261 Byronia. See Byroniida, byroniids Byroniida, byroniids, 182, 246. See also Phosphannulus

Byronia, 182 byssus, byssal threads. See Mollusca: Pelecypoda C I , C 2 , C 3 , C 4 , C 5 , C 6 sequence, 55-56, 63, 214, 231, 232-233, 240. See also sequence stratigraphy Calapoecia. See C n i d a r i a : Anthozoa: Tabulata calcareous. See c a l c i u m carbonate calcite, calcific. See c a l c i u m carbonate c a l c i u m carbonate, 6, 50, 67, 71, 101, 102, 117, 119, 134, 144, 147, 167, 281 aragonite, aragonitic, 6, 101, 117, 120, 129, 134, 280, 281, 283, 290 calcareous, calcareous shell, calcareous skeleton, 3, 56,

6 0 , 6 7 , 7 1 , 72, 79, 13?, 144, 145, 169, 188, 206, 239, 280, 2 8 1 , 2 8 9 , 2 9 2 calcite, calcitic, 6, 7,74, 89, 101, 117, 120, J24, 130, 137, 139, 163, 167, 168, 188, 280, 281, 283, 290 calcium phosphate, 6, 81, 102, 182, 196, 198, 219, 223 Calloway Creek Formation, 214

Calymene meeki. See meeki

Arthropoda: Trilobita:

Flexicalymene

C a l y m e n i d a , calymenids. Sec Arthropoda: Trilobita Calvptomatida. See hyolithids; Mollusca C a m b r i a n , xix, 2 , 4 , 6 , 30, 8 2 , 9 9 , 139, 147, 152, 160, 163, 167, 168, 182, 186, 191, 195, 199, 237, 249, 280, 282, 289 " C a m b r i a n Explosion," 2 "Cambrian Fauna," 2, 237 Camerata. See Echinodermata: Crinoidea Cameroceras. See Mollusca: C e p h a l o p o d a : Endoceratoidea camouflage, 9 6 , 280

campanulaeformis, Cyathochitina. See chitinozoans C a m p b e l l County, Kentucky, 59, 109, 203, 209 C a m p b e l l , Kenton S. W., 155, 160 Canada, 3,215,220, 233,249,253 Anticosti Island, 7 2 - 7 3 British C o l u m b i a , 6 C a n a d i a n Shield, 3, 216 Manitoba, 153

canadensis, Crewingkia. See

also eonchiolin; pseudochitin chitinozoans, 52, 68, 69, 247, 282. See also Algae

Conochitina: C. hirsuta, 66, 67 Cyathochitina, 66, 67; C. campanulaeformis, 66, 67 Hercochitina: H. turnbulli, 6 6 , 67 chitons. See Mollusca: Polyplacophora Chlorophyta. See Algae Chondrites. See taphonomy: trace fossils

Chordata, 38, 169, 196, 198, 282. See also conodonts Urochordata: tunicates, 210, 281. Vertebrata

See also Catellocaula

Agnat ha ("jawless fish"), 199, 200

Astraspis: A. desiderata, 199 gnathostome fish ("jawed fish"), 200 Osteichthyes ("bony fish"), 198

Reptilia: Virginia, 41

christyi, Platycoryphe. See C n i d a r i a : Anthozoa: Rugosa

C a n a d i a n System, 30, 31

capax, Hiscobeccus. See

Brachiopoda: Articulata Carboniferous, xix, 4, 23, 191, 198, 261, 270 Caritodens. See Mollusca: Pelecypoda Carley, S. T., 261

carfeyi, Retrorsirostra. See Brachiopoda: Articidata cadeyi, Rusophycus. See taphonomy: trace fossils Carlson, Sandra J . , 192, 321,322 Carneyella. See Echinodermata: Edrioasteroidea carpoids. See Echinodermata Carr, T i m , 54 Carriker, Melbourne R., 124 casei, Opisthoptera. See Mollusca: Pelecypoda casei, Plicodendrocrinus. See Echinodermata: Crinoidea: Cladida Caster, Kenneth E., 25, 27, 28, 30, 31, 3 5 , 4 8 , 9 3 , 138, 151, 158, 159, 162, 163, 186, 191, 209, 212, 215, 259, 275, 276, 299, 320, pl. 13

Catazyga. See Brachiopoda: Articulata 9 3 , 1 5 6 , 209, 210, 243, 281. See also bioclaustration; bioimmuration; Chordata: Urochordata: tunicates

C. vallata, 156, 209, 210, 281 Cenozoic Era, xix, 25, 237, 238, 248, 284 centipedes. See Arthropoda: Chilopoda Cephalopoda. See Mollusca Ceramopora. See Ectoprocta: Cystoporata Ceratopsis. See Arthropoda: C r u s t a c e a : Ostracoda Ceraurinus. See Arthropoda: Trilobita: Phacopida Ceraurus. See Arthropoda: Trilobita: Phacopida Chaetognatha, 198, 282

chambersi, Ceratopsis. See Arthropoda: Chappars, Michael S., 25

C h e e t h a m . A l a n H . , 185,298 Cheilostomata. See Ectoprocta Cheirocystis. See Echinodermata: Rhombifera Chelicerata. See Arthropoda C h i c a g o , University of. See University of C h i c a g o Chilopoda. See Arthropoda chitin, chitinous, 6 , 6 9 , 82, 119, 147, 148, 161, 181, 282. See

Christy, David, 262

cancellatus, Sinuites. See Mollusca: Monoplacophora

Catellocaula,

Charactoceras. See Mollusca: Cephalopoda

Crustacea: Ostracoda

Arthropoda: 'Trilobita: Phacopida chronostratigraphic units. See time-stratigraphic units 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. See also Findlav Arch; Kankakee Arch C i n c i n n a t i Group, 28, 47, 143, 266, 269, 277 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 C i n c i n n a t i School of Paleontology, 15-36 C i n c i n n a t i , Ohio Boudinot Avenue and Westwood Northern Boulevard, 151 C i n c i n n a t i Astronomical Society, 21 C i n c i n n a t i C o l l e g e of Pharmacy, 22 C i n c i n n a t i Historical Society, 21, 215 C i n c i n n a t i Taw C o l l e g e , 24 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, 87, 89, 9 0 , 9 5 , 105, 110, 111, 120, 123, 126, 134, 137, 142, 144, 151, 153, 155, 157, 158, 159, 160, 170, 172, 173, 174, 178, 180, 186, 203, 207, 208, 209, 229, 256, 263, pl. 8 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 Society of Natural History C i n c i n n a t i Normal School, 27, 262 C i n c i n n a t i Observatory, 21 C i n c i n n a t i Society for the Promotion of Useful Knowledge, 259, 261, 276 C i n c i n n a t i Society of Natural History, 15, 16, 21, 22, 23, 24, 26, 27, 28, 29, 30, 32, 33, 34, 259, 260, 261, 2 6 2 - 2 6 3 , 264, 265, 2 6 6 , 267, 268, 270, 272, 275, 276, pl. 13

Journal of the Cincinnati Society of Natural History, x, 17, 259, 276

Eden Park Reservoir, 29 M e d i c a l C o l l e g e of Ohio, 29 Ohio M e c h a n i c s Institute, 31, 262 Spring Grove Cemetery, 22, 23, 24, 25 Woodward High School, 18, 26, 32, 33, 34 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

Cincinnatidiscus. See Echinodermata: Edrioasteroidea

polyps, 74, 7 7 , 7 9 , 182, 206, 235 reefs. See biohcrms

cincinnatiensis, Acidaspis. See

Scyphozoa, 74, 82, 182. See also Byroniida

Echinodermata: C r i n o i d e a : Disparida Arthropoda: Trilobita:

Odontoplcurida

cincinnatiensis, Diplocraterion. See taphonomy: cincinnatiensis, Isorophus. See Echinodermata:

trace fossils

Edrioasteroidea circular is, Ischadites. See Algae: Chlorophvta: Dasvcladaceae C l a d i d a . See Echinodermata: C r in o idea c h i l l i s . See Mollusca: Pclccvpoda Clark, T h o m a s H., 3 Clarkson, Euan N. K., 160, 299 Clarksville M e m b e r . See Waynesville Formation class. See taxonomy: l . i n n a e a n Hierarchy c l . i s s i h i ation. Sec lavonoim clastic ratio, 49. See also shale-to-limestone ratio clavacoidea: Leptotrypa. See Ectoprocta: Trepostomata C l a y s Ferry Formation, 163, 214 C l e r m o n t C o u n t y , Ohio, 111, 126, 128, 153, 158, 163, 170, 186, 203, 207, 208, 209, 261 Stonelick C r e e k , 11,254 Climacograptus. See Hcmichordata: Graptoloidea Clinton County, Ohio, 60, 72, 75, 128, 136, 142 W i l m i n g t o n , 260, 273, 276 cluster analysis. See techniques 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 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 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, 131, 182, 196, 206, 219, 220, 221, 224, 233, 235,

240, 241, 242, 243, 247, 248, 281, 285. See also Byron iida a n e m o n e s . See Anthozoa Anthozoa, 74, 77 a n e m o n e s , sea a n e m o n e s , 73, 9 6 , 188 Rugosa, 74, 75, 7 7 , 7 8 , 7 9 , 2 0 6 , 220, 221, 235, 241, 247

Gvathophvlbides, '<>. ~~. 79 C. stellata, 78 Grewingkia, 41, 77, 206, 220, 224, 235, 242 G. canadensis, 74, 75, 77 Streptelasma, 11, 220, 224, 235, 240 S. divaricans, 75, 77

Foerstephyllum, 76,11, 79 F. vacuum, 78 Nyctopora, 76, 77, 79 Protaraea: P. richmondensis, 76,11, 79, 122, 123, 224, 240

hydroids. See Hvdrozoa Hydrozoa, 74, 93, 242, 245 fire coral, 74 hydroids, 74, 281 Siphonophorida

Index

Goleolus. See hyolitbids collagen, 195, 196 colonial organisms, colonies, 189, 281, 290. See C n i d a r i a , Ectoprocta, graplolites color patterns, 9 6 , 134, 135, 137, 252, pl. 6 Colorado, 199, 216 C o l u m b i a University Press, 112 C o l u m b i a n University See George Washington University Gomactinia. See Echinodermata: Crinoidea Comasterida. See Echinodermata: Crinoidea c o m m e n s a l i s m . See ecologic associations c o m m u n i t i e s . See ecology, ecologic needs: ecosystems competition, competitive interactions. See ecologic associations composite molds. See taphonomy: molds

Amorphognathus: A. ordovicicus, 196, 197 Aphebgnathus: A. grandis, 196, 197 Drepanoistodus: D. suherectus, 196, 197 Oulodus: O. oregonia, 196, 197 Phragmodus: P. undatus, 196, 197, pl. 5 Plectodina: P. tenuis, 196, 197 Prioniodus: P. dychei, 28, 277 Rhodesognathus: R. elegans, 196, 197 C o n r a d , T i m o t h y A., 109, 110, 111, 120, 123, 126, 127, 149 Gonstellaria. See Ectoprocta: Cystoporata constellata, Gonstellaria. See Ectoprocta: Cystoporata constrictus, Dystactocrinus. See Echinodermata: Crinoidea:

Tetradium, 76,11, 79, 220, 233, 240

328

planispiral coiling, 119, 120, 139, 289 C o l b a t h , G. Kent, 67, 69 C o l d Spring, Kentucky, 174

See also Cbordata

luhastraea: T. coccinea, pl. 3

m e d u s a , jellyfish, 7 1 , 74, 182, 209, 212, 251

sea anemones. See Anthozoa sea fans, 74 sea whips, 74 coarsening-upward cycle. See cycles, cyclicity coccinea, luhastraea. See C n i d a r i a : Anthozoa: Scleractinia cockles. See Mollusca: Pelecypoda C o d e of Stratigraphic Nomenclature. See stratigraphy (loclenterata. Sec I Inidaria coiling of shell, 117, 119, 120, 132, 134, 139

cones (the snails). See Mollusca: Gastropoda: Neogastropoda C o n n e c t i c u t A c a d e m y of Arts and Sciences, 149 Conochitin. See chitinozoans conodonts, 6, 28, 31, 35, 5 2 , 6 3 , 143, 183, 195, 196-199, 247.

Tabulate, 77, 79, 8 0 , 1 2 3 , 220, 221, 224, 247 Galapoecia, 76, 7 7 , 7 9

209,

Conularia C. formosa, 80, 82, 267, 269 C. miseneri, 270

conchiolin, 119, 282. See also chitin

Scleractinia

Portuguese Man-of-War, 74 Porpitidae: Palaeoscia: P. ftoweri, taphonomy: trace fossils jellyfish. See medusa

C o n u l a r i i d a , 74, 8 1 - 8 2 , 241, 247; but may not be cnidarians, 74, 8 1 - 8 2 , 241, 247

212.

See also

Disparida consumers. See ecology: ecosystems Gonularia. See C n i d a r i a : Scyphozoa: C o n u l a r i i d a C o n u l a r i i d a , conulariids. See C n i d a r i a : Scyphozoa convergent evolution. See organic evolution (lomvav Morris, S i m o n . 249 Cook, W. E., 263 Cooper, Dan L., 153, 309, pl. 6

Cooper, Edward M , 263 coprolites. See feces coprophagy. See food copulation, 162 coquina, 99, 283. See also shell pavements; shingled beds; Waynesville Formation: Marble Hill bed coral. See Cnidaria coral reef. See bioherm Corallidomus. See Mollusca: Pelecypoda coralline sponges. See Porifera: sclerosponges Cornulites, Sec: Annelida

correlation, 52. See also time Corryville Formation, M e m b e r , 12, 21, 53, 55, 60, 63, 68, 80, 90, 105, 110, 111, 114, 120, 126, 128, 142, 144, 151, 153, 158, 163, 170, 174, 175, 178, 180, 186, 203, 207, 208, 209, 211,214, 224, 251. See also: Grant Lake Limestone; also: M c M i l l a n Formation cotypes. See taxonomy: type specimens Covington Croup, 48 Covington, Kentucky, 19, 28, 127, 142, 256, 263, 266, 274 covmgroiiensis, Cylindrocoelia. See Algae C o w e n . Richard, 199 crabs. See Arthropoda: Crustacea: Malacostraca crateriformis, Lichenocrinus. See Echinodermata: Crinoidea: Disparida Crawfordsville, Indiana, 23, 261, 270 Hovey M u s e u m , Wabash College, 23 Cretaceous, xix, 2, 3,4, 72, 239, 292 Cricoconarida. See also Mollusca; Tentaculitoidea Crinoidea. See Echinodermata crosotus, Primaspis. See Arthropoda: Trilobita: Odontopleurida Crustacea. See Arthropoda Cruziana. See taphonomy: trace fossils cryptolithi, Rusophycus. See taphonomy: trace fossils Cryptolithus. See Arthropoda: 'Trilobita Cryptostomata. See Ectoprocta

Ctenobolbina. See Arthropoda: Crustacea: Ctenodonta. See Mollusca: Pelecypoda

Ostracoda

Ctenostomata. See Ectoprocta cubichnia. See taphonomy: trace fossils: behavior categories Cucumaria. See Echinodermata: Holothuroidea Cuffey, Roger J . , 9 1 , 95, 296, 302 cultrata, Ambonychia. See Mollusca: Pelecypoda Cupulocrinus. See Echinodermata: Crinoidea: C l a d i d a Curacao, Netherlands Antilles, pl. 3

curtus, Merocrinus. See

Echinodermata: Crinoidea: C l a d i d a

curvatum, Gorbyoceras. See Mollusca: Cephalopoda cuttlefish. See Mollusca: Cephalopoda Cuvier, Ceorges, 61 Cyanobacteria ("blue-green 'algae'"), 67

Cirvanella, 67 stromatolites, 67, 292 Cyanoplryta. See Algae

Cyathochitina, See: chitinozoans Cyathophylloides. See C n i d a r i a : Anthozoa: Rugosa cycles, cyclicity, 52-53, 55-57, 5 8 - 5 9 coarsening-upward cycle, 59 decameter-scale cycles, 59

eustatic sea-level c h a n g e , 55, 285 fining-upward cycle, fining-upward packet, 59 m e g a c y c l e , 55, 59, 288 meter-scale cycles, 55, 59, 61 Milankovitch Cycles, 64, 288 shoaling-upward cycles, 53 storm cycles, 55, 56, 292 Cycloconcha. See Mollusca: Pelecypoda Cyclocrinites, See: Algae: Chlorophyta: Dasycladaceae Cyclocystoidea. See Echinodermata Cyclonema. See Mollusca: Gastropoda Cyclora. See Mollusca: Gastropoda Cyclostomata. See Ectoprocta Cylindrocoelia. See Algae Cymaronora. See Mollusca: Pelecypoda C y n t l u a n a Formation, 123 Cyrroceras. See Mollusca: C e p h a l o p o d a Cyrtodontula. See Mollusca: Pelecypoda Cyrtolites. See Mollusca: Monoplacophora

Cystaster. See Echinodermata: Edrioasteroidea ( A s t o i d c a . Sec

Echinodermata

cystoids. See Echinodermata

Dalmanella. See

Brachiopoda: Articulata Dalve, Elizabeth A., 48, 230, 238, 263 d a m a g e (in life). See injuries Dana, James Dwight, 275 Darwin, C h a r l e s , 61

darwini, Cyclocrinites.

See Algae: Chlorophyta: Dasycladaceae

darwini, Pasceolus. See Algae: Cyclocrinites

Chlorophyta: Dasycladaceae:

Dasycladaceae. See Algae: Chlorophyta dating. See time Dattilo, B e n j a m i n E, 59, 60, 6 1 , 113, 231, 232, 308, 312 Davis, Richard Arnold, ix, 14, 89, 212, 229, 249, 255, 298, 304,320 D C A . See techniques: detrended correspondence analysis ( D C A) Dearborn County, Indiana, pl. 3, pl. 5 death assemblage, 7, 8, 230, 283, 287. See also life assemblage; taphonomy decadactylus, Glyptocrinus. See Echinodermata: C r i n o i d e a : Camerata decameter-scale cycles. See cycles, cyclicity Deceptrix. See Mollusca: Pelecypoda

Decoroproetus. See Arthropoda: Trilobita

decorus, Neocrinus. See

Echinodermata: Crinoidea deformation (in life), deformities. See pathology deformation (post-mortem). See taphonomy Deiss, C h a r l e s Frederick, 263

delicatum, Oncoceras. See Mollusca: Cephalopoda

demissa, Caritodens. See Mollusca: Pelecypoda demissa, Pterinea. See Mollusca: Pelecypoda: Caritodens demissa Dendroidea. See Hemichordata Dent, Ohio. See Ohio: Hamilton C o u n t y dentatum, Rhynchotrema. See Brachiopoda: Articulata dentition. See Brachiopoda deposit feeding. See food depositional sequences, 52-53, 55-57, 5 8 - 5 9

Index

329

desiderata, Astraspis. See

Chordata: Vertebrata: Ag n ath a detrended correspondence analysis (DCA). See techniques Devonian, xix, 4, 9, 62, 72, 79, 144, 151, 186, 191, 239, 251 diagnosis. See taxonomy: L i n n a e a n Hierarchy Dickhaut, H. E., 263

dickhauti, Lepidolites. See Algae: Chloropbyta: Dasycladaceae Diekmeyer, Sharon C. St. Louis, 232, 307

Diestoceras. See Mollusca: Cephalopoda Dill Robert F, 54 Dillsboro Formation, 44, 51 d i m o r p h i s m , sexual, 153, 283, 290 dinoflagellates. See Algae: Pyrrophyta "dioramas," pl. 13

Dystactophycus. See incertae sedis earthquakes, 7, 61, 291. See also seismites earwigs. See Arthropoda: Insecta

eatoni, Triarthrus. See Arthropoda:

Trilobita eccentricity (in Earth's orbit). See cycles: Milankovitch Cycles ecdysis, 6 , 9 2 , 9 3 , 117, 147, 148 Echinodermata, 24, 26, 144, 145, 1 6 6 - 1 9 2 , 2 1 9 , 2 2 3 , 233, 237,241,245,246,260,269 Asteroidea, 24, 167, 168, 169, 182,183, 184, 185, 186, 188, 189-190, 208, 210, 235, 241, 247, 248, 259, 260, 265, 266, 274 Asteriacites (See taphonomy: trace fossils)

Fromia: F. nodosa, pl. 9

diplobathrid camerates. See Echinodermata: Crinoidea: Camerata Diplocraterion. See taphonomy: trace fossils Diplopoda. See Arthropoda disarticulation of hard parts. See taphonomy Disparida. See Echinodermata: C r in o idea divaricans, Streptelasma. See C n i d a r i a : Anthozoa: Rugosa diversity, species diversity, taxonomic diversity, 2, 3, 4, 57, 6 1 , 69, 103, 113, 115, 118, 120, 130, 131, 137, 147, 1 6 1 , 1 6 3 , 1 6 7 , 204, 232, 234, 239, 241, 283 d'Orbigny, Alcide, 85, 9 0 , 9 5 , 271 dolomite, 3, 53, 217, 221, 222, 283 d o m i c h n i a . See taphonomy: trace fossils: behavior categories Donovan, Stephen K., 186 Dorfeuille, Joseph, 263, 272, 274, 276 "doughnuts." Sec "W'cichold doughnuts" Drake, Daniel, 16, 2 6 3 - 2 6 4 , 268, 273, 276 Drakes Formation, 44, 138 Preachersville M e m b e r , 214 Rowland M e m b e r , 127, 214

Goniasteridae, 189

Hudsonaster, 189; H. simp/ex, 185, 274

Lanthanaster, 189; L. intermedius, 184

Ophidiasteridae, 189

Palaeaster: P. simplex (See Hudsonaster simplex) Palaeasterina: P. approximata, 259, 265; P. speciosa, 260,266

Petraster: P. speciosus, 184, 189 Promopalaeaster, 189, 235 P. dyeri, 184 P. magnificus, 182, 183 P. speciosus, 182, 183 Salteraster, 189; S. grandis, 184 carpoids, 145, 168, 186, 275. See Stylophora C r i n o i d e a , 7, 23, 24, 2 8 , 4 3 , 56, 57, 93, 120, 121, 123, 137, 144, 1 6 2 , 1 6 6 , 167, 168, 169-178, 180, 181-183, 186, 188, 212, 223, 224, 225, 226, 231, 232, 236, 237, 241, 243, 245, 246, 247, 248, 249, 251, 252, 253, 261, 270, 271, 285 C a m e r a t a , 178, 180

Drepanoistodus. See conodonts

diplobathrid camerates, 176, 177, 180

drilling. See borings Droser, M a r y L., 1,237, 321 Dry Dredgers, 255, 263, 275, 300, 302, 317 dubius, hichenocrinus. See Echinodermata: Crinoidea: Disparida

monobathrid camerates, 175, 176, 180 Clyptocrinus, 4 2 , 4 3 , 120, 178, 224, 2 3 2 , 2 4 0 , 241, 252

d u m b e l l s . See taphonomy: trace fossils:

Diplocraterion

duncanae, Gorbyoceras. See Mollusca: Cephalopoda

dunni, Narthecoceras. See

Mollusca: C e p h a l o p o d a durophagy. See ecologic associations: predation: Durrell, Richard H., ix Dury, C h a r l e s , 105 Dury, Ralph, 105

duseri, Treptoceras. See Mollusca: Cephalopoda: Actinoceratoidea Dyche, D. T. D., 28, 277 dychei, Prioniodus. See conodonts Dyer, C. B., 14, 2 3 - 2 4 , 29, 31, 3 7 , 4 0 , 80, 82, 184, 185, 208, 210, 260, 261, 265, 266, 272, 274 dyeri, Allocotkhnus. See taphonomy, preservation: trace fossils dyeri, Clyptocrinus. See Echinodermata: C r i n o i d e a : C a m erata: Pycnocrinus dyeri, Promopalaeaster. See Echinodermata: Asteroidea dyeri, Pycnocrinus. See Echinodermata: C r i n o i d e a : Camerata Dysracrocr/nus. See Echinodermata: Crinoidea: Disparida

330

Index

Caurocrinus: G. nealli, 176, 177, 180

G. annularis, 272 G. decadactylus, 42, 171, 172, 175, 180 G. dyeri. See Pycnocrinus dyeri G. fornshelli, 176, 180 G. pattersoni, 272 G. richardsoni, 273, 276 Pycnocrinus, 2 4 , 4 3 , 120, 123, 178, 180,224 P. dyeri, 24, 3 7 , 4 0 , 4 2 , 4 3 , 1 7 5 , 176 Xenocrinus, 180, 240, 241 X. baeri, 176, 177, 228, 229 C l a d i d a , 178

Cupulocrinus, 178 C. polydactylus, pl. 11 Merocrinus, 178 M . curtus, 172, 176,177 Plicodendrocrinus, 178 P. casei, pl. 11 Comactinia, 166 Comasterida, 166 Disparida, 173, 174, 178, 186

Anomalocrinus, 178, 252 A. incurvus, 172, 174

Cincinnaticrinus,

113, 178, 182, 186, 226, 2 3 2 , 2 4 0 ,

241,253

C. pentagonus, 173 C. varihrachialus, 172, 173 Dystactocrinus: D. constrictus, 173, 183 Ectenocrinus, 113, 178, 226, 2 3 2 , 2 4 0 , 241, 253 E. geniculatus, 173 E. simplex, 172, 173, PL 10 locrinus, 178, 224 /. oehanus, 271 /. subcrassus, 110, 142, 170, 172, 174, PL 10 Lichenocrinus, 178 L. crateriformis, 172 L. dubius, 186 L. milleri, 172 Metacrinus, 182 Neocrinus: N. decorus, pl. 9 Pontiometra: P. andersoni, PL 9 Cyclocystoidea, 169, 190-191, 247

7.ygocycloides: Z. magnus, 185 cystoids, 237. See Rhombifera Echinoidea, 167, 238, 248 sand dollars, 167 sea urchins, 167, 168, 169, 237

Strongylocentrotus: S. franciscanus, PL 9 Edrioasteroidea, 5 6 , 6 0 , 114, 169, 182, 186, 188-189, 190, 241,243,247,250,251

Carneyella, 178, 179, 224 C. pilea, 180, 18J Cincinnatidiscus, 40, 41 Cysfasfer: C . stellatus, 180,181 lsorophus, 189, 224 J. cincinnatiensis, 40, J28, 180, 181, 189 Streptaster, 110, 188, 2 2 4 , 2 5 0 S.vorticellatus, 110, 180,181 Eocrinoidea, 168 Holothuroidea, 167, 248

Cucumaria: C. miniata, PL 9 Ophiuroidea, 6, 167, 168, 169, 189, 1 9 0 , 2 2 8 , 229, 247, 248

Ophiothrix, PL 9 Protasterina: P. flexuosa, 190 Taeniaster, 190 T. spinosus, 185, 190,228, 229 Rhombifera, rhombiferan "cystoids," 169, 186, 247

Cheirocystis: C. fultonensis, 178, 179, 186 Lepadocystis: L. moorei, 178, 179, 186 Stylophora, 169, 186, 191-192, 247. See also "carpoids"

Enoploura, 26, 145, 191, 192, 270, 275 {See also machaeridians)

E. balanoides, 191 E. popei, 186, 187, 191 Echinoidea. See Echinodermata ecologic associations. See chapters on individual groups of animals aegism, 243, 244, 245, 280, 282, 288, 289, 290 c o m m e n s a l i s m , 95, 123, 144, 189, 229, 234, 236, 243, 244, 245, 246, 280, 282, 286, 288, 289, 290, 292 competition, competitive interactions, 182, 234, 236, 237, 282 epizoa, epizoism, 9 3 , 9 4 , 9 6 , 103, 130, 182, 236, 242, 243,

244, 245, 284. See also encrustation

facultative, 285, 288 inquiline, i n q u i l i n i s m , 286. See also incolent m u t u a l i s m , 234, 236, 288, 290, 292 obligate, 123, 285, 288 parasite, parasitism, 8 6 , 94, 123, 182, 223, 229, 234, 236, 243, 246, 280, 281, 282, 285, 286, 288, 289, 290, 292 predation, 7 , 9 2 , 9 3 , 95, 123, 124, 125, 131, 132, 137, 143, 147, 151, 153, 154, 155, 161, 162, 182, 183, 186, 189,190, 205, 210, 229, 234, 2 3 5 - 2 3 6 , 241, 282,

283, 284, 285, 289, 290, 291, 293. See also boring durophagy, 131, 284 symbiosis, 210, 235, 282, 284, 288, 289, 290, 292 endosymbiont, 210, 284 ecologic succession, 56, 57, 115, 281, 292 ecology, ecologic needs. See also chapters on individual groups of a n i m a l s ; ecologic associations; environment autecology, 229, 280, 284, 292 ecosystems, x, 1, 67, 143, 222, 2 3 3 - 2 4 8 , 284 c o m m u n i t i e s , 4, 5 , 5 6 , 57, 169, 182,218, 230, 231, 237, 241 consumers, 234, 282, 285, 290 food c h a i n , food web, 67, 6 8 , 234, 282, 285, 290 guilds, 237-239, 247, 248, 286 m e g a g u i l d s , 237, 238 populations, 188, 189 primary producers, 67, 234, 247, 248, 285, 290 tiering, 182 paleoecology, 6 1 , 136, 188, 189, 230, 231 Economy M e m b e r . See Latonia Formation ecosystems. See ecology

Ectenocrinus. See

Echinodermata: Crinoidea: Disparida Ectoprocta, 2 1 , 24, 30, 31, 33, 34, 35, 3 6 , 4 0 , 52, 57, 72, 75, 7 7 , 7 9 , 8 5 - 9 7 , 9 9 , 107, 110, 111, 113, 114, 115, 120, 125, 127, 128, 130, 131, 137, 139, 1 4 4 , 1 5 6 , 162, 178, 180, 182, 188, 196, 2 0 6 , 2 0 9 , 210, 219, 221, 223, 224, 225, 231, 232, 233, 237, 240, 241, 242, 243, 244, 245, 247, 248, 250, 251, 252, 253, 260, 265, 270, 271, 272, 273, 275, 281, 285, 287, 290,293 "bryoherms," 91 "Weichold doughnut," 88, 89, 9 3 - 9 4 , 275 Cheilostomata, 237, PL 3

Cryptostomata: Escharopora, 87, 88

Ctenostomata: Ropalonaria: R. venosa, 87, 88 Cyclostomata, 107, 110 Cystoporata

Ceramopora; C. (?) beani, 260; C. nicholsoni, 271 Constellaria, 40, 87, 94 C. constellata, 275 C. florida, 87, 88 Fistulipora: F. oweni, 272 Heteropora: H. pacifica, PL 3 Trepostomata, 79, 87, 88, 8 9 , 9 0 , 237, 250

Heterotrypa, 91 H.frondosa, 9 0 , 9 5 , 271 Eeptotrypa: L. clavacoidea, 120, 121 Monticulipora 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), Spatiopora, 87, 88, 94

265, 270

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, ecologic 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 Arthropoda: Crustacea: Ostracoda

Eoleperditia. See

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 Florida, pl. 9

Echinodermata: Ophiuroidea

florida, Constellaria. See

Ectoprocta: Cvstoporata Flower, Rousseau H., 43, 133, 134, 138, 295 floweri, Fascifodina. See taphonomy: trace fossils floweri, Palaeoscia. See C n i d a r i a : Hydrozoa: Siphonophorida: Porpitidae; taphonomy: trace fossils F M N H . See Field M u s e u m of Natural History fodiniclmia. See taphonomy: trace fossils: behavior categories Foerste, August F., 18, 2 9 , 4 8 , 7 7 , 7 8 , 109, 149, 150, 151,153, 162, 207, 229, 264, 270, 271 Foerstephyllum. See C n i d a r i a : Anthozoa: Tabulata food supply. See food food chain, food web. See ecology: ecosystems food, 4 , 6 7 , 6 8 , 7 1 , 74, 85, 8 7 , 9 2 , 101, 123, 151, 155, 160, 168, 170, 172, 180, 188, 191-192, 198, 211, 229, 234, 235, 237-238, 239, 253, 280, 282, 283, 285, 286,

287, 290. See also chapters on individual groups of a n i m a l s ; ecologic associations: commensalism; ecologic associations: predation; ecology: ecosystems coprophagy, 121, 123,282 deposit feeding, 123, 131, 140, 163, 190, 204, 2 0 5 , 2 1 1 , 2 2 4 , 283, 285, 292 filter feeder, filter feeding, 7 1 , 72, 87, 101, 127, 131, 144, 155, 168, 172, 1 8 8 , 2 8 3 , 2 8 5 , 2 9 2 scavenging, 7, 151, 205, 223, 224, 225 suspension feeding, 21, 74, 123, 139, 163, 170, 182, 186, 190, 196, 205, 206, 211, 224, 237, 239, 283, 285, 292 Ford, John P., 21, 59 formation. See lithostratigraphic units formosa, Conularia. See C n i d a r i a : Scyphozoa: C o n u l a r i i d a fornshelli, Clyptocrinus. See Echinodermata: C r i n o i d e a : Camerata Fort Ancient M e m b e r See Waynesville Formation Fortey, Richard A., 1, 151, 155, 161, 303, 316 fossil succession. See biotal succession Foster, W i l l i a m , 3 1 , 2 6 4 Fox, W i l l i a m T., 231

franciscanus, Strongylocentrotus. See

Echinodermata: Echinoidea Franklin County, Indiana, 105, 107, 157, 184, 196, pl. 6, pl. 10 Franks' M u s e u m , 265 Frey, Robert C, 60, 130, 131, 136, 137, 138, 297, 307 Fromia. See Echinodermata: Asteroidea

frondosa, Heterotrypa. See Ectoprocta: Trepostomata fucoids. See taphonomy: trace fossils Fulton Beds, Fulton Shale, 144, 186. See also Kope Formation Fulton, Robert, 265 fultonensis, Cheirocystis. See Echinodermata: Rhombifera fungi, 242 Fusispira. See Mollusca: Gastropoda: Neogastropoda:

Subulites G a h n , Forest J . , 183, 297 G a m a c h i a n Stage, 214, 240 Garrard Formation, 214 Gastropoda. See Mollusca Gault, W i l l i a m , 265

Caurocrinus. See

Echinodermata: C r i n o i d e a : C a m e r a t a G e e , Henry, 191 generic n a m e . See taxonomy: nomenclature: Binomial System of Nomenclature genkulatus, Ectenocrinus. See Echinodermata: C r i n o i d e a : Disparida Ceniculograptus. See Hemichordata: Graptoloidea genus, genera. See taxonomy: I.innaean Hierarchy geologic time-scale, xix geologic time-units, 4 5 - 4 6 Geological Society of A m e r i c a , 5, 11, 30, 32, 36, 54, 59, 133, 257, 2 7 1 , 2 7 4 Penrose M e d a l , 30, 32 geopetal structures, 124 George Washington University (formerly C o l u m b i a n University), 22, 35 Gibson, Bruce, 105, 176 Gibson, Charlotte, 105, 176

gigantea, Anomalodonta. See

Mollusca: Pelecypoda gigas, Isotelus. See Arthropoda: Trilobita Gilbert Formation, 214 Girvanella. See Cyanobacteria

Clyptocrinus. See Echinodermata: Crinoidea: Glyptorthis. See Brachiopoda: Articulata

Camerata

G o l d m a n , Daniel, 195 G o n d w a n a . See paleogeography Goniasteridae. See Echinodermata: Asteroidea

Gorbyoceras. See Mollusca: Cephalopoda Grabau, A m a d e u s W., 18

gracile striatulum, Cyclonema. See Mollusca: Gastropoda gracile, Cyclonema. See Mollusca: Gastropoda gracile, Ordovicium. See acritarchs graded beds, 60 G r a h a m , George, 265 grainstone, 50, 55, 59, 114, 2 8 6 , 289 Grand Avenue Member. See Kope Formation grandis, Aphelognathus. See conodonts grandis, Salteraster. See Echinodermata: Asteroidea Grant Lake Formation, Limestone, 44, 79, 9 0 , 114, 123, 250 Corryville Member. See Corrvvillc Formation, M e m b e r

granulosa, Flexicalymene. See Arthropoda: Trilobita Graptoloidea, graptolites. See Hemichordata Arthropoda: Trilobita: C a l y m e n i d a Great B a h a m a Bank. See B a h a m a Platform Great Barrier Reef, Australia, 9 1 , 241, pl. 3. See also bioherm Great Limestone Deposite, 47 Grewingk, C. C. A., 41 Grewingkia. See C n i d a r i a : Anthozoa: Rugosa Grinndl, George Bird, 265 group. See lithostratigraphic units growth lines, 75, 77, 117, 120, 2 8 6 guilds. See ecology: ecosystems Gunderson, G. O., 82, 309 Gurley, W. F. E., 25, 157,313 gutter casts, 60

Gravicalymene. See

H a i m e , Jules, 265, 270 Haines, M. P., Mrs., 265 Hall, J . H . , 2 5 9 , 265 Hall, James (of C i n c i n n a t i ) , 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, 173, 175, 176, 180, 183, 195, 207, 210, 229, 262, 265-266,272,276,277 Hall, John W., Jr., 2 5 9 , 2 6 5 , 2 6 6 11.ill.mi. Anthony 3, 4 halli, Alloliehas. See Arthropoda: Trilobita: Lichida halli, Amphilichas. See Arthropoda: Trilobita: Lichida:

chas halli

12 Alloli-

Hallopora.

See Ectoprocta: Trepostomata: Parvohallopora 1 lamilton Comity, Ohio, 10, 8 9 , 9 0 , 106, 142, 151, 155, 156, 173, 180, 186, 277, pl. 10 'Trammel Eossil Park, Sharonville, 12, 124, 257 I l a m m o n d , W. E., 266 Hand, Greg, 306 hardgrounds, 6 0 , 6 4 , 105, 124, 125, 178, 188, 203, 2 0 6 , 211, 219,250, 2 5 1 , 2 8 6 I larding Sandstone, 199 Harper, George W., 17, 18, 26, 27, 32, 3 3 - 3 4 Harris, Frank W., 5 6 , 5 7 , 114, 115, 231 Harris, G. D., 25 Harris, I., 259, 266 I larvard University, xv, 18, 23, 32, 260, 273, 274 M u s e u m of Comparative /oology ( M C Z ) , xv, 18, 23, 32, 2 6 0 , 2 7 3 , 2 7 4 , pl. 13 Harvey, J . W „ 260, 266 Haucr, Kendall, 311 Hay, Helen B„ 44, 53, 55, 56, 57 Hayden, F. V., 266, 268 Uebertella, S e e : Brachiopoda: Articulata I ledcen, Stanley. i\ I [eimbrock, W i l l i a m , 67 Helcionopsis. See Mollusca: Monoplacophora Hemichordata, 169, 196 Dendroidea, 195

Muslivograptus, 196 graptolites, Craptoloidea, 52, 63, 161, J94, 195-196, 240, 241,247,251

Amheimograptus: A. anacanlhus. 196 Climacograptus. See Geniculograptw V.enkuhgmptus, 196; C. typicalis. 194, 195 Orthograptus: (). quadrimucronatus, 196 Pterobranchia, 196; Rhahdopleura, 196 I lendy, Austin, xiv I leran, M a g g i e , xiv

Hercochitina. See chitinozoans I Icrrlinger, Andrew, 267 Herzer, Henry, 28, 30, 267 Heteropora. See Ectoprocta

I lolothuroidca. See Echinodermata holotype. See taxonomy: type specimens I loltedahlina. See Brachiopoda: Articulata homeomorphy. See organic evolution: convergent evolution horseshoe crabs. See Arthropoda: Chelicerata: Xiphosurida Hosea, L. M., 24, 267 host, 77, 86, 87, 88, 9 3 , 9 4 , 103, 123, 144, 182, 189, 210, 234, 235, 236, 242, 243, 244, 245, 246, 282, 285, 286, 288, 289 Hovey M u s e u m , Wabash College. See Crawfordsville, Indiana Howe, A. J . , 267 I [udson River C r o u p , 266, 269, 277 I ludsonaster. See Echinodermata: Asteroidea I luffman D a m . near Davton, ()hio, 153 Hughes, Nigel C , 82, 153

humerosum. See Mollusca: Gastropoda: Cyclonema

humerosum, Cyclonema. See

Mollusca: Gastropoda

Hunda, Brenda, 151 Hunter, W. H. H., 267

huronensis, Labechia. See

Porifera hurricanes. See storms I lutchinson, G. Evelyn, 229 Huxley, T. H., 267 hydroids. See C n i d a r i a : Hydrozoa I Ivdrozoa. hvdrozoans. See C Inidaria I lyinan, L. H., 167 Hyolithes. See hyolithids hyolithids, 145, 247. See also Mollusca: Calyptomatida

Coleolus, 145 Hyolithes. 145 hyporeliefs. See taphonomy: trace fossils lapetus O c e a n , Sea. See paleogeography See Arthropoda: Trilobita: Phacopida Ice Age, ice cap, ice sheet, ix, 10, 53, 216, 217, 240, 269 ichnofossils. See taphonomy, preservation: trace fossils ichnogenus, ichnogenera. See taphonomy: trace fossils ichnospeeies. See taphonomy: trace fossils ICZN. See International C o d e of Zoological Nomenclature Illinois Geological Survey, 31 impedichnia. See taphonomy: trace fossils: behavior categories

icarus, Ceraurinus.

inaequahile, Cameroceras. See Mollusca: Cephalopoda:

Heterotrypa. See Ectoprocta: Trepostomata High Bridge C r o u p (middle Ordovician], 67. 139, 164 Tyrone I .iinestone. 139 Highland County. Ohio, 151, 153 1 Mil Quarry Beds, 47, 225. See also Fairview Formation Hill, H. H., 267 Himalayas, 3 Hints, Olle, 145 Hirnantian Stage, 63, pl. 2

lunula, Conochitina. See chitinozoans Hiscoheccus. See Brachiopoda: Articulata Hitz bed, 214 Hogstrom, Anette E. S., 145, 307

334

Holbrook, R. H., 267 boles (in shells). See taphonomy: trace fossils: borings Holland, Steven M., 53, 55, 5 9 , 6 3 , 113, 120, 130, 204, 2 1 5 - 2 2 6 , 229, 231, 232, 233, 241, 255, 312, pl.

Index

Endoceratoidea Inarticulata. See Brachiopoda incertae sedis, 211-212, 286. See

Blastophycus, 212 Dystactophycus, 212

also Palaeoscia

incolent, 286. See also ecologic associations Echinodermata: Crinoidea:

incurvus, Anomalocrinus. See Disparida index fossils, 195, 255 Indiana

Bedford, 10 Crawfordsville, 23, 2 6 1 , 2 7 0 Hovey M u s e u m , Wabash College, 23

Dearborn County, pl. 3, pl. 5 Franklin County, 105, 107, 157, 184, 196, pl. 6, pl. 10 Brookville, 184, 196 Indiana Geological Survey, 51, 214, 255, 263 Jefferson County, 111 Madison, 7 7 , 7 9 , 2 2 1 , 2 6 6 , 2 6 9 Richmond, 47, 120, 265, 270, 272 W a y n e County, 67, 75, 228 infauna, infaunal, 7, 131, 139, 153, 204, 205, 237, 238, 239, 247, 248, 284, 286 injuries, d a m a g e (in life), 137, 183, 186, 235, 236. See also ecologic associations: predation inquiline, inquilinism. See ecologic associations insculpta, Glyptorthis. See Brachiopoda: Articulata Insecta. See Arthropoda intermedins, Lanthanaster. See Echinodermata: Asteroidea International C o d e of Zoological Nomenclature ( I C Z N ) , xi International Geological Correlation Programme, 63, 255 International Stratigraphic C o d e . See stratigraphy interspecific interactions. See ecologic associations locrinus. See Echinodermata: Crinoidea: Disparida Ischadites. See Algae: Chlorophyta: Dasycladaceae Ischyrodonta. See Mollusca: Pelecypoda isochronous surfaces. See time Isopoda. See Arthropoda: Crustacea

Isorophus. See Echinodermata: Edrioasteroidea

Isorthoceras. See

Mollusca: Cephalopoda isotopes, 62, 290. See also time: absolute age: radiometric dating Jacobson, Stephen R., 69 James Book Store, 14, 17, 22, 31 James, Joseph A., 17 James, Joseph R, 2 2 - 2 3 , 29, 203, 207, 210 James, U. P., 14, 17-22, 2 3 , 2 8 , 29, 31, 209, 210, 260, 261, 264, 2 7 1 , 2 7 7 jamesi, Lepidocoleus. See Machaeridia Jefferson County, Indiana, 111 jellyfish. See C n i d a r i a : medusa Jennette, David C, 55, 59, 60 Johnson, T h o m a s T., 229, pl. 7

Journal of Geology, 55 journal of the Cincinnati Society of Natural History. See C i n cinnati Society of Natural History junior synonym. See taxonomy: nomenclature: synonym; taxonomy: nomenclature: synonymy Jurassic, xix, 4, 239 K-bentonite, K-bentonite beds, K-bentonite "zones," 62, 63, 163, 287 Bear Creek K-bentonite "zone," 62 Millbrig K-bentonite bed, 62 Seneca County, Ohio, 6 2 - 6 3 Kankakee Arch, Kankakee Branch of the C i n c i n n a t i Arch, 9.

See also Cincinnati Arch Kaplan, Peter, 124, 125 Kemper, Willis, 268 Kenton County, Kentucky, 67, 87, 90, 110, 123, 209 Kentucky Boone County, 87, 110, 111, 142, 180 Bracken County, 178, 209

C a m p b e l l County, 59, 109, 203, 209, pl. 5 Cold Spring, 174 Covington, 19, 28, 127, 142, 256, 263, 266, 274 F l e m i n g County, 72 Kenton County, 67, 87, 9 0 , 110, 123, 209 Kentucky Geological Survey, 10, 51, 214, 256 Lexington, 40, 67, 123, 272 Transylvania C o l l e g e , University, 40, 272 Madison County, 77 Marion County, 120 Maysville, 11, 5 9 , 7 9 , 9 1 , 180 Nelson County, 78 Trimble County, 124, 129 Bedford, 129

kentuckyensis, Rhytiodentalium. See

Mollusca: Scaphopoda Kesling, Robert V., 178 Kiessling, Wolfgang, pl. 2 kingdom. See taxonomy: L i n n a e a n Hierarchy Kjellesvig-Waering, Erik N., 158, 159, 162, 300 Knell, Simon J . , 265 Kope Formation, II, 19, 21, 44, 49, 51, 52, 53, 55, 56, 57, 59, 6 0 , 6 1 , 6 7 , 6 8 , 6 9 , 106, 113, 120, 123, 127, 130, 142, 144, 149, 151, 155, 156, 160, 173, 176, 178, 180, 186, 190, 196, 203, 208, 209, 211,214, 219,

225, 231, 232, 253, pl. 5. See also Eden Shale; Latonia Formation Fulton M e m b e r , 144, 186 Grand Avenue M e m b e r , 44 W e s s e l m a n Tongue, 4 4 Kozlowski, Roman, 196 Kritsky, G e n e , 267

Lahechia. See Porifera laelia, Philhedra. See Brachiopoda: Inarticulata lag deposit, 56, 114,287 land plants. See Plantae, plants Lanthanaster. See Echinodermata: Asteroidea lata, Cyclonema hilix. See Mollusca: Gastropoda:

hilix

Cyclonema

Lathrop, J. P., 268

laticosta, Platystrophia. See Brachiopoda Latonia Formation, 4 4 , 51. See also Eden Formation; Kope Formation Economy M e m b e r , Economy beds, 44, 144 McMicken Member, 44 Southgate M e m b e r , 4 4 latus, Isotelus. See Arthropoda: Trilobita Laurentia. See paleogeography laurentiensis, Sanctum. See taphonomy: trace fossils: borings layer-cake geology, 45 Lebanon Beds, 47; renamed Richmond, 47 L e b a n o n , Warren County, Ohio, 28, 176, 260, 267, 271, 273, 274, 277 Lebensspuren. See taphonomy, preservation: trace fossils Lepadocystis. See Echinodermata: Rhombifcra I.eperditicopida. See Arthropoda: C r u s t a c e a : Ostracoda Lepidocoleus. See M a c h a e r i d i a Lepidocyclus. See Brachiopoda: Articulata Lepidolites. See Algae: Chlorophyta: Dasycladaceae

Leptaena. See Brachiopoda: Articulata

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 : Licrophycus. See taphonomy: trace fossils

Disparida

life assemblage, 8, 230, 283, 287. See also ecology: ecosystems: 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 Bellevue 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: Flexicalymene 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 Michigan, University of. See University of M i c h i g a n Mickleborough, John, 26, 27, 153, 155, 262, 276, 313 micraulaxum, Multiplicisphaeridium, S e e : acritarchs microscopy. See techniques: microscopy i n f r a s t r u c t u r e , 6, 6 8 , 7 2 , 144, 166, 168, 192, 292 Middle (Eden) Shales, 47 Milankov itch Cycles. See cycles, cyclicity Millbrig K-bentonite bed. See K-bentonite, K-bentonite beds, K-bentonite "zones"

millepunctata, Trematis. See Brachiopoda: Inarticulata Miller, Arnold I., 113, 231, 232, 305, 308, 312 Miller, C. A., 269 Miller, S. A., 14, 17, 23, 2 4 - 2 5 , 28, 2 9 , 4 2 , 6 8 , 80, 82, 126, 127, 134, 137, 142, 144, 157, 160, 161, 176, 182, 184, 185, 203, 204, 205, 2 0 6 , 208, 209, 210, 259, 260, 261, 265, 266, 267, 270, 272, 274, 275 milleranus, Ceraurus. See Arthropoda: Trilobita: Phacopida milleri, Eichenocrinus. See Echinodermata: C r i n o i d e a : Disparida

milleri, Technophorus. See

Mollusca: Rostroconchia millipedes. See Arthropoda: Diplopoda Milne-Edwards, Henri, 265, 270

miniata, Cucumaria. See

Echinodermata: Holothuroidea Minnesota Geological Survey, 31 M i n t z , Leigh W., 178, 310 Miocene, xix, 4 Misener, John, 270 Misener, S. R., 270 miseneri, Conularia. See C n i d a r i a : Scyphozoa: C o n u l a r i i d a mode of life, 5,7, 52, 74, 160, 169, 188. See also chapters on individual groups of a n i m a l s modesta, Isygospira. See Brachiopoda: Articulata modiolaris, Modiolopsis. See Mollusca: Pelecypoda Modiolopsis. See Mollusca: Pelecypoda Mohawkian Stage, 167 Mohr.Paul, 261,270 molds, composite. See taphonomy: molds molecular clocks, 62. See also organic evolution Mollusca, 26, 3 1 , 8 7 , 9 3 , 9 9 , 101, 116-140, 144, 145, 162, 204, 206, 219, 223, 224, 225, 231, 233, 241, 244, 245, 259, 260, 2 6 1 , 2 8 2 , 2 8 8 , 2 9 0 , 2 9 1 anatomy and morphology: tentacles, 119, 131, 132, 138, 252 Aplacophora, 119,280 Calyptomatida, 145. See hyolithids Cephalopoda, 29, 43, 8 7 , 8 8 , 9 3 , 9 4 , 9 5 , 9 6 , 101, 118, 119, 131-138, 139, 144, 154, 196, 205, 220, 223, 225, 231,235, 238, 241, 247, 248, 249, 251, 252, 260, 264, 270, 280, 286 Actinoceratoidea, actinocerids, actinoceroids, 245

Beloitoceras: B. amoenum, 136, 137 Charactoceras, 138 C. haeri, 136, 137 cuttlefish, 131

Cyrtoceras: C. vallandighami, 275 Diestoceras, 138 D. eos, 136, 137 Endoceratoidca, endocerids, endoceroids, 134, 137, 154,245

Cameroceras, 137, 225 C. inaequabile, 134, 135, 137

Endoceras, 134 Corbyoceras, 138 G. curvatum, 137, 138 G. duncanae, 134, 135 lsorthoceras, 137 Manitoulinoceras M. tenuiseptum, 137 M. williamsae, 137 Narthecoceras: N. dunni, 138 nautiloid cephalopods, 60, 87, 88, 8 9 , 9 0 , 94, 132-139, 2 0 5 , 2 2 8 , 229, 235, 236, 238, 241, 244, 245, 249, 2 5 1 , 2 5 2 , 264, pl. 8 (includes Actinoceratoidea, Endoceratoidea, and Nautiloidea) Nautiloidea, 245 Ascocerida, 137

Nautilus, 95, 117, 131, 132, 134, 137, 139, 241. pl. 4 Orthocerida, 137 octopus, 117, 131, 139

Oncoceras, 137 O. delicatum, 137 Orthonyhyoceras, 43. See also Treptoceras Schuchertoceras O.obscurum, 137, 138 squid, 117, 118, 131, 137, 139,252 trace fossils, 138. See also taphonomy: trace fossils

Treptoceras, 43, 136, 137, 223,228, 229. See also Orthonyhyoceras T. duseri, 60, 134, 135, 136, 137, 138,228, 229

Zittelloceras: Z. russelli, 137; Z. williamsae, 137

chitons. See Polyplacophora C r i c o c o n a r i d a , 145. See Tentaculitoidea Gastropoda, 7, 24, 32, 101, 116, 117, 118, 119-125, 127, 129, 137, 139, 182, 223, 225, 231, 235, 237, 238, 240, 241, 243, 244, 246, 247, 248, 282

Archaeogastropoda, 123 (See also Cyclonema; Naticonema) abalones, 119, 123 limpets, 119, 123, 139 periwinkles, 123 bellerophontids, 120

Cyclonema, 120, 123, 144, 176, 223, 252 C. bilixlata, 122, 123 C. gracile striatulum, 122, 123 C. humerosum, 120, 122, 123 C. sublaeve, 122, 123 C. varicosum, 122, 123 Cyclora, 223 Liospira, 225 Lophospira, 223, 225 Loxoplocus: L. bowdeni. See Paupospira Naticonema, 120 Neogastropoda, 123 cones, 123 murexes, m u r i c e s , 123

Subulites (Fusispira), 122, 123 whelks, 123

Paupospira P. bowdeni, 125, 127 P. moorei, 122, 123, 127 P. tropidophora, 127 Index

337

platyceratids, 120 Ptcropoda, 145

Salpingostoma: S. richmondensh, 120, 12 J slugs, sea slugs, 117, 119 Monoplacophora, 93, 119, 120, 125, 139, 223, 244, 247 Archinacella: A. area, 120, 121

Cyrtolites, 125, 139 C. ornatus, 120, 121 Helcionopsis: H. striata, 120, 121 Neopilina, 139 Sinuites, 225 S. cancellatus, 120, 121

Pelecypoda, 7, 92, 93, 98, 99, 100, 101, 112, 113,117, 118, 119, 123,125, 128-131, 139, 144, 157, 168, 189, 206, 210, 223, 225, 231, 235, 237, 238, 240, 241, 242, 244, 245, 251, 269, 276, 280, 281, 283, 284, 291,293

Ambonychia, 88, 89, 126, 129, 130, 131, 223, 225, 240, 241, 252

A. cultrata, 126 Anomalodonta: A. gigantea, 126 Caritodens, 128, 130, 223, 241, 251 C. demissa, 126 cockles, 129

Corallidomus: C. scobina, 128, 206 Ctenodonta, 129, 131 Cycloconcha, 129 Cymatonota, 129 C. typicalis, 40 Cyrtodontula, 129 Deceptrix, 129, 225 giant c l a m . See Tridacna Ischyrodonta: 1. truncata, 126, 131 l.ockeia: 1,. siliquaria, 209, 210 l.yrodesma, 129 Modiolopsis, 129, 130, 131, 224, 225, 251 M. modiolaris, 127 mussels, 22, 32, 33, 117, 129, 131, 261 mvtilids, 131

Opisthoptera, 129 O. casei, 126 oysters, 114, 117, 129, 131

Pseudocolpomya, 131 Pterinea, 129 P. demissa. See Caritodens demissa Rhytimya, 225

monobathrid camcratcs. Sec Echinodennata: Crinoidea: Camerata Monoplacophora. See Mollusca Montgomery County, Ohio, 72, 157 monticule. See Ectoprocta: anatomy and morphology

Monticulipora. See Ectoprocta: Trepostomata Moore, Richard B., 17, 270, 279 Moore, T i m , 38

moorei, Lepadocystis. See Echinodermata: Rhombifera moorei, Paupospira. See Mollusca: Castropoda Morris, Robert W., 123 Morrow, Warren C o u n t y . Ohio, 175 Mount Auburn M e m b e r , Formation, 21, 44, 53, 55, 59, 63,

68, 160,214, 223, 250. See also M c M i l l a n formation Mount I lope Member. See Fairview Formation mountain building, orogeny. See tectonic activity mudstone, 50, 217, 218, 219, 222, 224, 225 M l ' C M . Sec Miami I University, ()hio, ( h l n r d multipartitum, Petalichnus. See taphonomy: trace Multiplicisphaeridium. See acritarchs murexes, murices. See Mollusca: Castropoda: Neogastropoda mussels. See Mollusca: Pelecypoda m u t u a l i s m . See ecologic associations mvtilids. See Mollusca: Pelecypoda myzostomids. See Annelida nannoplankton, 239

Narthecoceras. See Mollusca: Cephalopoda

Naticonema. See

Mollusca: Gastropoda National Academy of Sciences, 30, 32, 274 National M u s e u m of Natural History ( U S N M ) , xv, 19, 21, 75, 120, 127, 128, 153, 173, 175, 176, 182, 184,

185, 266. See also Smithsonian Institution; United States National M u s e u m Natural History M u s e u m , London, xv, 23, 26 natural selection. See organic evolution nautiloid ccphalopod. See Mollusca: C e p h a l o p o d a Nautiloidea. See Mollusca: C e p h a l o p o d a Nautilus. See Mollusca: Cephalopoda: Nautiloidea nealli, Caurocrinus. See Echinodermata: Crinoidea: Camerata

neglecta, Plectorthis. See

Brachiopoda: Articulata nekton, nektonic, 7, 138, 198, 280, 288, 290 Nelson County, Kentucky, 78

Neocrinus. See Echinodermata: Crinoidea

scallops, 112, 113, 117, 129

spiny oysters. See Spondylus

Neogastropoda. See Mollusca: Castropoda

Spondylus, 131 Tridacna, 131 Vanuxemia: V. waynesvillensis, 305

Neopilina. See Mollusca: Monoplacophora Neostrabops. See Arthropoda: Aglaspida

Polyplacophora, 117, 118, 119, 139 chitons, 117, 139, 145, 248 Rostroconchia, 119, 127, 139, 247, 290

Technophorus, 139 T. faberi, 127 T. milleri, 127 Scaphopoda, 119, 139, 140

Rhytiodentalium: R. kentuckyensis, 140 tusk shells. See Scaphopoda molting. See ecdysis

338

fossils

Nereigenys. See

Annelida: scolecodonts Nestor, Robert, 153 New C a l e d o n i a , pl. 4 New England, 3, 215 New York, 3, 17, 23, 27, 35,47, 4 8 , 149, 155, 160, 257, 265, 2 6 6 , 2 7 1 , 2 7 6 , 293 New York Geological Survey, 23, 32 New York State M u s e u m , 180 Newberry, John Strong, 24, 270, 271, 276, 277 Newton, A . J . , 270 Nicholson, H. Alleyn, 75, 77, 270-271

Index

nicholsoni, Ceramopora. See Ectoprocta: Cystoporata Nickles, John M., 1 8 , 1 9 , 2 0 , 27, 30, 32-33, 3 4 , 4 7 , 52, 230, 259, 260, 261, 263, 265, 266, 267, 269, 270, 272, 275, 277 Nitecki, Matthew H„ 68 nodosa, Fromia. See Echinodermata: Asteroidea nomen duhium (pl., nomina dubia), 42, 288. See taxonomy: nomenclature nomen nudum (pl., nomina nuda), 42, 288. See taxonomy: nomenclature nomenclature. See taxonomy normal wave-base, fair-weather wave-base. See wave base North Atlantic Province. See faunal provinces North Bend Tongue. See Fairview Formation numerosum, Trachomatichnus. See taphonomy: trace fossils nutrition. See food

Nyctopora. See

C n i d a r i a : Anthozoa: Tabulata

O ' N e a l l J . K., 271 Obata, Tadahiro, 43,317 obligate association. See ecologic associations obrution deposits, 60, 288 obscurum, Schuchertoceras. See Mollusca: C e p h a l o p o d a occidentalis, Hebertella. See Brachiopoda: Articulata octopus. See Mollusca: Cephalopoda Odontopleurida, odontopleurids. See Arthropoda: Trilobita Oeh, George, 271

oehanus, locrinus. See

Echinodermata: Crinoidea: Disparida

Ohio Adams County, 75, 158, 162, 163, 216, 229, pl. 7 Brown County, 123, 128, 151 Butler County; 68, 75, 80, 87, 89, 107, 108, 126, 127,134, 137, 153, 156, 160, 176, 195, 2 2 9 Clermont County, 111, 126, 128, 153, 158, 163, 170, 186, 203, 207, 208, 209, 261 Bear Creek Quarry, 62 Stonelick Creek, 11, 254 Clinton County, 6 0 , 7 2 , 75, 128, 136, 142 Dent, 176,180 Hamilton County, 10, 8 9 , 9 0 , 106, 142, 151, 155, 156, 173, 1 8 0 , 1 8 6 , 277, PL 10 Highland County, 151, 153 Huffman Dam (near Dayton), 153 Lebanon, 28, 176, 260, 267, 271, 273, 274, 277 Montgomery County, 72, 157 Morrow, 175 Oxford, 105, 153, 155, 156, 256, 269, pl. 11 Point Pleasant, 225 Portsmouth, 10 Preble County, 77, 105, 107, 108, 109, 111, 123, 126, 137, 178, 184 Trammel Fossil Park, Sharonville, 12, 124, 257 Seneca County, 63 Warren County, 60, 105, 126, 136, 172 Waynesville, 120, 182, 184, 185, 256, 259, 266 W i l m i n g t o n , 260, 273, 276 Ohio Division of Geological Survey, 92, 256. See also Ohio Geological Survey Ohio Geological Survey, 23, 269, 276. See also Ohio Division of Geological Survey Ohio M e c h a n i c s Institute, C i n c i n n a t i , Ohio, 31, 262

Ohio State University, C o l u m b u s , Ohio, xv, 2, 51, 256, 271 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

ohioensis, Megalograptus. See Arthropoda: Chelicerata: Eurypterida Oldroyd, John David, 231 Oligocene, xix, 4

Oncoceras. See Mollusca: Cephalopoda oncolites. See Algae

onealli, Acidaspis. See

Arthropoda: Trilobita: Odontopleurida onealli, Parvohallopora. See Ectoprocta: Trepostomata Ophidiasteridae. See Echinodermata: Asteroidea Ophiothrix. See Echinodermata: Ophiuroidea Opisthoptera. See Mollusca: Pelecypoda Orbigny, Alcide d', 85, 90, 95, 271 orbital parameters. See cycles: Milankovitch C y c l e s order. See taxonomy: L i n n a e a n Hierarchy Ordovician: "Ordovician Biodiversification Event," 2, 289; "Ordovician Radiation," 1, 2, 289

ordovicicus, Amorphognathus. See conodonts Ordovicium. See acritarchs Oregonia Formation, M e m b e r , 44, 53, 56, 63,214, 223. See

also Arnheim Formation

oregonia, Oulodus. See

conodonts organic evolution, 1, 2, 39, 6 1 , 62, 67, 82, 105, 115, 118, 131, 134, 139, 163, 167, 195, 196, 229, 233, 237, 267,

282, 286, 287. See also durophagy; molecular clocks convergent evolution, 105, 282, 286 evolutionary relationships. See chapters on individual groups of a n i m a l s homeomorphy, 105, 282, 286 organic evolution by natural selection, 134 ornatus, Cyrtolites. See Mollusca: Monoplacophora ornatus, Protoscolex. See Annelida orogeny (mountain building). See tectonic activity Orthocerida. See Mollusca: C e p h a l o p o d a : Nautiloidea Orthograptus. See Hemichordata: Graptoloidea Orthonybyoceras. See Mollusca: C e p h a l o p o d a : Actinoceratoidea Brachiopoda: Articulata Orton Geological M u s e u m . See Ohio State Universitv Orton, Edward, 2 4 , 4 7 , 52, 268, 271, 276 Orton, Edward, Jr., 271

Orthorhynchula. See

Osgood, Richard G., Jr., 138, 203, 204, 205, 206, 208, 209, 210,211,212 Osteichthyes ("bony fish"). See Chordata: Vertebrata Ostracoda. See Arthropoda: Crustacea O S U . See Ohio State University: Orton Geological M u s e u m Oulodus. See conodonts Owen, David Dale, 261, 269, 272 oweni, Fistulipora. See Ectoprocta: Cystoporata O w e n s , R. M., 151, 155, 161, 305 Oxford, Butler County, Ohio, 105, 153, 155, 156, 256, 269,

pl. 11 oxygen content. See environment oysters. See Mollusca: Pelecypoda ()zarkian System. 30, 31

pacifica, Heteropora. See

Ectoprocta 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-

pentagonus, Cincinnaticrinus. See

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

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

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

Perroxesfes. See taphonomv: trace fossils: borings Phacopida, phacopids. See Arthropoda: Trilobita Phaeophyta. See Algae Phanerozoic Eon, 2 , 4 , 4 6 , 62, 239, 284, 289

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

Echinodermata: Crinoidea: Disparida pera, Petroxestes. See taphonomy: trace fossils: borings Pericharax. See Porifera

Petrocrania. See Brachiopoda: luarticulata

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

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

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

Dasycladaceae

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: Prioniodus: P. dychei. See conodonts

Odontopleurida

priority. See taxonomy: nomenclature Probasco, Henry, 272

Proetus parviusculus. See Arthropoda: etus parviusculus Promopalaeaster. -See Echinodermata:

Trilobita:

Decoropro-

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 demissa

repichnia. See taphonomy: trace fossils: behavior categories

reticularis, Anomaloides. See Algae: Chlorophyta:

Mollusca: Pelecypoda:

Caritodens

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

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 Brachiopoda: Articulata

Rhynchotrema. See

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, 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

Quadrijugator. See Arthropoda: Crustacea: Ostracoda quadrimucronatus, Orthograptus. See Hemichordata: Graptoloidea Queenston Delta. See paleogeography

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 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

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; taphonomy: 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

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

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

Shideler, W i l l i a m Henry, 28, 262, 263, 265, 268, 274

shideleri, Megalograptus. See Arthropoda: Chelicerata:

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

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

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

simplex, Hudsonaster. See Echinodermata: Asteroidea simplex, Palaeaster. See Echinodermata: Asteroidea: Hudsonaster simplex

scovillei, Orthis. See Brachiopoda: Articulata: Austinella

Smithsonian Institution, xv, 19, 21, 30, 153, 269

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

soft-tissue preservation, 79, 142, 143, 168, 190. See also

scovillei

senaria, Calymene. See senaria

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 snails. See M o l l u s c a : (Gastropoda

Arthropoda: Trilobita:

Flexicalymene

taphonomy 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

Solenopora. See

Sphenothallus, 93, 243, 244

spiders. See Arthropoda: Chelicerata: Arachnida spinosity, 69, 89, 123, 131, 145,148, 153, 155, 160, 162, 163, 167

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

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

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

Station Hollow Member. See Brookville Formation Stearn, Colin W., 3, 301 Steinkern. See taphonomy: molds: internal molds

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

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

Cyclonema gracile Strimple, Harrell L., 172, 173, 320

Sole-

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

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

external molds, 6, 117, 129, 281, 285, 287, 288. See also

Streptelasma. See C n i d a r i a : Anthozoa: Rugosa striata, Helcionopsis. See Mollusca: Monoplacophora striatulum, Cyclonema gracile. See Mollusca: Gastropoda:

Rhodophyta:

188, 224, 231, 282, 289, 290, 293. See also chapters on individual groups of a n i m a l s ; death assemblage

composite molds, 120, 121, 129, 130

Streptaster. See Echinodermata: Edrioasteroidea

Stromatocerium richmondense. See Algae: nopora riclunondensis

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,

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, 223, 224, 225, 226, 235, 238, 293

Chondrites, 56, 205, 211, 224, 226 Chondrites type-A, 205, 2 0 8 Chondrites type-B, 205, 2 0 8 Chondrites t y p e - C , 205, 2 0 8 Cruziana, 202, 203,211

subspecies, 39, 285, 292, 294 taxon (pl., taxa), xi, 38, 39, 42, 43, 293 type-species, 40, 42, 43 variety. See subspecies Linnaeus, Karl, 38; Systema Naturae, 38 nomenclature

epircliefs, 204, 284

Binomial System of Nomenclature, 38, 204 binomen. See species n a m e generic n a m e , 37, 41, 42, 43, 286, 291 scientific n a m e . See species n a m e species n a m e , xi, 37, 4 1 , 4 3 , 198, 203, 281, 286, 291 specific n a m e , 37, 39, 43, 286, 291, 293 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 holes (in shells). See borings hyporeliefs, 2 0 4 , 2 0 7 , 2 1 2 , 286 ichnofacies, 2 1 0 - 2 1 1 , 2 8 6 ichnogenus, ichnogenera, 204, 211 ichnospecies, 204, 2 0 6 , 2 1 0 , 2 1 1

International C o d e of Zoological Nomenclature (ICZN),xi priority, 43, 290 pronunciation of scientific n a m e s , 41 synonym, synonymy, 151; junior synonym, 153, 182.

Diplocraterion, 60, 206, 211, 224, 226, 247 D. biclavata, 2 0 6 , 2 0 9 D. cincinnatiensis, 2 0 6 , 2 0 8 , 2 0 9 D. luniformis, 206 Dystactophycus. See incertae sedis

Licrophycus: L. flahellum, 261 l.ockeia: L. siliquaria, 209, 210. See also

See also priority Mollusca:

Pelecypoda

Palaeophycus, 204, 207, 224, 226 Palaeoscia: P. floweri, 209, 212. See also

Cnidaria:

Hydrozoa: Siphonophorida: Porpitidae

Paleodictyon, 202, 203, 204, 205 Petalichnus: P. multipart Hum, 204 Petroxestes. See borings Rusophycus, 151, 152, 155,202, 203,

204, 210, 226, 235.

See also Arthropoda: Trilobita R.carleyi, 155,207,210

R. cryptolithi, 155, 210 R. pudicum, 152, 1 5 3 , 2 0 7 , 2 1 0 Sanctum. See borings Spreiten, 206, 209. See also Diplocraterion Trachomatichnus: T. numerosum, 204 Trichophycus ("turkey tracks"), 138, 211, 224, 226

T. venosum, 202, 203, 205 Trypanites. See borings

type specimens, 22, 2 5 , 4 2 , 158, 260, 266, 272, 274, 286 cotypes, 42, 283, 286, 289, 292 holotype, 42, 72, 158, 173, 178, 182, 184, 186, 209, 283, 286, 289 paratype, 42, 158, 159, 186, 283, 286, 289 syntypes. See cotypes techniques. See also chapters on individual groups of animals acetate peels, 95, 96 acid dissolution, acid etching, 68, 9 6 , 143, 163, 195, 196, 223 cluster analysis, 231 detrended correspondence analysis (DCA), 231 factor analysis, 231 health, safety, 96 microscopy, 35, 36, 50, 89, 97, 187, 198, 275 polar ordination, 231 thin-sections, 2 9 - 2 0 , 31, 95, 9 6 - 9 7 , 275 longitudinal, 77, 95, 9 6 - 9 7 , 287, 293 tangential, 95, 9 6 - 9 7 , 287, 293 transverse, 95, 9 6 , 97, 287, 293

'Vechnophorus. See Mollusca: Rostroconchia

I appan, I lelen, 69 Tate Formation, 214 taxon, taxa. See taxonomy: L i n n a e a n Hierarchy taxonomic diversity. See diversity taxonomy, 3 7 - 4 3 , 89, 198, 2 0 3 - 2 0 4 , 282, 283, 286, 288, 289,

290, 292, 293. See also chapters on individual groups of a n i m a l s based on evolutionary relationships, 39 Linnaean Hierarchy, 3 8 - 3 9 class, 38, 39, 282, 289 diagnosis, 42, 283 family, xi, 38, 39, 285, 286, 289 genus, genera, xi, 38, 3 9 , 4 0 , 4 2 , 4 3 , 283, 285, 286, 288,

291. See also ichnogenus, ichnogenera ichnogenus, ichnogenera. See taphonomy: trace fossils ichnospecies. See taphonomy: trace fossils k i n g d o m , 38, 287, 289 order, 38, 39, 282, 285, 289 phylum, phyla, 38, 118-119, 282, 287, 289 species, xi, 37, 38, 3 9 , 4 2 , 4 3 , 283, 286, 291, 292. See

also ichnospecies

344

Index

tectonic activity, 3, 5, 8, 1 0 , 5 3 , 6 1 , 6 2 , 215, 217, 233, 292 Teichert, C u r t , 43 temperature. See environment tempestite, 60, 293

Tentaculites. See Tentaculitoidea Tentaculitoidea, tentacuhtoids, 144-145, 247

Tentaculites T. richmondensis, 142, 145 T. sterlingensis, 145 tenuis, Plectodina. See conodonts tenuiseptum, Manitoulinoceras. See Mollusca: Cephalopoda Terrill Formation, 214

tessellatus, Cryptolithus. See Arthropoda: Trilobita Tetradium. See C n i d a r i a : Anthozoa: Tabulata Tetraphalerella. See Brachiopoda: Articulata Thaerodonta. See Brachiopoda: Articulata thanatocoenose, thanatocoenosis. See death assemblage T h e Society for S e d i m e n t a r y Geology. See S E P M Thecidellina. See Brachiopoda: Articulata thin-sections. See techniques: thin-sections

Thompson, Esther H., 120 ticks. See Arthropoda: Chelicerata: Arachnida tiering. See ecology: ecosystems time. See also geologic time-units absolute age, absolute dating, 4 6 , 6 1 , 62, 63, 279, 290 radiometric dating, 1 , 4 6 , 6 2 , 6 3 , 279, 290 isochronous surfaces, 46 relative age, relative dating, 45, 4 6 , 6 1 , 62, 195, 281, 290.

See also biotal succession superposition, 46 time-averaged accumulations, 7, 230, 231, 234, 293 time-environment d i a g r a m , 240 time-scale, geologic. See geologic time-scale time-stratigraphic units, 46 Tobin, Rick C, 4 4 , 5 3 , 54, 55, 56, 57, 312 Townsend, M . ) . , 212, 297 trace fossils. See taphonomy Trachomatichnus. See taphonomy: trace fossils Trammel, R. L., 12 transverse thin-sections. See techniques: thin-sections Transylvania College, University. See Kentucky: Lexington

Treatise on Invertebrate Paleontology, xi, 35, 144, 145, 255, 297, 300, 301, 304, 307, 308, 319, 322

Trematis. See Brachiopoda: luarticulata Trepostomata. See Ectoprocta Treptoceras. See Mollusca: Cephalopoda: Actinoceratoidea Treptoceras duseri shale. See Waynesvillc Formation Triarthrus. See Arthropoda: Trilobita: Olenida Triassic, xix, 4, 82, 199 Trichophycus. See taphonomy: trace fossils Tricopelta. See Arthropoda: 'Trilobita: Phacopida

Tridacna. See Mcrllusca: Pelecypoda

trifidum, Asaphoidichnus. See taphonomy: trace fossils Trilobita. See Arthropoda Trimble County, Kentucky, 124, 129 Bedford, 129 trivial n a m e . See taxonomy: nomenclature: specific n a m e Trollope, Frances, 274 tropidophora, Paupospira. See Mollusca: Gastropoda truncata, Ischyrodonta. See Mollusca: Pelecypoda Trypanites. See taphonomy: trace fossils: borings Tubastraea. See C n i d a r i a : Anthozoa: Scleractinia tuberculata, Brachiospongia. See Porifera

tuberosa, Pattersonia. See Porifera

tunicates. See Chordata: Urochordata turbidity. See environment "turkey tracks." See taphonomy: trace fossils:

turnhulli, Hercochitina. See chitinozoans

Trichophycus

Turner, Peter, 199, 316 tusk shells. See Mollusca: Scaphopoda Twenhofel, W. H., 274 Twitchell, George B., 273, 274-275 type specimen. See taxonomy type-species. See taxonomy: L i n n a e a n Hierarchy typicalis, Cymatonota. See Mollusca: Pelecypoda typicalis, Geniculograptus. See Hemichordata: Graptoloidea Tyrone Limestone. See High Bridge Group Ulrich, E. O, 6, 18, 19, 20, 2 8 - 3 1 , 32, 33, 34, 3 5 , 4 7 , 6 7 , 6 8 , 87, 111, 120, 123, 126, 127, 142, 143, 144, 160, 172,173, 176, 261, 263, 267, 271, 273, 274, 277

undatus, Phragmodus. See

conodonts

undulata, Aulaeera. See Porifera: Stromatoporoidea United States Department of Agriculture, 22 United States Geological Survey, 20, 22, 29, 30, 32, 33, 35, 51, 2 6 1 , 2 6 6 United States National M u s e u m , 22, 30, 32, 35, 260, 261. See also National M u s e u m of National History; Smithsonian Institution University of C h i c a g o , 22, 25, 28, 262 Walker M u s e u m , 2^ University of C i n c i n n a t i , 14, 16, 18, 20, 21, 22, 25, 26, 27, 28, 29, 32, 33, 34, 3 5 , 4 8 , 8 0 , 9 0 , 9 3 , 110, 138, 142, 151, 155, 156, 170, 180, 203, 209, 225, 256, 260,

261, 263, 264, 272, 275, 298. See also M c Micken Hall University of M i c h i g a n , xiii, 178, 273 Utica Group, Utica S h a l e , 4 7 , 4 8 , 160

vacuum, Foerstephyllum. See

C n i d a r i a : Anthozoa: Tabulata Vail, Peter R., 4, 320 V a l l a n d i g h a m , George L., 259, 275 vallandighami, Cyrtoceras. See Mollusca: C e p h a l o p o d a

vallata, Catellocaula. See Catellocaula Van Cleve, J. W., 275 Vanuxemia. See Mollusca, Pelecypoda

varihrachialus, Cincinnaticrinus. See Echinodermata: Crinoidea: Disparida Mollusca: Gastropoda variety. See taxonomy: L i n n a e a n 1 Iierarchy: subspecies Vaupel, Ernst H., 28, 32, 273, 275 venosa, Ropalonaria. See Ectoprocta: Ctenostomata venosum, Trichophycus. See taphonomy: trace fossils Verne, Jules, 40 Veryhachium. See acritarchs Virginia, 3, 162, 315

varicosum, Cyclonema. See

Virginia. See Chordata: Vertebrata: Reptilia volcanic ash beds, 1, 7,62, 63, 163, 285, 287. See also K-bentonite volcanic eruptions, 7, 62

vorticellatus, Streptaster. See Echinodermata: Edrioasteroidea V u l i n e c , Kevina, 7 4 , 8 6 , 100, 117, 125, 147, 195 Wabash C o l l e g e . See Crawfordsville, Indiana wackestone. See limestone W a h l m a n , Gregory P, 120 Walcott, C h a r l e s D., 25, 199 Walker M u s e u m . See University of C h i c a g o Warder, John Aston, 275 Warn. John M . , 172, 173, 182 Warren County, Ohio, 60, 105, 126, 136, 172 Warshauer, Steven M., 163 Washington, pl. 3, pl. 9 Washington, George, 106 water movement. See environment W a u g h , David A , 9 1 , 304 wave base, 221. 223. 22> normal wave-base, fair-weather wave-base, 56, 2 2 1 , 222, 223, 240 storm wave-base, 56, 221, 240. See also storms W a y n e County, Indiana, 67, 75, 228 W a y n e State University, 176

Index

345

Wavncsvillc Formation, 48, 53, 56, 60, 63, 75, 79, 87, 107, 111, 115. 120, 124, 126, 128, 129, 130, 136, 138, 142, 149, 150, 151, 153, 157, 160, 162, 172, 176, 180, 184, 196,2J4, 225, 229, 307, 314, 317, pl. 6,

pl. 10. See also Brookville Formation: W a j nesx ilk- Member Blanchester M e m b e r , 44 Clarksville M e m b e r , 4 4 Fort Ancient M e m b e r , 44 Marble Hill bed, 124, 125, 127, 129 "Treptoceras duseri shale," 60, 136, 137, 138 Waynesville, Ohio, 120, 182, 184, 185, 256, 259, 266. See Warren County, Ohio waynesvillensis. Vanuxemia. See Mollusca. Pelecypoda Weaver, T h o m a s , 75, PI S W e e d o n , M . J . , 82, 309 Weichold, C h a r l e s , 93, 275 "Weichold doughnut." See Ectoprocta "Weichold ring." See Ectoprocta: "Weichold d o u g h n u t " W e l c h , James R., 182 Welch. I. B . 276

welchi, Megalograptus. See Arthropoda: Chelicerata: Furypterida

"Lower Member," 44, 2/4 "Upper Member," 44, 214 Saluda Member. See Saluda Formation, Member Whitfield, R. P., 134, 137, 142, 145, 229, 2 7 6 - 2 7 7 W h i t t i n g t o n , Harry B., 160 Whittlesey, C h a r l e s , 269, 277 Willet, 268, 277 \\ illiains. I Icnn Shaler. 23

williamsae, Manitoulinoceras. See Mollusca: williamsae, Megalograptus. See Arthropoda:

Cephalopoda Chelicerata:

Furypterida

williamsae, Zittelloceras. See

Mollusca: Cephalopoda W i l m i n g t o n , Ohio, 260, 273, 276 Wilson, Mark A., 124, 209, 210, 281, 296, 298, 311, 315 W i l s o n , W i l l i a m W., 277

wilsoni. Polygnathus, 277 Woodward High School, C i n c i n n a t i , Ohio, 18, 26, 32, 33, 34 Woodward, Henry, 26 worms and "worms," 5, 6, 77, 86, 103,142-145, 152, 153, 155, 182, 196, 206, 210, 222, 224, 241, 242, 244, 246,

282. See also Annelida Worthen, A. H., 47, 126, 137, 170, 172, 174, 266, 268, 269, 277,311

wellsi, Acanthochaetetes. See Porifcra: sclcrosponges, coralline sponges Wcsschnan Tongue. See Kope Formation Wessels, C h a r l e s , 276 Western Academy of Natural Sciences, 15, 16, 17, 21, 259, 260, 261, 262, 263, 265, 269, 272, 273, 275, 276 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 whelks. Sec Mollusca: Castropoda: Neogastropoda Whitewater Formation, 44, 48, 49, 53, 56, 63, 67, 68, 75, 79, 109. I l l , 120, 123, 126, 127, 134, 137, 1 3 8 , 2 0 8 , 214, 223

346

Index

Xenocrinus. See

F c h i n o d c r m a t a : Crinoidea: C a m e r a t a Xiphosurida. See Arthropoda: Chelicerata Yale University, 20, 32, 260, 274, 275 Yochelson, Ellis L. 34, 35, 36, 124, 300

/.itk'lloceras. See Mollusca: Cephalopoda /.one. See biostratigraphic units zooid. See Ectoprocta, graptolites

Zygocycloides. See Echinodermata: Cyclocystoidea

Zygospira. See

Brachiopoda: Articulata

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

paleoecology.

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