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Conodont paleontology of the Permian Sabine Bay, Assistance and Trold Fiord Formations, Northern Ellesmere… Henderson, Charles Murray 1981

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CONODONT PALEONTOLOGY OF THE PERMIAN SABINE BAY, ASSISTANCE AND TROLD FIORD FORMATIONS, NORTHERN ELLESMERE ISLAND, CANADIAN ARCTIC ARCHIPELAGO by CHARLES MURRAY HE-NDERSON B.Sc,  The U n i v e r s i t y , of B r i t i s h Cblumbia, 1979  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE  REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE' in  THE The  FACULTY OF GRADUATE STUDIES  Department of G e o l o g i c a l  Sciences  We a c c e p t t h i s t h e s i s as conforming to t h e r e q u i r e d  THE  standard  UNIVERSITY OF BRITISH COLUMBIA October 1981  C h a r l e s Murray Henderson, 1981  ... -  In p r e s e n t i n g  t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the  requirements f o r an advanced degree a t the U n i v e r s i t y of B r i t i s h Columbia, I agree t h a t  the L i b r a r y s h a l l make  it  and study.  f r e e l y a v a i l a b l e f o r reference  I further  agree t h a t p e r m i s s i o n f o r e x t e n s i v e copying o f t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by t h e head o f my department o r by h i s o r her r e p r e s e n t a t i v e s . understood t h a t  copying or p u b l i c a t i o n of t h i s t h e s i s  f o r f i n a n c i a l gain  s h a l l n o t be allowed without my  permission.  Department  of  Geological  Sciences  The U n i v e r s i t y o f B r i t i s h 2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5 Date  Iti s  October 2, 1981  Columbia  written  ii ABSTRACT A. s u c c e s s i o n o f l a t e E a r l y through M e d i a l Permian conodont faunas i s documented f o r t h e f i r s t  time from t h e c a l c a r e o u s , f i n e g r a i n e d ,  quartzose  sandstones o f t h e A s s i s t a n c e and T r o l d F i o r d Formations on n o r t h e r n mere I s l a n d , Northwest T e r r i t o r i e s . one  s p e c i e s and t h r e e s u b s p e c i e s  in c h r o n o l o g i c a l order:  minutus, N e o g o n d o l e l l a  Of t h e t a x a i d e n t i f i e d and d e s c r i b e d ,  a r e proposed as new.  Neogondolella  tognathodus p r a y i , N e o g o n d o l e l l a  The taxa i n c l u d e ,  i d a h o e n s i s subsp. i n d e t . , N e o s t r e p -  i d a h o e n s i s n.subsp. A, Anchignathodus  s e r r a t a ( ? ) , N. n.sp. B, N. p o s t s e r r a t a ( ? ) , N. b i t t e r i  n.subsp. C, and N. r o s e n k r a n t z i n.subsp. D. also associated with Neogondolella  s e p a r a t e form s p e c i e s .  Numerous ramiform elements a r e  i d a h o e n s i s n.subsp. A.  may comprise p a r t o f a multielement represent  Elles-  Neogondolella  These elements  apparatus or they  I n observation of t h e i r  may  questionable  s t a t u s , a somewhat u n s a t i s f a c t o r y d u a l taxonomy i s proposed f o r t h e s e elements, and i n c l u d e s t h e f o l l o w i n g t a x a : N. i d a h o e n s i s n.subsp. A - Xaniognathus t o r t i l i s , sis  N. i d a h o e n s i s n.subsp. A - E l l i s o n i a excavata,  N.  idahoen-  n.subsp. A - E l l i s o n i a t r i b u l o s a , and N. i d a h o e n s i s n.subsp. A - P r i -  oniodella  decrescens.  S t a t i s t i c a l work on t h e abundant p l a t f o r m elements of N. i d a h o e n s i s n.subsp. A p r o v i d e s evidence  f o r minor ' e v o l u t i o n a r y t r e n d s o f i n c r e a s i n g  s i z e and i n c r e a s i n g number o f d e n t i c l e s u p s e c t i o n .  Comparison o f these c o n -  odonts w i t h N. s e r r a t a . a n d N. p o s t s e r r a t a from t h e Great B a s i n o f SW USA suggests  t h a t t h e p h y l o g e n e t i c development o f Permian N e o g o n d o l e l l a  lowed an,, e v o l u t i o n a r y path.more a p p r o p r i a t e to punctuated to  phyletic  fol-  equilibria  gradualism.  The conodont taxa i n d i c a t e t h a t t h e A s s i s t a n c e Formation i s Upper  than  Leonardian  t o Uppermost Roadian i n age whereas t h e T r o l d F i o r d  i n c l u d e s most o f t h e Wordian.stage.  Formation  These two f o r m a t i o n s have been s e p a r -  a t e d i n t o f i v e s u b d i v i s i o n s on t h e b a s i s o f b o t h l i t h o l o g y . and t h e presence or absence o f v a r i o u s b i o t a 1  A s i x t h , s u b d i v i s i o n i s d e s c r i b e d f o r t h e Sa-  b i n e Bay Formation which u n d e r l i e s t h e A s s i s t a n c e and where conodonts a r e apparently  absent.  L i t h o l o g i c and b i o t i c evidence  ( i n c l u d i n g t r a c e f o s s i l s and mega- and  m i c r o b i o t a ) p o i n t t o shallow, o f f s h o r e marine c o n d i t i o n s w e l l w i t h i n t h e p h o t i c zone and c h a r a c t e r i z e d by low energy f o r most o f t h e conodont b e a r i n g s t r a t a .  and slow d e p o s i t i o n a l  rates,  A much t h i c k e r c o r r e l a t i v e  sec-  t i o n t o t h e south r e p r e s e n t s , i n l a r g e p a r t , a d e l t a f r o n t sequence.  The  Sabine Bay Formation,  on t h e o t h e r hand, i s composed o f s h o r e f a c e  possibly i n a barrier island  sandstones,  setting.  The r e s u l t s o f t h i s r e s e a r c h i n d i c a t e t h a t conodonts may be v e r y p r o m i s i n g f o r c o r r e l a t i o n o f Permian s t r a t a i n the Canadian A r c t i c pelago and f o r worldwide comparison.  More work w i t h i n t h e Sverdrup  ArchiBasin,  i n c l u d i n g both m a r g i n a l and b a s i n a l s e c t i o n s , i s n e c e s s a r y t o p r o v i d e a good b i o z o n a t i o n o f these marine Permian s t r a t a . brachiopods  i n combination  resolve t h i s zonation.  The use o f t h e abundant  w i t h t h e conodonts i s p r o b a b l y t h e b e s t way t o  The t a x o n o m i c . d e s c r i p t i o n s and s u b d i v i s i o n s p r o -  posed h e r e i n should p r o v i d e a f o u n d a t i o n f o r f u t u r e work.  iv  TABLE OF CONTENTS T i t l e page  i  Abstract  i i  Table o f Contents  iv  L i s t of Tables List  v i i  of F i g u r e s  viii  L i s t of P l a t e s  i  Acknowledgements  x  x  Introduction  1  L o c a t i o n and Scope of t h e Study  1  F i e l d Work  3  P r e v i o u s Work  4  L a b o r a t o r y and A n a l y t i c a l Methods.  6  S t r a t i g r a p h y and Paleoenvironment - General  Statement  10  Sabine Bay Formation  10  Hamilton  Peninsula area  10  McKinley  Bay a r e a  14  Tanquary F i o r d a r e a  14  A s s i s t a n c e Formation  15  Hamilton  Peninsula area  15  McKinley  Bay a r e a  16  Sawtooth Range area T r o l d F i o r d Formation Hamilton  P e n i n s u l a area  ".  17 18 18  McKinley. Bay a r e a  21  Sawtooth Range area  22  V  Age and C o r r e l a t i o n . . . R e l a t i v e value of v a r i o u s  23 f o s s i l biota........  "."25 :  P r e v i o u s Conodont work  28  E a r l i e r age assignments f o r A r c t i c Permian Formations  29  Sabine Bay Formation Assistance  29  Formation  -  T r o l d F i o r d Formation....  30 30  Age assignments r e s u l t i n g from t h i s work  30  S u b d i v i s i o n A....  31  Subdivision B  32  Subdivision  34  C  S u b d i v i s i o n D.  36  Subdivision E  37  Subdivision F  38  Summary  40  Q u a n t i t a t i v e A n a l y s i s o f Measurable c h a r a c t e r s  f o r Neogondolella  40  Introduction  40  R e s u l t s and D i s c u s s i o n  45  O v e r a l l length of platform  (Ll)  Number o f d e n t i c l e s per element  46 (#)  54  R a t i o o f l e n g t h t o number o f d e n t i c l e s (Ll/#)  54  Length from.'.tip:'."of cusp t o f o u r t h d e n t i c l e a n t e r i o r (L2) . ...  57  Maximum w i d t h (Wl)  57  Width?.at p o s t e r i o r end (W2)  57  H e i g h t from t i p o f cusp t o base o f t h e f l a n g e . (HI)  57  Ll/Wl r a t i o .  '"59  vi  ,L1/HI r a t i o  60  P o s t e r i o r area  60  D i s c u s s i o n o f E v o l u t i o n a r y t r e n d s and c o n c e p t s  61  Systematic Paleontology  69  Introduction  69  Anchignathodus .minutus.  73  Neostreptognathodus p r a y i  73  N e o g o n d o l e l l a i d a h o e n s i s subsp. iridet  74  Neogondolella  i d a h o e n s i s n.subsp. A  75  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A v a r , g r a c i l i s  84  Neogondolella  i d a h o e n s i s n.subsp. A v a r . r o b u s t u s  85  Neogondolella  i d a h o e n s i s n.subsp. A v a r . i n t e r m e d i a t u s  86  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A v a r , c o n s t r i c t u s  86  Neogondolella  87  i d a h o e n s i s n. subsp. A v a r . l o b a t u s  N e o g o n d o l e l l a s e r r a t a ( ?.) . . . N e o g o n d o l e l l a n.sp. B_  90 92  Neogondolella postserrata(?)  94  N e o g o n d o l e l l a b i t t e r i n.subsp.  Cj  95  N e o g o n d o l e l l a r o s e n k r a n t z i n.subsp. I).  98  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A - Xaniognathus t o r t i l i s  103  Neogondolella  i d a h o e n s i s n.subsp. A - E l l i s o n i a  104  Neogondolella  i d a h o e n s i s n.subsp. A - E l l i s o n i a . t r i b u l o s a  104  Neogondolella  i d a h o e n s i s n.subsp. A - P r i o n i o d e l l a d e c r e s c e n s . . . .  105  excavata  References.......  106  Plates 1 - 8  113  Appendix,!-.  •••  129-135  vii LIST OF TABLES T a b l e 1.  Statistics  T a b l e 2.  Values d e r i v e d from z - t e s t s f o r L l , # *d Ll/#  T a b l e 3.  Counts and percentages of p l a t f o r m s and N e o g o n d o l e l l a i n F48 - F54.  T a b l e 4.  T a b l e 5.  from d a t a g i v e n i n Appendix I . . . .  47 49  ai  S t a t i s t i c s from sample subsets denticles)  ramiforms.of  (elements w i t h 10 and  Values d e r i v e d from t - t e s t s on sample s u b s e t s  51 11 52 52  viii  LIST OF FIGURES F i g u r e 1.  Map showing l o c a t i o n s o f s e c t i o n s s t u d i e d  F i g u r e 2.  Generalized l i s t of c h a r a c t e r i s t i c s f o r v a r i o u s marine environments  F i g u r e 3.  F i g u r e 4.  F i g u r e 5.  2 shallow 11  L i t h o l o g i c and b i o l o g i c c h a r a c t e r i s t i c s and c o r r e l a t i o n of f o r m a t i o n s and s e c t i o n s .  12  C o r r e l a t i o n c h a r t f o r s e r i e s , stages and zones o f t h e Permian  24  Frequency d i s t r i b u t i o n o f number o f d e n t i c l e s and p l a t f o r m l e n g t h f o r N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A  43  F i g u r e 6.  Graphical r e p r e s e n t a t i o n of part of Table 1  48  F i g u r e 7. (a,b,c)  Graphs showing r e l a t i o n s h i p o f t h e p l a t f o r m l e n g t h t o number of d e n t i c l e s p e r element f o r F49, F54 and F96  53  F i g u r e 8.  Graph showing r e l a t i o n s h i p o f t h e p l a t f o r m l e n g t h t o number o f d e n t i c l e s per element o f N. i d a h o e n s i s n.subsp. A, !N. s e r r a t a , and _N. p o s t s e r r a t a ( i n c h r o n o l o g i c a l o r d e r ) . . .  56  Graphical representation of part of(Table 1  58  F i g u r e 9.  F i g u r e 10. Graph showing r e l a t i o n s h i p o f Length/Height t o Length of p l a t f o r m f o r F49  79  F i g u r e 11. Graph showing r e l a t i o n s h i p of .Length/Width to Length of p l a t f o r m f o r F49  80  F i g u r e 12. Graph showing r e l a t i o n s h i p o f P o s t e r i o r Area t o Length of p l a t f o r m for.F53  81  F i g u r e 13. Graph showing r e l a t i o n s h i p o f P o s t e r i o r Area t o Length of p l a t f o r m f o r F49  82  F i g u r e 14. Graph showing r e l a t i o n s h i p o f P o s t e r i o r Area t o Length/ number of d e n t i c l e s f o r F49  83  LIST OF PLATES All  f i g u r e s on p l a t e s a r e Scanning  P l a t e 1.  Electron  Micrographs.  Neostreptognathodus p r a y i , Anchignathodus minutus, Neog o n d o l e l l a i d a h o e n s i s s u b s p . i n d e t . , Neogondo1ella idahoens i s n.subsp. A - P r i o n i o d e l l a decrescens. and N e o g o n d o l e l l a i d a h o e n s i s n. subsp. A - E l l i s o n i a excavata.  113  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A - E l l i s o n i a t r i b u l o s a , N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A - Xaniognathus t o r t i l i s , and N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A  115  P l a t e 3.  Neogondolella. i d a h o e n s i s n.subsp.. A  117  P l a t e 4.  Neogondolella  i d a h o e n s i s n.subsp.. A  119  P l a t e 5.  Neogondolella.. i d a h o e n s i s n.subsp. A  121  P l a t e 6.  N e o g o n d o l e l l a n.sp. IS, N e o g o n d o l e l l a p o s t s e r r a t a Q ? ) , and Neog o n d o l e l l a s e r r a t a ( ?)  123  P l a t e 2.  P l a t e 7.  N e o g o n d o l e l l a b i t t e r i n.subsp. Cj and N e o g o n d o l e l l a r o s e n k r a n t z i n.subsp. I) 125  P l a t e 8.  N e o g o n d o l e l l a r o s e n k r a n t z i n.subsp. I)  127  X  ACKNOWLEDGEMENTS There a r e a number of people t o whom I became indebted d u r i n g the c o u r s e of t h i s r e s e a r c h . F i r s t of a l l ,  I would l i k e to thank my  a d v i s o r , Dr. R.V..  Best, f o r  h i s encouragement and p a t i e n t e d i t i n g of e a r l i e r d r a f t s of t h i s  thesis.  H i s c o n t r i b u t i o n has improved immensely the q u a l i t y of the f i n a l I would a l s o l i k e to express my  g r a t i t u d e to the o t h e r members of my  m i t t e e : Drs. G.E.  Rouse and P.L.  G e o l o g i c a l Survey  of Canada, Vancouver.  acknowledge Mike Orchard, c u s s i o n s we  Smith of UBC  and  Dr. M.  Orchard,  of  comthe  In p a r t i c u l a r , I would l i k e to  a f e l l o w conodont worker, f o r the f r u i t f u l  dis-  had w i t h r e g a r d s to conodonts and t h e i r taxonomic problems.  I g r a t e f u l l y thank the G e o l o g i c a l Survey f i n a n c i a l and ticular,  manuscript.  l o g i s t i c a l support d u r i n g the f i e l d  I would l i k e to acknowledge W.W.  for i n i t i a l l y  of Canada i n C a l g a r y f o r the  s u g g e s t i n g the p r o j e c t and  season of 1979.  Nassichuk,  In p a r -  A. Embry, and U. Mayr  l a t e r f o r p r o v i d i n g support  and  useful discussions. Foremost, I wish to express my who  not o n l y typed p a r t of the manuscript  source of encouragement and this  deepest  a p p r e c i a t i o n to my  wife, Betty,  but a l s o served as an  endured my many l a t e evenings  spent  endless completing  thesis. F i n a l l y , I would l i k e to d e d i c a t e t h i s t h e s i s to the l a t e Dr.  G. P e r r y .  Dave was  the o r i g i n a l a d v i s o r to my  E l l e s m e r e I s l a n d f o r p a r t of the c o l l e c t i n g m i s i n g l i f e was  t h e s i s and had j o i n e d me  i n early July,  cut s h o r t by a h e l i c o p t e r c r a s h b e f o r e my  even ended i n August, 1979. memory p r o v i d e d me  David  1979. field  on  His proseason  had  H i s death:.was a t r a g i c and b i t t e r blow but h i s  w i t h the i n s p i r a t i o n and m o t i v a t i o n to c o n t i n u e and com-  plete t h i s research.  1  INTRODUCTION T h i s t h e s i s r e c o r d s t h e r e s u l t s o f a study of t h e conodont i g r a p h y o f t h e Permian Sabine Bay, A s s i s t a n c e and T r o l d F i o r d of n o r t h e r n E l l e s m e r e I s l a n d , N.W.T. l i t h o f a c i e s were u t i l i z e d  biostrat-  Formations  To a l e s s e r extent b r a c h i o p o d s and  to e s t a b l i s h  correlations.  L o c a t i o n and Scope o f t h e Study E l l e s m e r e I s l a n d , t h e most n o r t h e r l y i s l a n d of t h e Canadian A r c h i p e l a g o , i s l o c a t e d between 76oahd 83°North  latitude.  Arctic  The study i s  based p r i m a r i l y on seven s e c t i o n s from f o u r d i f f e r e n t a r e a s i n c l u d i n g two from Hamilton P e n i n s u l a (80°10' N, 081°45' W), two from McKinley Bay  (81°10' N, 079°10'W), two from t h e h e a d o f  076°30' W) and one from t h e Sawtooth  Fiord  (81°25' N,  Range (79°30' N, 083°20' W)  A, B, C, and D r e s p e c t i v e l y on F i g . 1 ) . i n minor d e t a i l  Tanquary  (sections  Other s e c t i o n s have been s t u d i e d  ( F i g . 1) and"are o n l y r e f e r r e d t o where they proved v a l u a b l e  as support f o r any i n t e r p r e t a t i o n s . The l i t h o l o g y  s t u d i e d i n t h e above mentioned  sections represent the  m a r g i n a l f a c i e s f o r t h e Permian p a r t o f t h e Sverdrup B a s i n ; a b a s i n of d e p o s i t i o n from e a r l y C a r b o n i f e r o u s t o T e r t i a r y . and r e c o n n a i s s a n c e s t u d i e s by p r e v i o u s workers 1964;  Nassichuk and C h r i s t i e ,  Despite::the d e s c r i p t i v e  ( T h o r s t e i n s s o n , 1974; C h r i s t i e ,  1969; and Mayr, 1976) t h e Permian p a r t o f  the b a s i n remains t h e p o o r e s t understood of t h e Phanerozoic systems.  Fossil  c o l l e c t i o n s and age d e t e r m i n a t i o n s have been p r e v i o u s l y r e p o r t e d from t h e Sabine Bay, A s s i s t a n c e and T r o l d F i o r d Formations by Nassichuk et a l . (1965), Harker and T h o r s t e i n s s o n (1960), Nassichuk (1970), Nassichuk and S p i n o s a (1970) and by J.B. Waterhouse and R.E. Grant i n T h o r s t e i n s s o n (1974). r e p o r t e d on h e r e i n c o n s t i t u t e s t h e f i r s t  The m a t e r i a l  s y s t e m a t i c c o l l e c t i o n s through t h e  2  Figure 1. Map of northern Ellesmere Island, showing location of the sections studied.  3  complete  s e c t i o n s : the p r e v i o u s r e p o r t s were l a r g e l y o f i s o l a t e d o c c u r r e n c e s .  These p r e v i o u s s t u d i e s emphasized correlation.  ammonoids and brachiopods to f a c i l i t a t e  Both of t h e s e groups have t h e i r own  c i a t e d w i t h them (see p. 25)  p e c u l i a r problems a s s o -  t h a t h i n d e r c o r r e l a t i o n schemes.  emphasizes the use o f conodonts,  This report  a group whose once many problems h i n d e r i n g  c o r r e l a t i o n have been l a r g e l y i r o n e d out by i n t e n s i v e r e s e a r c h over the p a s t f i v e y e a r s ( C l a r k and Behnken, 1979; Collinson,  C l a r k et a l . , 1979  and Wardlaw and  19 79b).  I t i s because  o f t h e c o r r e l a t i o n problems f o r c e r t a i n f o s s i l groups,  the abrupt l i t h o l o g i c  changes over s h o r t d i s t a n c e s , and the presence of  d i s c o n f o r m i t i e s and t r a n s g r e s s i v e u n i t s t h a t the c o r r e l a t i o n and e n v i r o n mental r e l a t i o n s h i p s of the f o r m a t i o n s p e r t a i n i n g to t h i s r e p o r t a r e p o o r l y understood.  The o r i g i n a l i n t e n t i o n f o r the r e s e a r c h was  the age and c o r r e l a t i o n of t h e s e f o r m a t i o n s . accomplished  through the use of conodonts  to b e t t e r d e f i n e  T h i s seems to have been  a l t h o u g h the r e s u l t s should o n l y  be regarded as a b e g i n n i n g , but a s t a r t t h a t a t l e a s t j u s t i f i e s  optimism.  F i e l d Work Access to the study a r e a i s by Twin O t t e r or DC-3 o l u t e Bay,  aircraft  C o r n w a l l i s I s l a n d , to Eureka or Tanquary F i o r d  From t h e s e bases a c c e s s to the s e c t i o n l o c a l i t i e s was  from Res-  airstrips.  accomplished  through  the use of J e t Ranger h e l i c o p t e r s . The f i e l d work was August  11, 1979.  completed  d u r i n g t h r e e weeks between June 16 and  The s e c t i o n d e s c r i p t i o n was  completed  w i t h the a i d of an  a s s i s t a n t from f l y camps c o n s i s t i n g of a l o g a n and a pyramid of t h e l o c a l i t i e s .  tent at  Radio c o n t a c t was m a i n t a i n e d a t r e g u l a r times w i t h the  main base a t Eureka or Tanquary F i o r d to r e p o r t weather, and to move d a t e s and  each  supplies required.  indicate  The weather through t h e p e r i o d i n d i c a t e d above was and  c l o u d and  i n c l u d e d o n l y two weather r e l a t e d down-days.  seven day p e r i o d saw or r a i n . and  a, m i x t u r e  During  This  of  sun  fifty-  t h r e e days w i t h snow f l u r r i e s and n i n e days of showers  t h i s e n t i r e p e r i o d the s e c t i o n s s t u d i e d were f r e e of i c e  snow w i t h the e x c e p t i o n of the minor f l u r r i e s .  t h e o r d e r of t h e day w i t h temperatures but more t y p i c a l l y averaged  Otherwise sunshine  r e a c h i n g as h i g h as 19oC  3 to 10°C.  (July  was 30)  Daytime temperature f l u c t u a t i o n s  were minor as a t t h i s l a t i t u d e the sun remains above t h e h o r i z o n from A p r i l 15 to August 29  ( T h o r s t e i n s s o n , 1974).  P a r t of the camp remained a t  Tanquary F i o r d a f t e r the IT "* of August but snow began to f a l l on the 1  and camp was  1  f o l d e d f o r t h e season by t h e 16  when no break was  12  t h  in sight.  P r e v i o u s Work The  summary of p r e v i o u s work i n the a r e a as p r e s e n t e d h e r e i n , and  e s p e c i a l l y of the e a r l y h i s t o r y , G e o l o g i c a l Survey of Canada (GSC) is  i s l a r g e l y taken from R.L. Memoir 331  (1964) and  Christie's  to whom the  credit  due. The h i s t o r y of e x p l o r a t i o n and  g e o l o g i c a l i n v e s t i g a t i o n of n o r t h e r n  E l l e s m e r e I s l a n d i s a v e r y a u s p i c i o u s and  c o l o u r f u l one.  The f i r s t  geolo-  g i c a l s t u d i e s were by a B r i t i s h e x p l o r e r , C a p t a i n S i r George Nares, on a 1875-76 Royal Navy e x p e d i t i o n to Lady F r a n k l i n Bay. a n a t u r a l i s t on Nares' e x p e d i t i o n , and r o c k s and Peninsula.  f o s s i l s i n the r e g i o n between D i s c o v e r y Harbour and F e i l d e n L i e u t e n a n t Adolphus W.  G r e e l y of the .U.S.  Army e s t a b l i s h e d  E x p e d i t i o n s went t o Lake Hazen  G r e e l y F i o r d d u r i n g 1882-83 where g e o l o g i c a l and a r c h a e o l o g i c a l specimens  were c o l l e c t e d and gic  Feilden,  o t h e r s made e x t e n s i v e c o l l e c t i o n s of  F o r t Conger i n D i s c o v e r y Harbour i n 1881. and  C a p t a i n H.W.  c o p i o u s notes made.  T h i s success was  end of t h e e x p e d i t i o n where a l l but seven men  t a i n t e d by the  tra-  d i e d of s t a r v a t i o n because  5  a planned rendesvous w i t h a r e t u r n v e s s e l was The a r e a was  e x p l o r e d by a number of o t h e r s over the next seventy y e a r s  i n c l u d i n g Commander R.E. (1913-17) who  late.  Peary  c o l l e c t e d Permian  (1898-1909),  a g e o l o g i s t W.  Elmer  Ekblaw  f u s u l i n i d s from near the mouth of  Tanquary  F i o r d , and a g e o l o g i s t Dr. J.C. T r o e l s e n (1939-40). The f i r s t appearance of the GSC was n o r t h e a s t shore of E l l e s m e r e . and R.G.  B l a c k a d a r (GSC)  Lake Hazen a r e a . 1957,  and  1958  331.  In 1956 and  R.L.  i n 1948  by V.K.  G. H a t t e r s l e y - S m i t h (Defence Research Board)  l a t e r conducted g e o l o g i c a l r e c o n n a i s s a n c e i n the  C h r i s t i e of the GSC conducted f i e l d  i n n o r t h e a s t E l l e s m e r e p r o d u c i n g a map 1957  Ellesmere Island.  P r e s t a l o n g the  work i n  1954,  i n h i s GSC Memoir  R. T h o r s t e i n s s o n and E.T. Tozer i n v e s t i g a t e d  T h i s work and much of the p r e v i o u s work was  means of dog teams and canoe over extended f i e l d  seasons.  western  conducted  In 1961  and  by 1962  O p e r a t i o n Eureka, under the d i r e c t i o n of R. T h o r s t e i n s s o n o f the GSC, i n c l u d e d J . Wm.  K e r r , E.T. T o z e r , and H.P.  t r a n s p o r t a t i o n i n c l u d e d P i p e r Super In  1963  Trettin.  Cub a i r c r a f t and a G2A  R. T h o r s t e i n s s o n and P. Harker conducted f u r t h e r  s t u d i e s and mapping of E l l e s m e r e .  These f i v e f i e l d  t i o n f o r T h o r s t e i n s s o n ' s . G S C B u l l e t i n 224 major work on C a r b o n i f e r o u s and Permian P r e v i o u s f o s s i l work was indicated  stratigraphic  seasons a r e the founda-r.'.  s t r a t i g r a p h y i n the a r e a .  However, l a t e Lower  Permian  from the a r e a have o n l y been r e p o r t e d  (Kozur and Nassichuk,  (see p. 28  helicopter.  (1974) which remains today as the  i n the f i r s t p a r t of t h i s c h a p t e r .  once p r e v i o u s l y  period  l a r g e l y on b r a c h i o p o d s and ammonoids as  through M i d d l e Permian conodonts  lections  During t h i s  1977)  and t h i s was  of j u s t two  col-  for details) .  The d e s i g n a t i o n of the t h r e e f o r m a t i o n s of t h i s r e p o r t d a t e between 1960 and 1974.  The Sabine Bay Formation was  named by Tozer and  Thorsteins-  6  son  (1964) f o r a . s e c t i o n on Sabine P e n i n s u l a , M e l v i l l e I s l a n d .  tance Formation was for  named and d e f i n e d by Harker and  was  Assis-  T h o r s t e i n s s o n (1960)  a s u c c e s s i o n on G r i n n e l l P e n i n s u l a , Devon I s l a n d .  F i o r d Formation  The  F i n a l l y , the T r o l d  d e f i n e d by T h o r s t e i n s s o n (1974) and  includes a  type  s e c t i o n on a s m a l l , unnamed t r i b u t a r y of the East Cape R i v e r t h a t i s s u e s i n t o the n o r t h e a s t s i d e of Canon F i o r d on the west c o a s t of (very near the Hamilton The most  precise  C a r b o n i f e r o u s and  P e n i n s u l a . s e c t i o n s of t h i s way  Ellesmere  report).  to summarize the p r e v i o u s work i n the a r e a  Permian r o c k s i s to say t h a t the r e c o n n a i s s a n c e  completed but t h a t d e t a i l e d  s t u d i e s a r e merely  on  has  been  beginning.  L a b o r a t o r y and A n a l y t i c a l Methods L a b o r a t o r y work was A standard  conducted  from the f a l l of 1979  t e c h n i q u e of a c e t i c a c i d d i s s o l u t i o n , wet  quid s e p a r a t i o n (tetrabromoethane) from t h e i r host  was  to the s p r i n g of  1981.  s i e v i n g , and heavy l i -  used to c o n c e n t r a t e the conodonts  rocks.  The b u l k samples t h a t were processed were of two  types.  The f i r s t  con-  s i s t e d of l a r g e s i n g l e b l o c k s or a number of moderate s i z e d s l a b s t h a t weighed up to 25 kg  (55 l b s . ) but more t y p i c a l l y averaged  These b l o c k s were c o l l e c t e d f o r t h e i r f i n e l y s i l i c i f i e d  10 kg  (22  brachiopod  lbs.). content  which were to have been the major emphasis of the r e s e a r c h ( t h e emphasis switched  to conodonts about half-way  type c o n s i s t e d of 2 to 3 cm diameter  through  second  between 3.2  and 4.2  kg  (7 to 9 l b s . )  total. The  l a r g e b l o c k s were broken i n t o two  broken o f f the b l o c k s and to  The  c h i p s c o l l e c t e d from s i n g l e h o r i z o n s  s p e c i f i c a l l y f o r conodonts and weighing in  the p r o c e s s i n g ) .  3/4  of the t o t a l ) was  fractions.  Small fragments were  r e t a i n e d f o r conodonts w h i l e the remainder p l a c e d i n h y d r o c h l o r i c a c i d baths  (diluted,  (1/2 but  7 not  to s p e c i f i c percent  was not t o be so s t r o n g s i l s ) to separate placed  as t h e o n l y c r i t e r i o n to.be met was t h a t  bubbling  as t o cause f u r t h e r breakage o f t h e s i l i c i f i e d  the s i l i c i f i e d  brachiopods.  fos-  The conodont samples were  i n p l a s t i c buckets which were subsequently  filled  with a s o l u t i o n  of 60% g l a c i a l a c e t i c a c i d a t a d i l u t i o n o f 1 p a r t a c i d t o 6 t o 9 p a r t s water ( t o keep t h e a c i d a t o r below 10% - s t r o n g e r a c e t i c a c i d tends t o e t c h t h e conodonts w h i l e any s t r e n g t h o f h y d r o c h l o r i c w i l l d i s s o l v e t h e conodonts) .  The samples were l e f t i n a fume hood f o r up t o two months but  more t y p i c a l l y f o r two t o t h r e e weeks w i t h  t h e a c i d being changed weekly.  The  f o r such work was r e q u i r e d be-  l o n g e r than normal d i s s o l u t i o n period,  cause t h e r o c k s , b e i n g c a l c a r e o u s s o l v e and c o n t a i n e d  quartzose  sandstones, were slow t o d i s -  a h i g h p e r c e n t a g e of i n s o l u b l e s (as opposed to pure  c a r b o n a t e s which a r e more commonly sampled f o r c o n o d o n t s ) .  Even a f t e r  these  l o n g p e r i o d s , t h e samples were r a r e l y e n t i r e l y d i s s o l v e d and d i s s o l u t i o n was u s u a l l y d i s c o n t i n u e d a f t e r i t was f e l t s u f f i c i e n t i n s o l u b l e r e s i d u e had been separated.  As a r e s u l t , i t i s i m p o s s i b l e  of i n s o l u b l e r e s i d u e .  I t was n e c e s s a r y  to r e p o r t t h e . a c t u a l p e r c e n t a g e  t o use a m o d i f i e d  l a t i n g t h e conodonts due to t h e l a r g e i n s o l u b l e f r a c t i o n s .  procedure f o r i s o A f t e r the  samples were d i s s o l v e d , they were wet s i e v e d and washed through a f o u r s i e v e s t a c k c o n s i s t i n g o f 20 (.841 mm), 200 mesh (.075;mm) standard  35 (.500 mm),  21 cm diameter s i e v e s .  100 (.150 mm)  On top o f t h i s  and stack  was a 1.2 mm n y l o n s c r e e n to r e t a i n t h e c o a r s e s t p a r t i c l e s and u n d i s s o l v e d chunks.  The two c o a r s e s t s i e v e s were used t o s e p a r a t e any c o a r s e  small undissolved  sand o r  fragments from.the f i n e sand and, h o p e f u l l y , conodonts  which would be trapped  i n t h e f i n e s t two s i e v e s .  T h i s s t a c k was  because o f t h e h i g h percentage o f i n s o l u b l e m a t e r i a l .  necessary  Normally, a s i n g l e  8 150 or 200 mesh s i e v e w i t h a n y l o n s c r e e n on top i s s u f f i c i e n t f o r the i n s o l u b l e s o f r e l a t i v e l y pure c a r b o n a t e s . r e t a i n e d and l e f t  A l l of the i n s o l u b l e r e s i d u e was  to a i r d r y i n p o r c e l a i n c r u c i b l e s .  f r a c t i o n that f i l t e r e d  through the f i n e s t  p l a s t i c buckets and a l l o w e d to s e t t l e . of the excess water was p l a s t i c containers.  poured  o f f and  The s i l t  and  clay  s i e v e was. also, c o l l e c t e d i n  A f t e r the sediment the wet  sediment  had  s e t t l e d most  s t o r e d i n covered  Some of these samples were l a t e r a n a l y z e d f o r t h e i r  palynomorph c o n t e n t i n c o n j u n c t i o n w i t h a graduate c o u r s e w i t h G.E. at  UBC. A f t e r d r y i n g , the 100 and  200 mesh i n s o l u b l e f r a c t i o n s were p l a c e d i n  s e p a r a t o r y f u n n e l s f i l l e d w i t h tetrabromoethane  ( s p e c i f i c g r a v i t y = 2.89).  The remaining c o a r s e r i n s o l u b l e s were p l a c e d i n a c o n t a i n e r and cabinets. a light and  Rouse  The  i n s o l u b l e s i n the gegaratory f u n n e l s d i v i d e d  stored i n  i n t o two  fractions:  f r a c t i o n f l o a t i n g on top and c o n s i s t i n g of q u a r t z , c h e r t , g l a u c o n i t e  silicified  or s i l i c e o u s m i c r o f o s s i l s and a heavy f r a c t i o n s i n k i n g t o the  bottom and c o n s i s t i n g o f opaques, i r o n coated g r a i n s , f i s h d e b r i s ( t e e t h , p l a t e s . . . ) and conodonts ( s p e c i f i c g r a v i t y = 2.84  to 3.10:  Ellison,  1944).  These heavy f r a c t i o n s were then allowed to run out of the f u n n e l onto a filter  paper.  paper.  S i m i l a r l y , the l i g h t f r a c t i o n was  The tetrabromoethane  was  f i l t e r e d onto a s e p a r a t e  c o n s t a n t l y reused owing t o the h i g h c o s t  of the mat e r i a l .  These f r a c t i o n s were then t h o r o u g h l y washed w i t h  and l e f t  to d r y .  The acetone w i t h i t s d i s s o l v e d  t i o n was  p l a c e d i n an. open beaker and allowed to evaporate i n a fume hood  u n t i l the tetrabromoethane  was  tetrabromoethane  acetone in solu-  c o n c e n t r a t e d (acetone evaporates more r a p i d l y ) .  T h i s procedure allowed o n l y minimal  l o s s of heavy l i q u i d w i t h each s e p a r a t i o n .  A f t e r d r y i n g , the conodonts were p i c k e d from the heavy f r a c t i o n s w i t h the  9  a i d of a b i n o c u l a r microscope if  and a wet, v e r y f i n e p a i n t b r u s h .  However,  the heavy f r a c t i o n was l a r g e and c o n t a i n e d abundant i r o n m i n e r a l s , the  sample was passed through a magnetic  s e p a r a t o r where the conodonts a r e  f u r t h e r c o n c e n t r a t e d i n the non-magnetic heavy f r a c t i o n . saves unnecessary  This  procedure  time spent p i c k i n g n o n - p r o d u c t i v e r e s i d u e s .  The conodonts h a v i n g been c o n c e n t r a t e d from t h e i r r o c k i n abundances, when p r e s e n t , r a n g i n g available for detailed  from  one-  study.  to as many as 150 per kg (F49), were thus  The a n a l y s i s of these, faunas f i r s t c o n s i s t e d  of s i m p l e o b s e r v a t i o n under the b i n o c u l a r microscope Secondly,  and d e s c r i p t i o n .  the samples were measured f o r v a r i o u s parameters  meter mounted on a . b i n o c u l a r microscope.  with a micro-  These measurements were used to  enhance d e s c r i p t i o n s and s u b j e c t e d to v a r i o u s s t a t i s t i c a l procedures as out'^ lined  i n a l a t e r chapter.  Microscope the SEM.  T h i r d l y , the samples were putaonl. Scanning  (SEM) s t u b s , coated w i t h g o l d - p a l l a d i u m , and photographed  Electron with  These photos b e s i d e s p r o v i d i n g t h e i l l u s t r a t i o n s f o r the p l a t e s  facilitated  even more d e t a i l e d d e s c r i p t i o n .  would be i m p o s s i b l e without t h e SEM.  Indeed,  adequate d e s c r i p t i o n  The i d e n t i f i c a t i o n and  to o t h e r s i m i l a r conodonts to determine  comparison,  the age r e l a t i o n s h i p s and c o r r e l a -  t i o n of the s t u d i e d s e c t i o n s concluded t h e a n a l y s i s o f the conodonts. The biota.  s e c t i o n s were a l s o a n a l y z e d i n terms of t h e i r l i t h o l o g y and o t h e r D e s c r i p t i o n s of the l i t h o l o g y were r e s t r i c t e d to the f i e l d  and a c l o s e i n s p e c t i o n o f hand specimens. prepared  notes  Although a few t h i n s e c t i o n s were  i t was d e c i d e d t h a t time was i n s u f f i c i e n t  to do an adequate study,  nor d i d i t seem n e c e s s a r y i n a p a l e o n t o l o g i c a l t h e s i s .  The remaining  biota  were i d e n t i f i e d a t h i g h taxonomic l e v e l s and used a s . a rough guide to changing b i o f a c i e s . l e s s e r degree,  Some b r a c h i o p o d genera were i d e n t i f i e d as they a i d e d , to a the age d e t e r m i n a t i o n s of the s t r a t a h e r e i n d e s c r i b e d .  10 STRATIGRAPHY AND  PALEOENVIRONMENT - GENERAL STATEMENT  A d i s c u s s i o n of the s t r a t i g r a p h y f o r the m a r g i n a l f a c i e s of the youngest Permian.on E l l e s m e r e I s l a n d f o l l o w s . the megascopic  T h i s c h a p t e r i s based  on  d e s c r i p t i o n of the s t r a t a and sediments, and on the mega-  b i o t a and t r a c e - f o s s i l s p r e s e n t .  The p a l e o e n v i r o n m e n t a l i n t e r p r e t a t i o n s  :  r e c o r d e d h e r e i n a r e not meant to be d e f i n i t i v e as they a r e founded on " rough d a t a .  The expected c h a r a c t e r i s t i c s f o r v a r i o u s environments  as  d e s c r i b e d i n Brenner and Davies (1974), Davies et a l . (1971), D i c k i n s o n et a l . (1972), G o l d r i n g and B r i d g e s (1973), Harms et a l . (1975), Howard (1972) and M i a l l  (1978) a r e summarized i n F i g u r e 2. Sabine Bay  Formation  A. Hamilton P e n i n s u l a a r e a Here the Sabine Bay Formation, which o v e r l i e s the B e l c h e r Channel Formation and o v e r l a i n by the A s s i s t a n c e , i s c h a r a c t e r i z e d by c y c l i c mentary environments .(Fig. 3 shows f e a t u r e s mentioned ;  in this  chapter).  The f o r m a t i o n c o n s i s t s of 180 metres of medium, c l e a n , w e l l f r i a b l e q u a r t z o s e sandstones w i t h some f i n e and c o a r s e sand and g r a n u l e s and p e b b l e s .  The sandstones a r e porous  (10 to 15%  sorted, sparse  estimated)  and u s u a l l y uncemented a l t h o u g h l o c a l c a l c i t e cement i s p r e s e n t . s u r f a c e s a r e g e n e r a l l y white to l i g h t b e i g e i n c o l o u r w h i l e  Fresh  weathered  s u r f a c e s a r e dominantly y e l l o w i s h brown to brownish orange but may medium brown and p a l e r e d or creamy p i n k . bedded t o massive but e x h i b i t f a i n t  also  The sandstones a r e g e n e r a l l y  i n t e r n a l laminae upon c l o s e r  Crossbedding i s not common but l o c a l l y c o n s p i c u o u s . a c o u p l e of dykes  sedi-  The sequence  be ' :  thick  examination. i s c u t by  (up to 3 m t h i c k ) , the d e l i n e a t i o n of which would be  im-  p o r t a n t i n terms of hydrocarbon p r e s e r v a t i o n as T h o r s t e i n s s o n (1974) r e p o r t s  11  shallow offshore  t r a n s i t i o n lower shoreface  upper shoreface  transition  foreshore  - f i n e to medium, clean| sandstone -well s o r t e d -trough crossbedding -minor amount of burrowing -beds t r u n c a ted -ripple laminae  - f i n e to medium, clean sandstone - w e l l sorted -trough crossbedding -minor amount of burrowing  -very clean, well sorted, f i n e to med ium sandstone| -low dipping, parallel to subparallel bedding -some convolute| laminae -no t r a c e fossils  - i n c r e a s i n g depth - m c r e a s i n l g d i s t a n c e from shore  t h i n beds \-f i n e , and - f i n e to mediuml few s e d i - c l e a n sand d i r t y sandmentary stone stone s t r u c t u r e s f i n e sand -30 to 45 cm siltstone beds t h i c k t h i c k beds extensive en upwards - p a r a l l e l bioturba- |-siltstone laminae tion inter-lens l i k e •low energy) bedded crossbedding ripple organic -abundant laminae rich large c ross conodonts distinct abundant bedding trace f o s s i l s rieogondol-j -mega ellids ripples occur i n deeper lower en^ ergy, lessfc nutrient r i c h environments thah i d i o g n a t h o d i d s and gn|a t h o d i d s .  t  d e p o s i t and sediment feeders burrows more horizon-| t a l and develop branches higher d i v e r s i t y F i g u r e 2.  1  -low d i v e r s i t y of burrows, s u s p e n s i o n feeders -unbranched, v e r t i c a l to s t e e p l y i n c l i n e d burrows  G e n e r a l i z e d l i s t of c h a r a c t e r i s t i c s f o r v a r i o u s s h a l l o w marine environments.  F i g u r e 3.  L i t h o l o g i c and b i o l o g i c c h a r a c t e r i s t i c s and s e c t i o n s p e r t i n e n t to t h i s r e p o r t .  c o r r e l a t i o n of the f o r m a t i o n s  and  13 bituminous, r e s i d u e s i n an o u t c r o p on Hamilton P e n i n s u l a . A c o q u i n o i d u n i t o f r u g o s o c h o n e t i d b r a c h i o p o d s o u t c r o p s 30 metres from t h e top o f t h e formation".  Coarse r i b b e d S p i r o p h y t o n i s p r e s e n t i n  t h i s u n i t and i n t h e o v e r l y i n g 30 m, but absent below. The type o f r i b b i n g o r laminae p r e s e n t on S p i r o p h y t o n seems to be v e r y u s e f u l f o r p a l e o e n v i r o n m e n t a l i n t e r p r e t a t i o n f o r t h e Permian Ellesmere.  r o c k s of  M a r i n t s c h and F i n k s (1978) i n a study of Devonian  (a t r a c e f o s s i l . s i m i l a r to Spirophyton) demonstrated  Zoophycus  environmental  signi-  f i c a n c e f o r t h e mean and maximum d i a m e t e r . o f t h e t r a c e and f o r t h e meniscus h e i g h t ( r e l a t e d t o r i b b i n g diameter o f the t r a c e and maximum body diameter o f t h e organism c r e a t i n g t h e burrow).  They found t h a t t h e animal  i s l a r g e s t near t h e c e n t r e o f i t s environmental range deep o f f s h o r e m a r i n e ) , s m a l l e s t near t h e margins  (quiet,  relatively  (shallower, higher  and absent i n t h e s h a l l o w e s t water beds w i t h i n t h e i r sequence.  energy)  Observa-  t i o n s f o r t h e Permian o f E l l e s m e r e suggest t h a t t h e c o a r s e r i b b e d , s m a l l e r diameter  (10 t o 20 cm) S p i r o p h y t o n a r e found i n s h a l l o w s h o r e f a c e e n v i r o n -  ments w h i l e t h e f i n e r i b b e d , l a r g e r diameter  (20 t o 35 cm) a r e found i n ."  deeper, q u i e t e r o f f s h o r e marine c o n d i t i o n s .  Any environments  interpreted  as f o r e s h o r e or t r a n s i t i o n a l between f o r e s h o r e and s h o r e f a c e do not c o n t a i n any S p i r o p h y t o n .  A p p a r e n t l y , S p i r o p h y t o n i s a l s o absent from t h e Van Hauen  and Degerbols Formations which a r e the b a s i n a l e q u i v a l e n t s o f t h e A s s i s tance and T r o l d F i o r d Formations.  Brachiopods were never found i n abun-  dance i n beds c o n t a i n i n g S p i r o p h y t o n a l t h o u g h a few may be p r e s e n t near by. Two o t h e r f e a t u r e s a r e noteworthy w i t h r e g a r d s t o t h e Sabine Bay F o r mation.  The f i r s t  i s an'unusual u n i d e n t i f i e d h e l i c a l burrow (5 t o 12 cm  diameter) found on bedding s u r f a c e s w i t h l a r g e s c a l e r i p p l e s  (wavelength =  0.9 to' 1.15 m, Amplitude = 20 t o 30 cm) and p r o b a b l y r e p r e s e n t i n g an upper  14  s h o r e f a c e environment.  These burrows a r e f a i r l y , evenly spaced  apart) suggesting high competition f o r resources. been found a t Hamilton Formation  P e n i n s u l a and McKinley  and a t H e n r i e t t a - N e s m i t h  These burrows have  Bay w i t h i n the Sabine Bay  ( F i g . 1) i n p o s s i b l e s h a l l o w water  e q u i v a l e n t s o f t h e T r o l d F i o r d Formation.  Secondly,  o t h e r m i c r o b i o t a o t h e r than palynomorphs a r e p r e s e n t B. McKinley  (0.3 m  no conodonts or any in this  formation.  Bay a r e a  Here 33 metres o f Sabine Bay Formation  unconformably o v e r l i e t h e Nan-  sen and a r e o v e r l a i n in. t u r n by a t h i n s e c t i o n o f t h e A s s i s t a n c e The  Sabine Bay begins w i t h d i r t y ,  f i n e to v e r y f i n e q u a r t z o s e  Formation.  sandstone  w i t h c o a r s e r i b b e d S p i r o p h y t o n and carbonaceous m a t e r i a l t o c l e a n , v e r y f i n e a r e n i t e w i t h a coquina o f r u g o s o c h o n e t i d . b r a c h i o p o d s i s i n t e r p r e t e d as a lower progrades  s h o r e f a c e environment.  a l l o f which  This unit  apparently  i n t o an upper s h o r e f a c e environment ( f i n e t o medium g r a i n e d ,  c l e a n q u a r t z o s e a r e n i t e s ) , which i n t u r n t r a n s g r e s s e s i n t o a lower s h o r e f a c e environment (brachiopod and b i v a l v e c o q u i n o i d q u a r t z o s e s t o n e ) , and f i n a l l y  progrades  very c l e a n quartzose a r e n i t e ) . C. Tanquary F i o r d The  i n t o a f o r e s h o r e environment  ( f i n e grained,  -  area  f o r m a t i o n ranges i n t h i c k n e s s from 36 to 70 metres, t h i n n i n g  towards t h e n o r t h and o n l a p p i n g t h e Tanquary s t r u c t u r a l h i g h . Bay  sand-  Formation  The Sabine  unconformably o v e r l i e s t h e Canyon F i o r d Formation  t u r n o v e r l a i n unconformably by t h e T r i a s s i c . B j o r n e Formation tance o r T r o l d F i o r d . : e q u i v a l e n t s a r e p r e s e n t .  and i s i n  as no A s s i s -  The Sabine Bay can be d i -  v i d e d i n t o t h r e e u n i t s I n c l u d i n g upper and'"lower,'uM'ts-of c l e a n , f i h e - t o medium g r a i n e d q u a r t z o s e a r e n i t e s r e p r e s e n t i n g s h a l l o w s h o r e f a c e t o f o r e -  15  shore environments, and a m i d d l e u n i t o f p o o r l y s o r t e d , v e r y f i n e  sand-  stone  to s i l t s t o n e and minor s h a l e w i t h a d i s t i n c t r o o t zone ( w i t h c o a l -  ified  r o o t s o r o t h e r p l a n t fragments) s u g g e s t i n g  marsh environment a t l e a s t p a r t l y emergent.  a b a c k b a r r i e r lagoon o r  Except f o r r o o t s no o t h e r mega-  f o s s i l s o r t r a c e s were observed. Assistance A. Hamilton P e n i n s u l a The  Formation  area  A s s i s t a n c e Formation a t Hamilton P e n i n s u l a  i n c l u d e s between 162 and  178 metres o f s e c t i o n depending on t h e p o s i t i o n o f t h e t a l u s - c o v e r e d dary w i t h t h e o v e r l y i n g T r o l d F i o r d Formation.  The lowest  part of the  A s s i s t a n c e c o n s i s t s of a f i n e grained, poorly sorted, quartzose w i t h carbonaceous m a t e r i a l , t r a c e f o s s i l s  boun-  sandstone  ( c o a r s e r i b b e d S p i r o p h y t o n and  S k o l i t h o s ) , minor rounded pebbles and fragmented b i o t a a t t h e top o f one bed,  c a l c i t e cement, and g l a u c o n i t e .  The A s s i s t a n c e i s s i m i l a r to p a r t s o f  the Sabine Bay, except f o r t h e g l a u c o n i t e . of g l a u c o n i t e i s abrupt,  t h e remaining  Although t h e appearance u p s e c t i o n  l i t h o l o g y suggests t h a t t h e boundary  between t h e A s s i s t a n c e and t h e u n d e r l y i n g Sabine Bay may be continuous.  Thorsteinsson  (1974) i n t e r p r e t s t h i s boundary as a  i t y which, i f p r e s e n t , must be o f s h o r t The  gradational'and disconform-  duration.  pebbly sandstone u n i t i s f o l l o w e d by s t r a t a t h a t t y p i c a l l y weather  y e l l o w i s h grey t o g r e y i s h orange w i t h f r e s h s u r f a c e s b e i n g v a r i o u s df grey,  and composed o f v e r y f i n e q u a r t z o s e  shades  sandstone to s i l t s t o n e  v a r i a b l e amounts o f c a l c a r e o u s  cement.  aceous m a t e r i a l , t r a c e f o s s i l s  (Asterosoma, f i n e r i b b e d Spirophyton,  lites  with  These r o c k s c o n t a i n abundant c a r b o n Plano-  (3 t o 5 mm; diameter) and o t h e r u n i d e n t i f i e d t y p e s ) , and abundant mega-  and m i c r o b i o t a . G l a u c o n i t e : i s p r e s e n t  i n a l l o f these r o c k s but never as  abundant as i n t h e o v e r l y i n g T r o l d F i o r d Formation.  A l a r g e part of the  16 s e c t i o n i s s o f t and f r i a b l e and was  g e n e r a l l y measured as cover o r t a l u s  w h i l e harder more c a l c a r e o u s sandstones stand out p r o m i n e n t l y . i s g e n e r a l l y t h i n t o medium but p o o r l y d e f i n e d .  Few  Bedding  o t h e r sedimentary  s t r u c t u r e s were observed a l t h o u g h carbonaceous m a t e r i a l i s o f t e n arranged as i r r e g u l a r l a m i n a t i o n s .  A l l f a c t o r s p o i n t to the p r e v a l e n c e of s h a l l o w  o f f s h o r e marine c o n d i t i o n s away from s h o r e f a c e environments few f i n e g r a i n e d , c l e a n e r q u a r t z o s e sandstone beds may t i o n a l beds between o f f s h o r e and lower s h o r e f a c e .  although a  represent t r a n s i -  The e x t e n s i v e b i o t u r b a -  t i o n and p r e s e a c e o f .- t f r e - a u t n i g e n i c ~ m i r i e r a l - g l a u c o n i t e suggest t h a t :  of . d e p o s i t i o n were .considerably l e s s than f o r the Sabine  Bay.  Conodonts a r e abundant i n the lower h a l f of the f o r m a t i o n and N e o g o n d o l e l l a i d a h o e n s i s n.subsp.  A and Anchignathodus  rates  minutus.  include  The Neogon-  d o l e l l a fauna i s v e r y abundant and i n c l u d e a s i g n i f i c a n t p r o p o r t i o n of comp l e t e specimens.  A l a r g e number o f ramiform elements o c c u r i n a s s o c i a t i o n  with Neogondolella. Formations has none.  No o t h e r assemblage  i n the A s s i s t a n c e or T r o l d  Fiord  as many ramiforms compared to p l a t f o r m s ; i n f a c t , most had  The c o l o u r s of these conodonts a r e brown to dark brown and have an  a l t e r a t i o n index o f 2.0  a c c o r d i n g t o E p s t e i n et a l . (1977).  This  indicates  metamorphic temperatures o f 60 to 140°C and a f i x e d carbon range o f 55 t o 70%, w e l l w i t h i n the l i m i t s f o r petroleum p r e s e r v a t i o n . upper h a l f o f the f o r m a t i o n a r e fragmented and a c o u p l e of ramiform  Conodonts i n the  and r a r e and i n c l u d e N.  serrata(?)  fragments.  B. McKinley Bay a r e a The A s s i s t a n c e Formation, which was  not p r e v i o u s l y r e c o g n i z e d a t Mc K i n -  l e y Bay, i s d e f i n e d here as a t h i n (3 to 4 m) u n i t of. g r e y i s h y e l l o w weatheri n g , f i n e to medium g r a i n e d q u a r t z o s e sandstone f o l l o w e d by a u n i t of m a t r i x supported, dark grey c h e r t - p e b b l e conglomerate.  Large b r a c h i o p o d s and  bryozoan fragments cite.  o c c u r w i t h i n t h i s u n i t which -is v a r i a b l y cemented by  R e c r y s t a l l i z e d conodonts  ( p o s s i b l y as a r e s u l t of i n t e n s e h e a t i n g  by a nearby dyke) were found i n one sample (F100) and i d a h o e n s i s and Neostreptognathodus tognathodus  fragments  prayi.  included Neogondolella  The o c c u r r e n c e  i s unique to t h i s sample.  of two  Neostrep-  C l a r k (1974) i n d i c a t e d  gnathodids t h r o v e i n v e r y s h a l l o w n u t r i e n t r i c h water of moderate and normal  perhaps a t the l i m i t of the p h o t i c zone.  c a t i o n s from t h e l i t h o l o g y or a s s o c i a t e d b i o t a a t F100 a t e s i z e d p r o d u c t i d s and environments  small s p i r i f e r i d s ) ,  There a r e no  indi-  (dominantly moder-  o c c u r r e n c e s i n the  The most s i g n i f i c a n t d i f f e r e n c e i s the  p o s i t i o n w i t h i n the b a s i n t o - t h e extent t h a t the McKinley Bay  section i s  c l o s e r t o the b a s i n . m a r g i n than the Hamilton P e n i n s u l a s e c t i o n s . the obvious c y c l i c i t y of environments gnathodus specimens were found.  ' .  to suggest t h a t t h e r e p r e s e n t e d  a r e s h a l l o w e r than those of o t h e r conodont  Assistance at.Hamilton Peninsula.  that  energy  s a l i n i t y , whereas g o n d o l e l l i d s arid a n c h i g n a t h o d i d s p r e f e r r e d  deeper water,  cal-  a t Hamilton P e n i n s u l a no  Despite  Neostrepto-  Those samples w i t h N e o g o n d o l e l l a were  n e a r l y always a s s o c i a t e d w i t h 3 to 5 mm  diameter P l a n o l i t e s ; u n t r a n s p o r t e d  megafauna i n t e n s e l y bored, by an e n d o l i t h i c c h l o r o p h y t e a l g a i n d i c a t e a low energy,  s h a l l o w marine environment  v a t i o n s and  w e l l w i t h i n the p h o t i c zone.  i n t e r p r e t a t i o n s suggest t h a t e i t h e r the d i f f e r e n t  These o b s e r -  environmental  c o n d i t i o n s c o n t r o l l i n g the d i s t r i b u t i o n of these two genera a r e s u b t l e and as yet u n r e c o g n i z e d f o r t h i s a r e a o r t h a t Neostreptognathodus  was  very rare  i n the Permian o f the Sverdriip B a s i n . D. Sawtooth Range a r e a T h i s s e c t i o n of A s s i s t a n c e r o c k s i s v e r y t h i c k (515 to 545 m a f t e r r e moving the 88 m s i l l )  and can be d i v i d e d  i n t o two u n i t s .  The lower 450  m  18  t h i c k u n i t i s c h a r a c t e r i z e d by y e l l o w - g r e y ose sils  weathering, f i n e g r a i n e d  quartz-  sandstone, w i t h v a r i a b l e amounts of carbonaceous m a t e r i a l , t r a c e f o s ( S k o l i t h o s and c o a r s e  ribbed Spirophyton),  cements and p r a c t i c a l l y no m e g a f o s s i l s .  calcareous  and/or  siliceous  Pendants w i t h i n t h e s i l l  and a few  beds above i t contain, b r a c h i o p o d s (Jakutoproductus(?) f i c a n c e ) and s m a l l p e l e c y p o d s .  - see p.32 f o r s i g n i -  As g l a u c o n i t e i s not p r e s e n t  rocks r e l a t i v e l y r a p i d d e p o s i t i o n i s implied.  The proposed  i n any o f these paleoenviron-  mental i n t e r p r e t a t i o n f o r t h i s u n i t i s o f some s o r t o f a d e l t a complex. d e p o s i t i o n i s i n t e r p r e t e d as o c c u r r i n g a t t h e d e l t a f r o n t i n lower and  t r a n s i t i o n a l environments.  to s e c t i o n s t o t h e n o r t h e a s t ,  The t h i c k n e s s o f these d e p o s i t s  The  shoreface  compared  t h e r a p i d i t y o f d e p o s i t i o n , dominance o f dep-  o s i t f e e d e r s , and t h e abundance o f carbonaceous m a t e r i a l a l l c o n f i r m a deltaic  environment (Weimer, 1970).  The upper 95 m t h i c k u n i t i s c h a r a c t e r i z e d by f i n e t o v e r y grained,  fossiliferous  (almost  coquinoid),  quartzose  amounts o f carbonaceous m a t e r i a l and g l a u c o n i t e . t e n s e l y burrowed  ( f i n e ribbed Spirophyton,  p l a t y bedding and a r e v e r y Hamilton P e n i n s u l a .  fine  sandstone w i t h  varying  These sediments a r e i n -  and Asterosoma), have i r r e g u l a r  s i m i l a r t o t h e l i t h o l o g y ..of t h e A s s i s t a n c e a t  The presence o f abundant b r a c h i o p o d s and bryozoans as  w e l l as g l a u c o n i t e suggests lower energy c o n d i t i o n s and slower d e p o s i t i o n . Apparently  t h i s u n i t i s t r a n s g r e s s i v e over the lower u n i t .  Transgressions  a r e o f t e n i n i t i a t e d when a l l o r p a r t o f a d e l t a system i s abandoned so t h a t subsidence i n c r e a s e s r e l a t i v e to t h e d e p o s i t i o n . T r o l d F i o r d Formation A. Hamilton P e n i n s u l a The  area  T r o l d F i o r d Formation i s c h a r a c t e r i z e d by g l a u c o n i t e r i c h ,  fine  quartzose stone,  sandstones b u t . a l s o c o n s i s t s of minor b i o g e n i c arenaceous l i m e -  chert-pebble  spicules).  conglomerate, and  chert  (major components a r e sponge  On the b a s i s of r e g i o n a l o v e r s t e p p i n g  o l d e r formations  from NW  to SE, T h o r s t e i n s s o n  (1974) i n d i c a t e d a  i t y at the boundary between the A s s i s t a n c e and However, t h i s a u t h o r saw Peninsula.  Unless  no evidence  disconform-  T r o l d F i o r d Formations.  f o r such an i n t e r p r e t a t i o n at Hamilton  the seas t o t a l l y v a c a t e d  s e c t i o n s to show a n e a r l y continuous  by the T r o l d F i o r d on  the b a s i n one would expect some  r e c o r d w h i l e o t h e r s may  show a major  hiatus. The  lowest  by s i l i c i f i e d  p a r t s of the T r o l d F i o r d ( S u b d i v i s i o n D) a r e c h a r a c t e r i z e d  coquinas of b r a c h i o p o d s i n a g l a u c o n i t i c , f i n e  quartzose  sandstone which c o n t a i n s o n l y minor amounts of carbonaceous m a t e r i a l ( i n d i s t i n c t i o n to the more carbonaceous A s s i s t a n c e )  presumably due  i n t r o d u c t i o n of t e r r i g e n o u s p l a n t m a t e r i a l f u r t h e r o f f s h o r e . which a r e t y p i c a l l y of shallow s m a l l , unfragmented, and  to  The  decreased coquinas,  s u b t i d a l o r i g i n , a r e composed of dominantly  sometimes a r t i c u l a t e d p r o d u c t i d s , . a n d  of fragmented l a r g e p r o d u c t i d s and  spiriferids.  l e s s e r amounts  Other b i o t a form o n l y a  s m a l l f r a c t i o n of the t o t a l biomass w h i l e conodonts a r e a p p a r e n t l y  absent.  T h i s u n i t weathers dusky y e l l o w w i t h minor r e d w h i l e f r e s h s u r f a c e s g r e y i s h yellow-green  and  are  only o c c a s i o n a l l y red.  The m a j o r i t y o f the g l a u c o n i t e formed i n the s m a l l pores of echinoderm fragments and  p r o g r e s s i v e l y r e p l a c e d the s t r u c t u r e u n t i l a s o l i d b l e b of  glauconite resulted. and  G l a u c o n i t e a l s o formed i n the chambers of s m a l l forams  i n the z o o e c i a of bryozoa.  I t i s g e n e r a l l y regarded  that  forms by the a l t e r a t i o n of k a o l i n i t e c l a y s i n l o c a l l y r e d u c i n g ( p r o v i d e d by the s m a l l pores of b i o t i c  glauconite conditions  elements, presumably owing to  con-  20  c e n t r a t i o n s of d e c a y i n g o r g a n i c matter) but i n a g e n e r a l l y o x i d i z i n g environment  ( p r o v i d e d by a c t i o n o f s h a l l o w marine waves).  As w e l l a s b e i n g  most common on t h e o u t e r edge o f t h e s h e l f and on t o p o g r a p h i c h i g h s , g l a u c o n i t e i s i n v a r i a b l y a s s o c i a t e d w i t h low s e d i m e n t a t i o n r a t e s because o f the:, low c o n c e n t r a t i o n o f s o l u b l e i r o n i n t h e ocean and because t h e g l a u c o n i t i z a t i o n p r o c e s s stops a f t e r b u r i a l owing t o t h e l o s s of t h e proper  chemical  environment ( B u r s t , 1958). A 5% metre u n i t o f c h e r t g r a n u l e and pebble conglomerate  i s a char-  a c t e r i s t i c p a r t of S u b d i v i s i o n E . i n t h e middle of t h e T r o l d F i o r d The lower  Formation.  f i v e metres a r e c o n c e n t r a t e d i n l e n s e s o r channels and grade  l a t e r a l l y i n t o burrowed (8 t o 10 mm diameter g r a i n e d g l a u c o n i t i c q u a r t z o s e sandstone.  P l a n o l i t e s ) , v e r y f i n e to f i n e  The upper 0.5 metres i s r e p r e s e n t e d  by a s o l i d bed o f r e d and y e l l o w c h e r t pebble conglomerate sent a t r a n s g r e s s i v e l a g d e p o s i t . are present i n t h i s i n t e r v a l .  which may r e p r e -  Large unbroken p r o d u c t i d s (Thamnosia)  D e s p i t e t h e i r t h i c k s h e l l s , t h e energy  of the  d e p o s i t i o n a l environment must have been low f o r t h e s h e l l s t o remain unbroken, which seems somewhat anomalous i n l i g h t o f t h e g r a i n s i z e .  The b r a c h -  iopods a r e a l s o i n t e n s e l y bored by p o l y c h a e t e s and. b a r n a c l e s s u g g e s t i n g t h a t they were exposed t o t h e marine environment f o r a s i g n i f i c a n t  time  b e f o r e t h e i r i n c o r p o r a t i o n i n t o t h e conglomerate.  which  The conodonts,  a r e r a r e and fragmented i n t h i s u n i t , i n c l u d e N e o g o n d o l e l l a n.sp.  13 and N.  postserrata(?). The upper p a r t s of'.the T r o l d F i o r d of s i l i c i f i e d  brachiopods  s i m i l a r t o those lower  beds a r e dominated by bryozoans d r i c a l trepostomes  ( S u b d i v i s i o n ! F) b e g i n w i t h  comprise  i n the section.  coquinas Some  where these e n c r u s t i n g s h e e t - l i k e and c y l i n -  up t o 50% o f t h e r o c k volume.  A minor amount o f  b l u e - g r e y c h e r t a l s o occurs, i n t h i s i n t e r v a l .  Carbonaceous  a r e s p o r a d i c and u s u a l l y a s s o c i a t e d w i t h burrowing.  b l e b s and  The sandstones  out t h i s u n i t are. v e r y f i n e to f i n e g r a i n e d and moderately t o w e l l Weathering  films  throughsorted.  c o l o u r s i n c l u d e g r e y i s h orange, g r e y i s h y e l l o w and g r e y i s h y e l l o w  -green whereas f r e s h s u r f a c e s a r e g r e y i s h o l i v e to dusky y e l l o w - g r e e n . Abundance o f conodonts. i n t h i s u n i t dolella  n.subsp.  A assemblages.  populations are sparse. which a r e complete,  Between these two assemblages  The conodonts  B.  conodont  i n t h i s T r o l d F i o r d u n i t , some of  i n c l u d e N e o g o n d o l e l l a b i t t e r i n.subsp.  e l l a r o s e n k r a n t z i n.subsp. ramiform element was  is.comparable to t h a t of Neogon-  Only one s m a l l fragmented  found i n a s s o c i a t i o n .  C and  and  Neogondol-  unidentified  C o l o u r s of these conodonts  almost o r i g i n a l amber and have a c o l o u r a l t e r a t i o n index o f 1.5 brown) a c c o r d i n g to E p s t e i n et a l . (1977).  are  (very p a l e  T h i s i n d i c a t e s metamorphic  temperatures of 50 t o 90°C and a f i x e d carbon range of 55 to 70%, w i t h i n the l i m i t s f o r petroleum g e n e r a t i o n and  well  preservation.  The h i g h e s t beds o f the T r o l d F i o r d a r e burrowed  (medium r i b b e d  S p i r o p h y t o n ) , f i n e g r a i n e d , q u a r t z o s e sandstone w i t h s m a l l s p i r i f e r i d s and p h o s p h a t i c nodules c o n t a i n i n g moderately l a r g e i n a r t i c u l a t e b r a c h i o p o d s ( L i n g u l a ) and f i s h d e b r i s .  The environments  r e p r e s e n t e d by t h i s u n i t a r e  l a r g e l y s h a l l o w s u b t i d a l marine to p o s s i b l y t r a n s i t i o n a l s h o r e f a c e a t the top.  This unit  i s o v e r l a i n unconformably  by the T r i a s s i c B j o r n e  sandstone.  B. McKinley Bay a r e a Here the T r o l d F i o r d Formation ranges i n t h i c k n e s s from 24 t o 61 over a d i s t a n c e of 1.6  km  (1 m i l e ) .  The base of the f o r m a t i o n c o n s i s t s of  one metre o f r e d weathering c h e r t g r a n u l e and pebble conglomerate l a r g e s p i r i f e r i d s and moderate s i z e d p r o d u c t i d s (Thamnosia?). conglomerate,  silicified  metres  with  Above the  coquinas of b r a c h i o p o d s (mostly of s m a l l s i z e )  22  o c c u r i n a v e r y f i n e , g r a i n e d sandstone to arenaceous p a r t of t h i s u n i t demonstrates  carbonate.  f e a t u r e s s u g g e s t i n g hardground  At  least  development;  b r a c h i o p o d coquinas (most a r e i n l i f e p o s i t i o n ) t o t a l l y surrounded by c r u s t i n g trepostome  bryozoans w i t h the z o o e c i a l a t e r f i l l e d  en-  with glauconite.  The b r a c h i o p o d s a r e i n t e n s e l y bored by e n d o l i t h i c a l g a e and a c r o t h o r a c i c a n b a r n a c l e s , the d e n s i t y of which may time.  be an i n d i c a t o r of r e l a t i v e  exposure  The remainder o f the s e c t i o n c o n s i s t s o f f i n e g r a i n e d , g l a u c o n i t i c ,  q u a r t z o s e sandstones w i t h minor burrowing and g a s t r o p o d s .  (Spirophyton) and r a r e  brachiopods  I n a r t i c u l a t e b r a c h i o p o d s c h a r a c t e r i z e the h i g h e s t beds.  Only a c o u p l e of u n i d e n t i f i a b l e N e o g o n d o l e l l a fragments were found a t  this  locality. D.  Sawtooth Range a r e a T h i s sequence of T r o l d F i o r d s t r a t a i s t h i c k e r and c o n t a i n s l e s s , b i o t a  then the Hamilton P e n i n s u l a s e c t i o n .  The lower p a r t s o f the f o r m a t i o n a r e '  c h a r a c t e r i z e d by f i n e to v e r y f i n e g l a u c o n i t i c , q u a r t z o s e sandstones s h a l y l a y e r s , minor burrowing, and r a r e b r a c h i o p o d s .  with  The few b r a c h i o p o d  h o r i z o n s t h a t do o c c u r have been l e a c h e d o f a l l o r i g i n a l s h e l l . 169 metres a r e p a r t i c u l a r l y low i n b i o c l a s t i c d e b r i s .  The lowest  Most of t h i s  interval  weathers dark green and has more g l a u c o n i t e than quartz g r a i n s , however, two  samples (L38 and L46) weathered  glauconite.  b l u e - g r e y i n c o l o u r and had f a r l e s s  The cements i n t h e s e sandstones are"dominated  by s i l i c a  but  minor c a l c i t e i s a l s o p r e s e n t . Once a g a i n the m i d d l e p a r t of the T r o l d F i o r d g r a n u l e and pebble conglomerates.  The conglomerate  i s c h a r a c t e r i z e d by  chert  i n t e r v a l i s 20 metres  t h i c k but d i s c o n t i n u o u s as the g r a n u l e s and pebbles a r e c o n c e n t r a t e d i n l e n s e s or c h a n n e l s .  The m a j o r i t y of t h i s u n i t  i s r e d weathering and  has  23/.; abundant abraded  and bored b i o c l a s t s  ( p o l y c h a e t e s , sponge(?), and b a r n a c l e  b o r i n g s ) i n c l u d i n g a few Thamnosia and many l a r g e s p i r i f e r i d s . Planolites  (8 to 10 mm  are also present.  d i a m e t e r ) , i d e n t i c a l to those a t Hamilton P e n i n s u l a ,  Above the c o n g l o m e r a t i c u n i t a r e about 40 metres of  stone, s i m i l a r to the lowest 169 metres. consist,  Large  The next 56 metres of  sand-  section  of f i n e g r a i n e d , v a r i a b l y g l a u c o n i t i c q u a r t z o s e sandstones t h a t  .  weather green w i t h o c c a s i o n a l p u r p l i s h l a y e r s , l a c k i n g i n carbonaceous m a t e r i a l , and c o n t a i n a few s m a l l P l a n o l i t e s burrows.  There appears to be  e v i d e n c e f o r c h a n n e l i n g i n t h e s e sandstones as coquinas o f b r a c h i o p o d s a r e lens-like in distribution.  The next 30 metres c o n s i s t of f i n e g r a i n e d  sandstone and minor dark g r e e n i s h grey c h e r t t h a t would be b e s t d e s c r i b e d as.a s p i c u l i t e .  The s e c t i o n ends w i t h monotonous l i g h t green weathering,  f i n e g r a i n e d q u a r t z o s e sandstone which,  except f o r one l o c a l i t y w i t h  t i c u l a t e b r a c h i o p o d s , i s g e n e r a l l y d e v o i d o f megabiota. to d i s t i n g u i s h p a r t i c u l a r environments  It i s d i f f i c u l t  but the m a j o r i t y of the s e c t i o n i s  undoubtedly of s h a l l o w s u b t i d a l s h e l f o r i g i n .  I t i s p e r p l e x i n g why b r a c h i o -  pod l o c a l i t i e s a r e so few d e s p i t e the slow d e p o s i t i o n a l r a t e s denoted the presence of abundant No  samples  largely  faunas were p r e s e n t . AGE  AND  CORRELATION  A - . c o r r e l a t i o n c h a r t f o r Permian 4;  by  glauconite.  from t h i s s e c t i o n were p r o c e s s e d f o r conodonts,  because no s i l i c i f i e d  inar-  stages and zones i s p r o v i d e d i n F i g u r e  i t i n c l u d e s schemes from Grant and Cooper ( 1 9 7 3 ) , . F u r n i s h (1973), Ward-  law and C o l l i n s o n  (1979a) y Water house (197.6), and the scheme adopted f o r  t h i s study which i s a combination o f the o t h e r s . The r e s e a r c h conducted f o r t h i s study has s i g n i f i c a n t l y r e f i n e d the age  AA  BB  CCl  l  Dorashamian Djulfian  Changhsinglan Chhidruan Araksian  Tartarian  Ochoan  Changhsingian Chhidruan Araksian  •H  a)  3  N  C  X  Dorashamian  TH  u a) cn  >u tH  Djulfian Amarassian Capitanian  Amarassian Capitanian  Amarassian Capitanian  Punjabian  EE  PD  CC2  Zone 6 ttj •rH  Kazanian  Kazaniau  Zone 5  p.  3  to nj (V n) u  iH  Guadalupian Wordian  Wordian  5  01  tn  Wordian  Roadian  Zone 3  Roadian  Road l a n  Subdivision  E  Subdivision  D  Subdivision  C  Subdivision  B  Subdivision  A  Assistance Formation  'Zone 2 X Zone 1  a  F  Trold Fiord Formation  barren interval  Kungurian  Kungurian (Ufimian)  M  Zone A  Subdivision  n)  •H  Baigendzinian  Leonardian  Leonardian A r t i n s k i a n Leonard i a n  Sakmarian  Sakmarian Asselian  F i g u r e 4.  Asselian  •H  H <  Aktastinian Sterlitamakian Tastubian  Aktastinian Sterlitamakian Tastubian  Ai tn  Wolfcampian  Leonard i a n  tn tu to  a N. i d a h o e n s i s e N. p r a y i (0 X. a b s t r a c t u s V X. t r i b u l o s u s P N. b i t t e r ! 8 N. r o s e n k r a n t z i * N. p o s t s e r r a t a X N. s e r r a t a  S a b i n e Bay Formation  Aktastinian  Sakmarian  Asselian  C o r r e l a t i o n c h a r t f o r s e r i e s , s t a g e s and z o n e s o f t h e P e r m i a n . L = L o w e r P e r m i a n , M W a t e r h o u s e ( 1 9 7 5 ) , BB = F u r n i s h ( 1 9 7 3 ) , CC = Moore e t a l . ( 1 9 6 5 ) 1 = E u r o p e , 2 AA This study. EE  M i d d l e P e r m i a n , U « Upper P e r m i a n N o r t h A m e r i c a , DD = W a r d l a w a n d C o l l i n s o n  (1979a,b).  25  r e l a t i o n s h i p s of t h e A s s i s t a n c e and T r o l d F i o r d Formations through the use of conodonts as the b i o s t r a t i g r a p h i c it  index.  Before d i s c u s s i n g the r e s u l t s  seems a p p r o p r i a t e t o r e l a t e the s t a t e of t h e a r t p r i o r to t h i s  R e l a t i v e V a l u e of V a r i o u s F o s s i l  research.  Biota  F i v e groups have l e d the way over a l l  o t h e r s f o r the d e t e r m i n a t i o n of  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 s o f t h e Permian. .These i n c l u d e b r a c h i o p o d s , ammonoids, f u s u l i n i d s , palynomorphs and conodonts. B r a c h i o p o d s were the predominant marine megafauna o f t h e Permian. I t i s because o f t h i s dominance  t h a t t h e y can be used t o c o r r e l a t e more  r o c k s and on a wider b a s i s than any o t h e r group. a u t h o r s t h a t as b r a c h i o p o d s a r e l a t i t u d i n a l l y  I t i s the o p i n i o n of many  (climatically)  controlled  and r e l a t i v e l y l o n g r a n g i n g , t h e i r c o r r e l a t i o n . p o t e n t i a l i s d e c r e a s e d . However, Waterhouse  (1976) s t a t e s t h a t t h i s i s a w i d e l y r e i t e r a t e d  misap-  p r e h e n s i o n and t h a t brachiopod s p e c i e s and genera, d u r i n g t h e Permian, were l e s s l a t i t u d i n a l l y o r f a c i e s c o n t r o l l e d than f u s u l i n i d s o r ammonoids, and j u s t as s h o r t - l i v e d .  Even i f Waterhouse  i s proven c o r r e c t by t h i s  '  statement, t h e extreme d i v e r s i t y of the group makes worldwide mastery of t h i s group, e s p e c i a l l y a t t h e s p e c i f i c l e v e l , v e r y d i f f i c u l t Waterhouse  indeed.  (1976) a l s o i n d i c a t e s t h a t c e r t a i n c o r r e l a t i o n problems w i l l  not be s o l v e d u n t i l p a l e o n t o l o g i s t s i n the USSR reexamine b r a c h i o p o d u l e s i n the Permian type or standard s e c t i o n s of the U r a l s . t i o n s p e r t i n e n t t o t h i s study b r a c h i o p o d s ( p r o d u c t i d s and  In the forma-  spiriferids)  a r e by f a r the dominant b i o t a but need to be s t u d i e d a t t h e s p e c i f i c s i n c e many genera range throughout the e n t i r e s e c t i o n .  faun-  level  Because of the  problems a t the s p e c i f i c l e v e l , age d e t e r m i n a t i o n s have been of minor v a l u e o r c o n f l i c t i n g to o f f i c e r s of the GSC concerned w i t h . t h e C a r b o n i f e r o u s and  26  Permian o f A r c t i c  Canada.  Ammonoids a r e v e r y s h o r t - l i v e d , o f t e n even a t t h e g e n e r i c l e v e l , thus prove of i n f i n i t e v a l u e f o r c o r r e l a t i o n s  ( F u r n i s h , 1973).  and  However,  t h e i r v a l u e i s q u i c k l y d i m i n i s h e d when one c o n s i d e r s the r a r i t y of t h e i r : occurrence.  A c c o r d i n g to Waterhouse (1976; f i d e . R.E.  d e t a i l e d s t u d i e s i n West Texas produced  Grant, p e r s . comm.)  o n l y 5,000 ammonoids from  l o c a l i t i e s compared to some 3,000,000 brachiopods from about Waterhouse (1976) a l s o i n d i c a t e d t h a t i n over 1500 Yukon T e r r i t o r y o n l y f i v e y i e l d e d ammonoids. found p r e v i o u s l y i n the Sabine Bay,  800  97 localities.  l o c a l i t i e s from the  Although ammonoids have been  A s s i s t a n c e and T r o l d F i o r d  (however, r a r e l y ) t h i s author found none i n h i s d e t a i l e d  Formations  sections.  F u s u l i n i d s a r e s h o r t - l i v e d and o f t e n , when p r e s e n t , as abundant as brachiopods.  In o p p o s i t i o n to t h e s e p o s i t i v e a s p e c t s i s the s t r o n g l a t i -  t u d i n a l or c l i m a t i c r e s t r i c t i o n to t h e i r d i s t r i b u t i o n . generally restricted  F u s u l i n i d s are  to warm waters and a l t h o u g h p r e s e n t i n the C a r b o n i -  f e r o u s and E a r l i e s t Permian of E l l e s m e r e I s l a n d they a r e absent from the l a t e Lower and M i d d l e Permian s e c t i o n s covered by t h i s  report.  C a r b o n i f e r o u s and T e r t i a r y systems have l o n g dominated  palynological  s t u d i e s a l t h o u g h the Permian i s becoming i n c r e a s i n g l y important Jansonius,  (Hart,  1965;  1962).  Palynomorphs c o u l d prove v e r y v a l u a b l e f o r c o r r e l a t i o n of those p a r t s of s e c t i o n s where o t h e r b i o t a a r e r a r e or absent. A s s i s t a n c e Formation a t Hamilton dominated age.  by V i t t a t i n a which,  Two  samples from the  -•Peninsula y i e l d e d palynomorph assemblages  a c c o r d i n g to Hart  (1965), i s Kungurian i n  T h i s form i s s i m i l a r t o V. simplex d e s c r i b e d by J a n s o n i u s (1962)  from the Permian B e l l o y Formation of the Peace R i v e r a r e a , Canada.  Two  27  samples from t h e T r o l d F i o r d Formation a t Hamilton P e n i n s u l a y i e l d e d ' V . V i t t a t i n a c f . V. l a t a which was f i r s t d e s c r i b e d from t h e Guadalupian Flowerpot Formation o f Oklahoma, USA (Wilson, 1962).  These f l o r a l  differ-  ences suggest t h a t t h e s e palynomorphs may be u s e f u l f o r d i s c r i m i n a t i n g A s s i s t a n c e and Trold. F i o r d e q u i v a l e n t s .  These palynomorphs were found i n  the same samples, as t h o s e c o n t a i n i n g conodonts.  A c o r r e l a t i o n scheme  combining t h e d i s t r i b u t i o n s o f b o t h o f t h e s e groups would be u s e f u l f o r r e s o l v i n g s t r a t i g r a p h i c problems.in c o r r e l a t i v e rocks a t Henrietta-Nesmith (Fig.  1) which l a c k conodonts e n t i r e l y . Furthermore, palynomorphs  (as w e l l as conodonts) can be used to e s t i -  mate t h e temperatures t h a t t h e i r host s t r a t a were s u b j e c t e d t o . The two A s s i s t a n c e samples had an average Thermal A l t e r a t i o n Index (TAI) o f between 2.8 and 3.0 ( S t a p l i n 1969, 1974) whereas t h e T r o l d F i o r d samples a v e r aged 2.8.  According to Epstein et a l .  (1977) conodont c o l o u r  does n o t b e g i n u n t i l l a t e stages o f palynomorph d i a g e n e s i s ;  explaining  the v e r y minimal d i f f e r e n c e s f o r palynomorph.TAI's u p s e c t i o n . to S t a p l i n  alteration  According  (1969, 1974). T A I s on t h e o r d e r o f 2.8 i n d i c a t e hydrocarbon 1  p o t e n t i a l f o r o i l and wet gas and a mature o r g a n i c metamorphic  facies.  Temperatures o f 100°C a r e t y p i c a l f o r t h i s f a c i e s which, f a l l s w i t h i n t h e range o f 50 t o 140°C suggested by t h e conodonts.  Conodont c o l o u r  alter-  a t i o n indexes o f 1.5 t o 2.0 and palynomorph TAI's of 2.8 t o 3.0 a r e e n t i r e l y c o n s i s t e n t w i t h comparisons p r e s e n t e d ' i n E p s t e i n e t a l . (1977). Conodonts, compared  to t h e s e o t h e r groups, a r e s t i l l  i n their  i n terms o f t h e i r u s e as a b i o s t r a t i g r a p h i c index f o r t h e Permian.  infancy As  many problems a r o s e from t h e e a r l y i n t e n s i v e study o f Permian conodonts Waterhouse  (1976) was.;led t o express h i s doubt t h a t they w i l l  ever be a b l e  28  to p r o v i d e worldwide c o r r e l a t i o n f o r :the marine Permian.  In f a c t ,  conodonts have been found from the c o l d water Permian of east despite i n t e n s i v e search ( N i c o l ,  1975).  no  Australia  However, b i o s t r a t i g r a p h i c  schemes  based on conodonts have improved s u b s t a n t i a l l y d u r i n g the l a s t f i v e y e a r s , l a r g e l y t h r o u g h t h e work of Clark,,Behnken, Wardlaw and C o l l i n s o n . f a c t i n c o m b i n a t i o n w i t h the abundance  This  and e x c e l l e n t p r e s e r v a t i o n of  conodonts found i n t h e c o l d water faunas of the A s s i s t a n c e and T r o l d Formations ( t h i s study) which a r e c l o s e l y a l l i e d West Texas, Wyoming, Montana,  Fiord  to warm water faunas of  Utah, and Idaho ( t h i s cannot be s a i d f o r the  b r a c h i o p o d s from t h e same a r e a s ) j u s t i f y optimism t h a t conodonts w i l l enjoy a v e r y b r i g h t f u t u r e f o r w o r l d w i d e . c o r r e l a t i o n of the Permian. P r e v i o u s Conodont Work The o n l y p r e v i o u s Permian conodont work on E l l e s m e r e I s l a n d was based on a s i n g l e sample from the A s s i s t a n c e Formation on H a m i l t o n P e n i n s u l a (near the p o s i t i o n of F54) and a s i n g l e sample from t h e base of the Degerb o l s Formation ( b a s i n a l e q u i v a l e n t o f the T r o l d F i o r d Formation) from near Otto F i o r d as summarized  by Kozur and Nassichuk (1977).  The A s s i s t a n c e sample y i e l d e d s i x conodont specimens, t h r e e o f which were a s s i g n e d to N. i d a h o e n s i s and the remainder to an. i n t e r m e d i a t e p o s i t i o n between N. i d a h o e n s i s and N. s e r r a t a or N. n a n k i n g e n s i s .  These samples  seem v e r y much l i k e the p o p u l a t i o n samples d e f i n e d i n t h i s r e p o r t as N. i d a h o e n s i s n.subsp. A  which a r e i n t e r m e d i a t e between N. s e r r a t a (not N.  n a n k i n g e n s i s s i n c e s e r r a t i o n s a r e not p r e s e n t on the p o s t e r i o r p a r t s ) and N. i d a h o e n s i s .  The a u t h o r s ( i b i d . ) p l a c e d t h e i r fauna i n the Upper Roadian  ( t h i s r e p o r t p l a c e s i t i n t h e Lower R o a d i a n ) . The Degerbols  sample y i e l d e d s e v e r a l fragmentary conodont  specimens  29 a l l b e l o n g i n g to s i n g l e . s p e c i e s N. c f . N. g r a c i l i s and p o s s i b l y i n t e r m e d i a t e between _N.. i d a h o e n s i s and N. g r a c i l i s t o which the a u t h o r s a s s i g n e d an> Upper Roadian age stricto).  (younger than type Roadian but o l d e r than Wordian sensu  As i n d i c a t e d  i n t h i s r e p o r t , forms s i m i l a r , to N. g r a c i l i s occur  i n p o p u l a t i o n s of N. i d a h o e n s i s n.subsp. Roadian.  A which a r e a s s i g n a b l e to the Lower  However, as these forms a r e few i n number they a r e t h e r e f o r e  l i k e l y to be the o n l y forms p r e s e n t i n a s m a l l sample. it  As i n d i c a t e d  un-  earlier  seemed r e a s o n a b l e to suggest t h a t the v a r i e t i e s w i t h i n N. i d a h o e n s i s  n.subsp.  A, i f s e p a r a t e d as p e r i p h e r a l i s o l a t e s c o u l d , f o l l o w i n g  l e a d t o p o p u l a t i o n s o f _N. g r a c i l i s .  speciation,  I f t h i s i s indeed the case then the  Upper Roadian age i s e n t i r e l y c o n s i s t e n t .  However, t h e r e may  be some e c o l -  o g i c a l r e q u i r e m e n t s t h a t i n c r e a s e the number of one v a r i e t y or another i n different  environments.  In o t h e r words, i t cannot be d i s c o u n t e d t h a t  g r a c i l i s v a r i e t y of _N. i d a h o e n s i s n.subsp. the more b a s i n a l environment  A becomes the dominant member i n  of the Degerbols Formation (as opposed  s u b o r d i n a t e member on the margin^) l e a d i n g to an age assignment and Nassichuk's N. c f . N. g r a c i l i s of Lower Roadian. n e c e s s a r y to c l a r i f y t h i s problem.  the  to a  f o r Kozur  More work i s indeed  However, the Upper Roadian age more  c l o s e l y f i t s the s t r a t i g r a p h i c framework f o r the a r e a as i t i s understood at p r e s e n t . E a r l i e r Age Assignments  f o r A r c t i c Permian  Formations  Sabine Bay Formation: ( T h o r s t e i n s s o n , 1974) In 1974 no f o s s i l s had been observed i n the Sabine Bay on E l l e s m e r e I s l a n d but i t s age was  g i v e n e a r l y A r t i n s k i a n because of i t s p o s i t i o n above  the B e l c h e r Channel Formation and below the A s s i s t a n c e .  A r t i n s k i a n ammonoids  i d e n t i f i e d as S v e r d r u p i t e s - w e r e r e p o r t e d from b a s a l beds o f the Sabine Bay Formation on M e l v i l l e  Island.  30  A s s i s t a n c e Formation  ( T h o r s t e i n s s o n , 1974)  On the B j o r n e P e n i n s u l a the A s s i s t a n c e Formation i s dated as  early  A r t i n s k i a n o r A k t a s t i n i a n i n age on the b a s i s o f ammonoids (Nassichuk et al.,  1965)  and b r a c h i o p o d s ( i d e n t i f i e d " by J.B. Waterhouse as b r a c h i o p o d  fauna "E" o f the N. Yukon and p r o b a b l y to the Jakutoproductus z o n e ) . However, t h i s age assignment  i s c o n s i d e r a b l y o l d e r than the type A s s i s -  tance ( G r i n n e l l P e n i n s u l a , . Devon I s l a n d ) and the A s s i s t a n c e i n the v i c i n i t y of  Hamilton Peninsula.  Harker and T h o r s t e i n s s o n (1960) suggested an  age  e q u i v a l e n t t o the B a i g e n d z h i n i a n s u b s e r i e s (upper A r t i n s k i a n ) on the b a s i s of  brachiopods.  Brachiopods from Hamilton P e n i n s u l a ( i d e n t i f i e d by  Waterhouse) i n d i c a t e an age o f U f i m i a n o r Kungurian. 1970;  Nassichuk et a l . ,  Artinskian  1965)  J.B.  Ammonoids ( N a s s i c h u k ,  i n d i c a t e both l a t e s t E a r l y Permian and  latest  ( B a i g e n d z h i n i a n ) age. T r o l d F i o r d Formation ( T h o r s t e i n s s o n , 1974)  Brachiopods ( i d e n t i f i e d by J.B. Waterhouse) from the T r o l d Formation i n d i c a t e a Kazanian age  (Wordian  substage).  Fiord  A s i n g l e ammonoid  (Nassichuk et al..., 1965), Neogeoceras macnari, i n d i c a t e s a  Guadalupian  age. Age Assignments  R e s u l t i n g From T h i s Work  The c o r r e l a t i o n s and age assignments r e s u l t i n g from t h i s study a r e based on conodonts  - as. opposed to the p r e v i o u s work i n the a r e a w i t h abun-  dant b r a c h i o p o d s and r a r e ammonoids. tically alter, Six of  The r e s u l t s r e f i n e , but do not d r a s -  t h e ages;.assigned by p r e v i o u s workers.  s u b d i v i s i o n s a r e proposed f o r the l a t e Lower and M i d d l e Permian  E l l e s m e r e I s l a n d r e p r e s e n t e d by the Sabine Bay, A s s i s t a n c e and  Fiord  Formations.  Trold  The t r a c e a b i l i t y of these s u b d i v i s i o n s f o r the e n t i r e  A r c t i c A r c h i p e l a g o i s i m p o s s i b l e to a s s e s s a t t h i s time because o f the r a t h e r i n f a n t stage o f Permian conodont  work i n the a r e a and because  of  the  e n v i r o n m e n t a l and r e s u l t i n g l i t h o l o g i c changes i n t o t h e b a s i n a l e q u i v a l e n t s of the d e s c r i b e d s e c t i o n s . r a t h e r than zones because  The s i x u n i t s a r e r e f e r r e d to as  subdivisions-,  they a r e i n p a r t based on l i t h o l o g y and e n v i r o n -  ment and i n p a r t on p a l e o n t o l o g y . The w r i t e r i s o p t i m i s t i c t h a t f u t u r e work w i l l  e v e n t u a l l y l e a d to  f u r t h e r r e f i n e m e n t and  s y n t h e s i s of >aj' good t r a c e a b l e b i o z o n a t i q n based  p r i m a r i l y on conodonts  but a l s o supplemented  ;  by b r a c h i o p o d s .  Subdivision A T h i s s u b d i v i s i o n i s a s s i g n e d t o the Sabine Bay as e s s e n t i a l l y a l i t h o s t r a t i g r a p h i c u n i t .  Formation and  regarded  Shallow marine tongues do occur  near the top of the f o r m a t i o n a t both the McKinley Bay and Hamilton P e n i n ^ sula sections.  Brachiopods and a few pelecypods were c o l l e c t e d a t both of  these l o c a l i t i e s - a p p a r e n t l y the f i r s t r e p o r t e d f o s s i l s i n E l l e s m e r e I s l a n d exposures of the f o r m a t i o n . r e p o r t e d from the Sabine Bay a r e dominated, Svalbardia).  Marine tongues w i t h ammonoid f o s s i l s were  Formation on M e l v i l l e I s l a n d .  The  brachiopods  almost e x c l u s i v e l y , by r u g o s o c h o n e t i d s (Neochonetes  or  A l t h o u g h the range i s much g r e a t e r , these brachiopods may  r e l a t e d t o Waterhouse's (Bamber and Waterhouse, 1971) of /Leonardian t o Roadian age from the N. Yukon.  be  b r a c h i o p o d ;fauna  Because of t h e i r  position  below the A s s i s t a n c e Formation a n ' e a r l y L e o n a r d i a n or B a i g e n d z i n i a n age i s assigned.  P r e v i o u s age assignments  t o the Sabine Bay Formation  from  M e l v i l l e I s l a n d i n d i c a t e d an A k t a s t i n i a n age a t the base o f the f o r m a t i o n . Rocks a s s i g n e d t o t h e b a s a l p a r t o f . t h e A s s i s t a n c e Formation on B j o r n e P e n i n s u l a , E l l e s m e r e I s l a n d i n d i c a t e d a c o r r e l a t i o n w i t h Waterhouse's  (ibid)  32  Jakutoproductus zone o f the.N. Yukon and a n ' A k t a s t i n i a n age. t i o n s from near the base of the Sawtooth  Fossil  Range s e c t i o n ( F i g . 3) c o n t a i n  p r o d u c t i d s u n l i k e any seen a t Hamilton P e n i n s u l a and t e n t a t i v e l y as J a k u t o p r o d u c t u s . of  collec-  identified  These c o l l e c t i o n s a r e f o l l o w e d by a few hundred  l a r g e l y u n f o s s i l i f e r o u s s e c t i o n b e f o r e abundant  metres  brachiopod f o s s i l s are  once a g a i n encountered - these faunas b e i n g v e r y s i m i l a r to t h o s e a s s i g n e d a L a t e L e o n a r d i a n to the  E a r l y Roadian age a t Hamilton P e n i n s u l a .  b a s a l p a r t s o f : t h e B j o r n e P e n i n s u l a and Sawtooth  The age f o r  Range s e c t i o n s suggest  g r e a t e r c o r r e l a t i o n . t o the Sabine Bay Formation than to the A s s i s t a n c e F o r mation. to  L i t h o l o g i c d i f f e r e n c e s have r e s u l t e d i n . t h e s e s t r a t a b e i n g a s s i g n e d  the A s s i s t a n c e Formation but they might be b e t t e r d e s c r i b e d as a  formation.  Furthermore, i t would appear that the Sabine Bay  on H a m i l t o n P e n i n s u l a and a t McKinley Bay i s younger P e n i n s u l a or a t the Sawtooth  new  Formation  than t h a t a t B j o r n e  Range.  Inosummary, a l t h o u g h the u n i t i s based l a r g e l y on l i t h o l o g y , the few f o s s i l c o l l e c t i o n s i n d i c a t e t h a t the u n i t ranges i n age from A k t a s t i n i a n to  E a r l y and p o s s i b l y M e d i a l L e o n a r d i a n o f B a i g e n d z i n i a n . Subdivision B T h i s s u b d i v i s i o n i s d e f i n e d as a l i t h o s t r a t i g r a p h i c u n i t at the base  and a b i o s t r a t i g r a p h i c range zone a t the top".. all  In o t h e r words, i t i n c l u d e s  t h a t s e c t i o n above the top o f the Sabine Bay Formation A s s i g n a b l e to the  A s s i s t a n c e f o r m a t i o n up to the top of the range f o r N. i d a h o e n s i s n.subsp. In  the H a m i l t o n P e n i n s u l a s e c t i o n t h i s i n c l u d e s the s t r a t a from the base  of  the A s s i s t a n c e Formation t o the top o f F54  ( F i g . 3).  At McKinley Bay  A.  this  s u b d i v i s i o n comprises a l l the s t r a t a a s s i g n e d to the A s s i s t a n c e Formation. N e o g o n d o l e l l a i d a h o e n s i s has been r e p o r t e d e x t e n s i v e l y i n s e c t i o n s from  33 the western U n i t e d States..  S i m i l a r assemblages  to the conodont  fauna of  s u b d i v i s i o n B i n the A s s i s t a n c e Formation have been r e p o r t e d from the Meade Peak P h o s p h a t i c S h a l e Member o f the P h o s p h o r i a Formation o f Idaho, Wyoming, and Utah (Youngquist et a l . ,  1951;  C l a r k and E t h i n g t o n , 1962;  C l a r k and Behnken, 1971; Wardlaw and C o l l i n s o n ( f i g . 3 ) , 19 79b)and a s s i g n e d ;  a Roadian age.  Other s i m i l a r faunas occur i n the Bone S p r i n g  V i c t o r i o Peak Formation, Guadalupe  Mountains, Texas  Limestone,  ( L e o n a r d i a n i n age  a c c o r d i n g t o Behnken (1975) and Upper L e o n a r d i a n t o Lower Roadian by Wardlaw and C o l l i n s o n (1978)).  A c c o r d i n g t o Wardlaw and C o l l i n s o n  (1979a)  N-. i d a h o e n s i s c e r t a i n l y ranges through the l a t e s t p a r t of the L e o n a r d i a n and d o u b t f u l l y i n t o the e a r l y p a r t of the Roadian.  N. s e r r a t a on the o t h e r  hand has been r e p o r t e d from the C u t o f f , Brushy Canyon and Getaway Member of the Cherry Canyon Formation o f West Texas and from the Meade Peak Phosp h a t i c s h a l e member of the P h o s p h o r i a Formation and a s s i g n e d a Roadian to E a r l y Wordian  age  E t h i n g t o n , 1962).  ( C l a r k and Behnken, 1979;  Behnken, 1975;  and C l a r k and  Neostreptognathodus p r a y i has been r e p o r t e d from the K a i -  bab of Nevada and Utah and the Bone S p r i n g Limestone and the V i c t o r i o Peak Formation of West Texas where i t i s a p p a r e n t l y r e s t r i c t e d  to t h e L a t e  Leonardian. I t appears t h a t s u b d i v i s i o n B can be c o r r e l a t e d w i t h the L a t e Leonardi a n to E a r l y Roadian of the western U n i t e d S t a t e s . ( F i g . 3) a t McKinley Bay.which  The assemblage  f o r F100  has N. i d a h o e n s i s subsp. i n d e t . i n associa ?  t i o n w i t h Neostreptognathodus p r a y t suggests a L a t e L e o n a r d i a n age. faunas q u a n t i t a t i v e l y s t u d i e d i n d e t a i l a t Hamilton P e n i n s u l a (F48 t o  1  The F54)  appear t o be m o r p h o l o g i c a l l y i n t e r m e d i a t e between N e o g o n d o l e l l a i d a h o e n s i s and N.  s e r r a t a s u g g e s t i n g t h a t an e a r l y Roadian age i s more a p p r o p r i a t e .  34 No s e r r a t i o n s were noted on samples from F100 and because of t h e a s s o c i a - . t i o n w i t h Neostreptognathodus  p r a y i these r e p r e s e n t a t i v e s of N e o g o n d o l e l l a  i d a h o e n s i s a r e c o n s i d e r e d o l d e r than those from F48 t o F54. s i n c e the d i s t r i b u t i o n s of Neostreptognathodus  However,  p r a y i and N e o g o n d o l e l l a  i d a h o e n s i s a r e f a c i e s c o n t r o l l e d to the extent t h a t they r a r e l y occur bedded i n a s i n g l e s e c t i o n , i t i s d i f f i c u l t Neostreptognathodus  p r a y i without  t o a s s e s s the s i g n i f i c a n c e of  i t s presence a t Hamilton P e n i n s u l a .  w e l l i t i s i m p o s s i b l e to a s s e s s whether Lower Roadian and  inter-  As  s t r a t a are present  condensed, were o r i g i n a l l y p r e s e n t and eroded, o r u n r e c o g n i z e d , o r were  never d e p o s i t e d a t McKinley Bay. T h i s s u b d i v i s i o n can be c o r r e l a t e d i n p a r t w i t h the Kapp S t a r o s t i n Formation of S p i t s b e r g e n ( S z a n i a w s k i and Malkowski,  1979) where the a u t h o r s  r e p o r t N. i d a h o e n s i s , N. c f . N. g r a c i l i s , and Neostreptognathodus b a r d e n s i s , . S w e e t o c r i s t a t u s a r c t i c u s and s e v e r a l ramiform  sval-  elements.  T h i s s u b d i v i s i o n i s a l s o c o r r e l a t e d w i t h Wardlaw and C o l l i n s o n ' s (1979a)well d e f i n e d b i o z o n a t i o n f o r the Great Basin-Rocky USA as i n c l u d i n g t h e i r Zone 1 ( P e n i c u l a u r i s i v e s i -  Mountain  Region  Neostreptognathodus  p r a y i zone) and the lower p a r t of Zone 2 ( P e n i c u l a u r i s b a s s i - N e o s t r e p t o gnathodus s u l c o p l i c a t u s  zone).  As a r e s u l t of t h e s e age assignments  the base o f the A s s i s t a n c e Forma-  t i o n a t Hamilton P e n i n s u l a can be regarded as no o l d e r than L a t e L e o n a r d i a n . Subdivision C T h i s s u b d i v i s i o n i s v e r y l o o s e l y d e f i n e d and i s r e p r e s e n t e d by a c o u p l e of s p a r s e conodont c o l l e c t i o n s a t F63 and F73 from the Hamilton P e n i n s u l a section.  The base of the u n i t , i s d e f i n e d by the top of s u b d i v i s i o n B w h i l e  the upper l i m i t  i s d e f i n e d as the h i g h e s t p a r t of the A s s i s t a n c e Formation  35 ( a l t h o u g h covered by  talus).  A l a r g e p a r t of s u b d i v i s i o n C i s covered  t a l u s r e s u l t i n g i n sparse c o l l e c t i o n s .  The  a l s o owing to the decreased abundance and b r a c h i o p o d s apparent i n the r o c k s .  by  low number of c o l l e c t i o n s i s  p o s s i b l e lower- d i v e r s i t y of  Conodonts show an  equal~. i f not more  d r a m a t i c , d e c r e a s e i n numbers when they a r e p r e s e n t at a l l . Those conodonts r e p o r t e d because of t h e i r s m a l l e r idahoensis  n.subsp. A,  are q u e s t i o n a b l y  s i z e at a p p a r e n t l y  s i m i l a r , growth stages to  i n the d i s c u s s i o n  f o r s u b d i v i s i o n B, N.  assignment to the o v e r l y i n g s u b d i v i s i o n s , M e d i a l to L a t e Roadian S u b d i v i s i o n .C can biozonation  as  s i n c e Wardlaw and  and! cusp.  a range age  t h i s s u b d i v i s i o n C can be  assigned  age.  be c o r r e l a t e d w i t h Wardlaw and  b a s s i - Neostreptognathodus sp.  at Hamilton P e n i n s u l a  s e r r a t a has  Because of the  i n c l u d i n g the upper p a r t of Zone 2,  top of the Roadian.  N.  d e n t i c l e s j u s t a n t e r i o r of the  d i s t r i b u t i o n of M e d i a l Roadian to E a r l y Wordian.  a  serrata(?)  sharper more compressed p o s t e r i o r d e n t i c l e s ,  the l a c k of f o u r d i s t i n c t i v e n o d e - l i k e As r e p o r t e d  a s s i g n e d to N.  C zone), and  Perhaps t h i s p a u c i t y i s r e a l and  not  Collinson's  Zone 3  (1979a)  (Peniculauris  t h e i r b a r r e n i n t e r v a l at  of conodonts seen f o r the  r e l a t e d to p r e s e r v a t i o n a l  The  conodonts may  Roadian  factors  C o l l i n s o n (1979a) a l s o r e p o r t a b a r r e n i n t e r v a l i n  Upper Roadian to lowermost Wordian.  the  the  have undergone a  c r i s i s l i k e t h a t ofpre-Wolfcampian conodonts ( C l a r k ,  1972)  p a r t e x p l a i n the  d i v e r s i t y i n the m i d d l e  and  subsequent i n c r e a s e d  abundance and  which would i n  upper p a r t s of the T r o l d F i o r d Formation (as i s o f t e n the case a f t e r a  near-extinction). The  top!of the A s s i s t a n c e  Upper Roadian and Peninsula  can  t h e r e f o r e be regarded as no younger than  the f u l l range f o r the A s s i s t a n c e  Formation at Hamilton  i s from the Upper L e o n a r d i a n to Upper Roadian.  Subdivision. D T h i s s u b d i v i s i o n i s a l s o l a c k i n g i n conodonts but dominated by t h e brachiopod C a n c r i n e l l o i d e s .  Above u n i t D the presence of N.  postserrata(?),  N. b i t t e r i n,. subsp.::C and N. r o s e n k r a n t z i  n.subsp. J) i n d i c a t e a L a t e  Wordian or Kazahian age, so t h a t D i t s e l f  i s regarded as E a r l y Wordian or  L a t e Kungurian i n age. Waterhouse,  i n Bamber and Waterhouse  zone i n the N. Yukon which he a s s i g n s however, t h a t  (1971), r e p o r t s a  a Kazanian age.  Cancrinelloides  He does i n d i c a t e ,  C a n c r i n e l l o i d e s can o c c u r i n s l i g h t l y lower beds  i n h i s Thamnosia zone; Thamnosia i s the dominant genus i n t h i s s u b d i v i s i o n E. Thamnosia zone.  Waterhouse a s s i g n s  included/ report's  a L a t e U f i m i a n or Kungurian age to t h e  For t h i s reason i t seems r e a s o n a b l e to a s s i g n an E a r l y  Wordian age t o u n i t  D.  Wardlaw and C o l l i n s o n (1979a)also d e f i n e a Thamnosia d e p r e s s a zone which they a s s i g n an E a r l y Wordian age. at H a m i l t o n P e n i n s u l a species  I f the o v e r l y i n g s u b d i v i s i o n E  i s dominated, by Thamnosia and v a r i o u s  then s u b d i v i s i o n D cannot be as young as K a z a n i a n .  s i n c e the base of u n i t D i s d e f i n e d  Neogondolella Furthermore,  by t h e base of the T r o l d F i o r d Forma-  t i o n then a t l e a s t p a r t of the T r o l d F i o r d i s o l d e r than Kazanian; u n l i k e p r e v i o u s r e p o r t s which r e s t r i c t e d The f o u r d i s t i n c t i v e f e a t u r e s  i t t o the K a z a n i a n . of t h i s u n i t a r e the apparent l a c k of  conodonts, t h e l a c k of. l a r g e t h i c k - s h e l l e d p r o d u c t i d s  l i k e Thamnosia, the  presence of C a n c r i n e l l o i d e s and o c c u r r e n c e i n the base of the green sandstones of t h e T r o l d F i o r d Formation.  The e n v i r o n m e n t a l s i g n i f i c a n c e of the  d i s t r i b u t i o n of t h e s e b r a c h i o p o d s i s not w e l l understood (Waterhouse, U n t i l more s e c t i o n s  i n the a r e a a r e s t u d i e d ,  1973).  encompassing a wide v a r i e t y  37 o f environments, both m a r g i n a l and cance of t h i s u n i t D cannot be  b a s i n a l , the b i o s t r a t i g r a p h i c s i g n i f i -  established. Subdivision E  U n i t E i s the c l o s e s t to a r t r u e on the p r e s e n c e and rence of r e a s o n a b l y  range or acme biozone as i t i s based  dominance of the brachiopod abundant conodonts.  Thamnosia and  the  A l s o c h a r a c t e r i s t i c of  occurthis  s u b d i v i s i o n i s a conglomeratic  u n i t t h a t , except f o r v a r y i n g t h i c k n e s s , i s  i d e n t i c a l i n the McKinley Bay,  Hamilton P e n i n s u l a  tions.  I f t h i s conglomerate can be shown to be  and  Sawtooth Range s e c -  synchronous then i t c o u l d  prove a v e r y u s e f u l marker h o r i z o n ( F i g . 3) - p o s s i b l y r e l a t e d to a s i n g l e t e c t o n i c p u l s e or o t h e r  s h o r t - d u r a t i o n p h y s i c a l phenomenon.  At Hamilton P e n i n s u l a to F90  and  F35  Neogondolella ble  to F44. n. sp. J3.  conglomeratic  unit.  subdivision E includes f o s s i l  F36  and  c o n t a i n specimens of a new  Both of these c o l l e c t i o n s occur F87  f e r a b l e to N. p o s t s e r r a t a ( ? ) . tic unit.  F83  N. p o s t s e r r a t a has  c o l l e c t i o n s F83 species;  below the  chert-peb-  i n c l u d e s conodont fragments q u e s t i o n a b l y  T h i s . - c o l l e c t i o n o c c u r s w i t h i n the conglomerap r e v i o u s l y been r e p o r t e d  from the  Southwells  Member of the Cherry Canyon Formation to the Lower McCombs Member of B e l l Canyon Formation of Idaho and t a n i a n age  (Behnken, 1975;  (1979a)describe  Great B a s i n and  Texas and  C l a r k and  Behnken, 1979) .  Rex  Chert  a Wordian and Wardlaw and  Rocky Mountain r e g i o n of the western USA  or l e s s e q u i v a l e n t  the Capi-  Collinson the  Member of the P h o s p h o r i a Formation i n the  a s s i g n an E a r l y to M e d i a l Wordian.age.  Behnken, 1971;  assigned  a Thamnosia d e p r e s s a zone from the upper p a r t of  Plympton Formation, and  re-  to which they  They i n d i c a t e t h a t t h i s zone i s more  to the Neospathodus a r c u c r i s t a t u s assemblage ( C l a r k  C l a r k et a l . , 197.9) which C l a r k et a l . a s s i g n a Wordian  and age.  38  C l a r k e t a l . (1979) i n d i c a t e t h e e q u i v a l e n c e o f t h i s zone t o t h e Neogond o l e l l a d e n t i c u l a t a fauna o f West Texas d e s p i t e g i v i n g i t a C a p i t a n i a n t o Amarassian  age i n C l a r k and Behnken (1979).  Wardlaw and C o l l i n s o n  (1979a)  and C l a r k e t a l . (1979) both d e s c r i b e o v e r l y i n g faunas o f N. b i t t e r i and _N. r o s e n k r a n t z i a s s i g n e d to a L a t e Wordian and C a p i t a n i a n age.  F o r these  reasons N. p o s t s e r r a t a  (which i s o l d e r than N. d e n t i c u l a t a ) cannot be any  younger than Wordian.  Assuming ages a s s i g n e d to t h e type N. p o s t s e r r a t a  and f o r t h e Thamnosia d e p r e s s a zone t h e c o l l e c t i o n s below and w i t h i n t h e ;-  *  conglomerate  o f s u b d i v i s i o n E can be regarded as M e d i a l Wordian o r Upper-  most Kungurian  (Ufimian) i n age.  S u p p o r t i n g t h i s assignment  i s another  Thamnosia zone d e s c r i b e d by Waterhouse (Bamber and Waterhouse, 1971) from the N. Yukon.  Here Waterhouse a s s i g n s a L a t e U f i m i a n age s t a t i n g t h a t t h e  Thamnosia o f h i s zone-Ft a r e more e v o l v e d than those found i n t h e A s s i s t a n c e Formation.  Thamnosia i s abundant throughout  t h e c o n g l o m e r a t i c u n i t and  s l i g h t l y above i t and thus d e f i n e s t h e top o f u n i t E, L a t e Kungurian or M e d i a l Wordian i n age. Subdivision F S u b d i v i s i o n F o c c u r s above t h e zone w i t h dominant Thamnosia and i n c l u d e s t h e f o s s i l c o l l e c t i o n s F91 t o F97 and F45 t o F47 .  The conodonts  i d e n t i f i e d from t h i s zone i n c l u d e N e o g o n d o l e l l a b i t t e r i n.subsp. C and N. r o s e n k r a n t z i n.subsp. I). B r a c h i o p o d s . i n c l u d e v a r i o u s s p i r i f e r i d s and p r o d u c t i d s i n v a r y i n g degrees o f abundance; e s p e c i a l l y common a r e s p e c i e s of Y a k o v l e v i a and K u v e l o u s i a .  However, both o f these genera occur i n many  u n d e r l y i n g zones and r e q u i r e s p e c i f i c  i d e n t i f i c a t i o n b e f o r e they c o u l d be  <used :for range-zone..determination. The fauna j u s t d e s c r i b e d compares v e r y w e l l w i t h t h a t p r e s e n t i n Wardlaw and C o l l i n s o n ' s (1979a)zones  5 ,and6 (KuveTousi'a^'•'•leptosa zone and  39 Yakovlevla m u l t i s t r i a t a . - Neogondolella -Rocky Mountain r e g i o n o f western USA  b i t t e r i zone) from the Great  and  a s s i g n e d a l a t e Wordian  Basin  age.  The conodonts a t hand a r e v e r y s i m i l a r to faunas f i g u r e d i n Wardlaw and Collinson  (1979b) from the R e t o r t Phosphatic  Shale Member of t h e Phosphoria  Formation from Montana ( i n the case of N. r o s e n k r a n t z i ) and Wyoming ( f o r _N. b i t t e r i ) .  Representatives  f o r both.species  from the G e r s t e r  Formation  appear more advanced. N_. r o s e n k r a n t z i has a l s o been d e s c r i b e d from East Greenland i t was  named by Bender and  c l e a r a t the time. a C a p i t a n i a n and and  Stoppel,l965)though  t h e age r e l a t i o n s h i p s were not  C l a r k and Behnken (1979) and  Amarassian age  (where  C l a r k et a l . (1979) a s s i g n  to faunas d e s c r i b e d from the Radar, McCombs,  Lamar Members of the B e l l Canyon.Formation of Texas and  Formation of Nevada and Wyoming.  The main reason  the  Gerster  f o r c o n t r a d i c t i n g age  assignments a r e the d i f f e r e n c e s of o p i n i o n f o r the r e c o g n i t i o n of N. and  N. r o s e n k r a n t z i by Wardlaw and  (1979) i d e n t i f i e d Collinson,  C o l l i n s o n and  C l a r k et a l .  bitteri  C l a r k et a l .  specimens of N. r o s e n k r a n t z i ( a c c o r d i n g to Wardlaw and  1979b)as N. b i t t e r i .  T h e r e f o r e , N. r o s e n k r a n t z i can occur down  i n t o the L a t e Wordian,(see C l a r k et a l . , 1979  f o r d i s c u s s i o n and  descrip-  tion) . Having r e s o l v e d these c o n t r a s t i n g age  assignments and  stated that  conodonts at hand a r e l e s s advanced than C a p i t a n i a n specimens from Gerster limestone, One  other  I a s s i g n s u b d i v i s i o n F to  b r a c h i o p o d s near the top.  the  L a t e Wordian or Kazanian  f e a t u r e t h a t i s c h a r a c t e r i s t i c of t h i s f i n a l  t h e Sawtooth Range and H a m i l t o n P e n i n s u l a  mentally  a  the  age.  subdivision at  i s t h e dominance of  A l t h o u g h such a f e a t u r e i s p r o b a b l y  inarticulate environ-  c o n t r o l l e d , t h e h o r i z o n c o u l d prove to be a u s e f u l marker ( F i g . 3 ) .  40  The absence of i n a r t i c u l a t e s a t McKinley  Bay  suggests  t h a t a l l o r p a r t of  t h i s s u b d i v i s i o n , i s m i s s i n g h e r e : perhaps as a r e s u l t of r e g r e s s i o n i n the l a s t part of the T r o l d F i o r d  Formation.  The top "of u n i t F i s c o i n c i d e n t w i t h the top of the T r o l d F i o r d mation, the l a s t Permian s t r a t a i n the! A r c t i c . Formations  The B l i n d F i o r d or  of Lower T r i a s s i c age r e s t unconformably on the T r o l d  I n d i c a t i o n s a r e t h a t the T r o l d F i o r d Formation  ranges i n age  t h e r e i s no d i r e c t evidence  o r never d e p o s i t e d a t a l l .  10 m i l l i o n  have been.present  I t i s f a i r to say t h a t the time  by the T r o l d F i o r d / B j o r n e u n c o n f o r m i t y  represented  cannot be e n t i r e l y  out s i n c e N. b i t t e r i and N . / r o s e n k r a n t z i can range t h i s h i g h . i m p o s s i b l e to say whether younger r o c k s may  Fiord.  Kazanian.  f o r i t , a C a p i t a n i a n age  i n the upper p a r t s of the T r o l d F i o r d Formation  Bjorne  from the Lower  Wordian or Upper Kungurian to the Uppermost Wordian or Upper Although  i s considerable: l  n  For-  ruled  It i s also and  eroded,  represented the o r d e r of  years. Summary  The  Sabine Bay,  A s s i s t a n c e and  T r o l d F i o r d Formations  have been  a t e d i n t o s i x s u b d i v i s i o n s which can be c o l l e c t i v e l y r e f e r r e d to as t i n i a n to L a t e s t Wordian o r Kazanian a r e separated by u n c o n f o r m i t i e s them, "  i f present at a l l ,  to be  age.  Although  the t h r e e  ( T h o r s t e i n s s o n , 1974) Q  f short"duration.  s e c t i o n s c o u l d i n d i c a t e continuous  this  separAktas-  formations  author.considers  Perhaps more b a s i n a l  s e d i m e n t a t i o n w i t h the t r a n s g r e s s i o n s and  r e g r e s s i o n s o n l y a f f e c t i n g the margins of the b a s i n . QUANTITATIVE ANALYSIS OF MEASURABLE CHARACTERS. FOR  NEOGONDOLELLA  Introduction Q u a l i t a t i v e o b s e r v a t i o n of the conodont p o p u l a t i o n s i n samples F49,  F52,  F53 and'F54 i n d i c a t e d  F48,  t h a t t h e r e a r e no d i s t i n g u i s h a b l e d i f f e r -  ences w i t h r e s p e c t to o v e r a l l p l a t f o r m shape and d e n t i c l e c o n f i g u r a t i o n . A d e s c r i p t i o n based on these o b s e r v a t i o n s was s u b s p e c i e s of N e o g o n d o l e l l a  new  i d a h o e n s i s but d i d n o t h i n g to e l u c i d a t e any  e v o l u t i o n i n the p o p u l a t i o n s . j u s t my  s u f f i c i e n t to e r e c t a  I t a l s o seemed expedient  s u b j e c t i v e o p i n i o n on which to base the new  to have more than  subspecies.  As a.  r e s u l t , a q u a n t i t a t i v e a n a l y s i s was.^undertaken to determine i f t h e r e any demonstrable e v o l u t i o n i n the p o p u l a t i o n s and criteria  to p r o v i d e  unbiased  on which to base the taxonomyr(see Appendix I f o r data)  The measurable parameters a n a l y z e d  was  v  i n c l u d e the o v e r a l l p l a t f o r m  l e n g t h ( L l ) , the l e n g t h from the p o s t e r i o r cusp to the f o u r t h d e n t i c l e a n t e r i o r of t h e cusp ( L 2 ) , the h e i g h t from the t i p of the cusp to the base of the f l a n g e ( H i ) , the maximum w i d t h end of the p l a t f o r m (W2) , and  (Wl), the w i d t h a t the p o s t e r i o r  the number of d e n t i c l e s  (//) on the p l a t f o r m .  Some of these parameters have been measured on s i m i l a r conodont p o p u l a t i o n s by o t h e r workers ( L l and  # by Behnken, 1975;  L l , HI and  // by D z l k  Trammer, 1980).  In a d d i t i o n a number of r a t i o s were determined  L l / H l , Ll/Wl and  Ll/#.  The  f i n a l parameter a n a l y z e d  i n the p o s t e r i o r end of the p l a t f o r m determined  and  including  i s a f u n c t i o n of a r e a  by the e q u a t i o n L2(Wl +  W2)%  These v a r i o u s parameters were a n a l y z e d by c a l c u l a t i n g the mean, s t a n dard e r r o r f o r the mean, the standard d e v i a t i o n of the mean and, a comparison u s i n g z- and of  t - t e s t s to determine the s t a t i s t i c a l  any d i f f e r e n c e s between p o p u l a t i o n s .  finally,  significance  B e f o r e l i s t i n g and d i s c u s s i n g the  r e s u l t s of t h e s e c a l c u l a t i o n s the background, assumptions and i m p l i c a t i o n s of  such s t a t i s t i c a l  t e s t i n g should be o u t l i n e d .  Above a l l one must remember t h a t i n each, case we  are d e a l i n g with a  sample of the p o p u l a t i o n and not the p o p u l a t i o n i t s e l f .  Any  two  samples  42-  from t h e same p o p u l a t i o n w i l l vary; s i z e t h i s d i f f e r e n c e isminimal.  however, i n samples o f s u f f i c i e n t  Since the population  i s the u n i t of e v o l -  ution,-, ::the: samples s t u d i e d h e r e i n can o n l y be regarded as approximating the e v o l u t i o n a r y t r e n d s . of t h e p o p u l a t i o n  A requirement f o r a sample to be r e p r e s e n t a t i v e  i s t h a t i t be  s e l e c t e d a t random.  f o r t h i s study were a l l . those ( l a r g e and small) t h a t were s u f f i c i e n t l y p r e s e r v e d characters.  The o n l y p r o c e s s e s  The conodonts s e l e c t e d  i n each f o s s i l  collection  t o a l l o w f o r t h e measurement o f t h e v a r i o u s i n t h e s e l e c t i o n of t h e specimens were  those as a r e s u l t o f t h e environment o f d e p o s i t i o n , the d i a g e n e t i c h i s t o r y and  t h e sample p r o c e s s i n g .  Although these p r o c e s s e s  can be non-random  ( p r e f e r e n t i a l breakage o r e t c h i n g o f more f r a g i l e specimens), t h e r e s u l t i n g frequency curves ( F i g . 5) r e a s o n a b l y suggesting  that the processes  approximate a normal d i s t r i b u t i o n ,  were too s m a l l to be s e l e c t i v e .  p r e s e n t a t i v e approximation of the population, for s t a t i s t i c a l analysis.  Having a r e -  the sample i s now a v a i l a b l e  C a l c u l a t i o n o f t h e mean o f any c h a r a c t e r  involves  EX the simple  s o l v i n g o f t h e equation'M'= —  (M = mean, ZX = sum o f the v a l u e s  f o r a c h a r a c t e r , and N = the number of specimens w i t h i n t h e sample). However, Burma (1948). p o i n t s o u t t h a t t h e mean o f a sample c o n s i s t i n g o f a growth s e r i e s i s merely t h e mean s i z e o f h a l f grown specimens, t h e minimum s i z e i s t h a t o f t h e s m a l l e s t o f t h e youngest specimens and t h e maximum  s i z e i s t h e l a r g e s t o f t h e o l d e s t specimens.  Burma (1948) f u r t h e r m o r e  s t a t e s t h a t such a procedure i s meaningless and l a c k i n g i n b i o l o g i c a l n i f icance: .and i t should  t h a t i f one dictum i s e s t a b l i s h e d i n q u a n t i t a t i v e  be t h a t comparisons o f one c h a r a c t e r ,  a t comparable growth stages  only.  sig-  paleontology  t o be v a l i d , must be made  H i s p o i n t i s w e l l , taken but one . d i f f i c u l t  to heed i n many groups, e s p e c i a l l y t h e conodonts.  Such a procedure would  Element F48  F49  Representation F52  F53  r5 E :  8  13  15  8  13  Element F48  I in  I  F49  o  F i g u r e 5.  LO  7  8  16  6  O io  S £  7  g o  o in io  *  F53 + F54  overall  iii 15  F 4 9+F52  .lk.  Q o u> io  Denticles  for Platform  F54  JUL  of  F49+F52  15  Representation F53  id O •0  F54  Ju A  F52  I1 • I  for Number  in  v  6  Length  16  (pm)  F53+F54  o o to eo co  io  16  overall  o o 05 io a co  o m *  44  be easy w i t h a group l i k e the ammonites where f e a t u r e s a r e p r e s e n t t h a t a l l o w one t o d i s t i n g u i s h an a d u l t conch. f o r the conodont  However, no such f e a t u r e s e x i s t  p l a t f o r m ( D z i k and Trammer,  f o r e even b e g i n n i n g the a n a l y s i s ? .  1980).  Are we d e f e a t e d be-  Other workers have shown t h a t  phores l i k e N e o g o n d o l e l l a Have a complex ontogeny.  conodonto-  M e r r i l l and P o w e l l '  (1980) demonstrated an ontogeny of P e n n s y l v a n i a n G o n d o l e l l a where the app a r a t u s began as ramiform elements  o n l y and subsequently developed  p l a t f o r m ( " j u v e n i l e " ) and ramiform elements and f i n a l l y o n l y ("mature") a p p a r a t u s .  into  into a platform  Other workers have a l s o suggested t h a t more than  one p a i r of p l a t f o r m s , each of which a r e a t : : d i f f e r e n t developmental comprise  the a p p a r a t u s .  p l a t f o r m may  In o t h e r words, the developmental  stages,  stage of the  bear no r e l a t i o n s h i p to the a c t u a l age o f the conodont  Unless the samples d i s p l a y some unusual m o r t a l i t y r a t e , i t may  animal.  be v a l i d  to compare the e n t i r e range of p l a t f o r m s i z e s as the m a j o r i t y of them p r o b a b l y reached a c e r t a i n s t a g e i n development b e f o r e death.  With  c o n s i d e r a t i o n s i n mind I proceeded w i t h the s t a t i s t i c a l comparisons e n t i r e sample but a l s o separated, the d a t a i n t o two w i t h 10 d e n t i c l e s and those w i t h 11)  s u b s e t s (those  these of the  elements  of p o s s i b l e p a r t i c u l a r growth s t a g e s .  A f t e r c a l c u l a t i n g the mean of each sample the standard e r r o r of the mean was  c a l c u l a t e d a t the 95%  confidence l e v e l  (a  = —Swhere a-== standard m . N d e v i a t i o n or measure of c e n t r a l tendency o f v a r i a b i l i t y ) . F i n a l l y , the l i m -  + i t s of v a r i a b i l i t y were s e t a t the 75% 2.0a  respectively).  sumes a near normal graphed  and 95%  l e v e l s (M - 1.15a  + arid M -  Proper use and f u l l v a l u e of. t h e s e c a l c u l a t i o n s a s d i s t r i b u t i o n of the c h a r a c t e r s .  These v a l u e s were then  f o r v i s u a l impact and compared i n the cases o f l e n g t h and  the  r a t i o length/number of d e n t i c l e s u s i n g z- and t - t e s t s t o determine the s i g -  45  n i f i c a n c e of the d i f f e r e n c e s . equal standard 1975)  and  The  t - t e s t assumes a normal d i s t r i b u t i o n  d e v i a t i o n s f o r the two  i s valuable  samples being  f o r samples of low number.  pend on the same assumptions but  and  compared (Hodges et a l . ,  The  z - t e s t does not  i s o n l y u s e f u l f o r samples of  de-  approxi-  mately 20 or more specimens. It  i s g e n e r a l l y thought t h a t the q u a n t i t a t i v e approach i n s c i e n c e i s  the o n l y t r u l y o b j e c t i v e approach (Raup and t h i s a s s e r t i o n the t e c h n i q u e i s o f t e n met gists.  ple,  Secondly, a n y t h i n g  w i t h o b j e c t i o n by  Despite  paleontolo-  However, s t a t i s t i c a l methods are a v a i l a b l e  apply  specimens or w i t h 1000  behind them ( i b i d . ) .  be minimized.  The  the f u l l range of v a r i a t i o n of any  So  long  as  information  v a l u e of such an  to d e f i n e minute changes i n the specimens through time, and  weighs t h e s e o b j e c t i o n s .  those  i s o f t e n e i t h e r owing to l a c k of knowledge  the l i m i t s of i n t e r p r e t a t i o n f o r the d e r i v e d  t h i s l a s t o b j e c t i o n should  s p e c i e s being  founded  Another o b j e c t i o n i s t h a t misuse of s t a t i s t i c s by  them to p a l e o n t o l o g y  recognizes  or more (Burma,  s t a t i s t i c a l a n a l y s i s w i l l p r o b a b l y be  or l a c k of a p p r e c i a t i o n , of the p h i l o s o p h y one  p. 42).  which a person attempts to do w i t h a s m a l l sam-  which he c o u l d not do by  on e r r o r ( i b i d . ) . who  f o r a few.  study o f samples w i t h a v e r y few  1948) .  1971;  Many workers s t a t e t h a t the method i s good f o r a l a r g e number of  specimens but not for  Stanley,  analysis  to:.indicate  species or i n f r a s p e c i f i c u n i t f a r out-  A l l s p e c i e s t h a t a r e c r e a t e d w i t h the concept o f  an e n t i t y of l i t t l e v a r i a t i o n should  be viewed w i t h  suspicion  (ibid.). Results The All  and  Discussion  measurements f o r the specimens s t u d i e d are  included  i n Appendix  measurements were completed w i t h a micrometer set i n a b i n o c u l a r  A.  micro-  scope a t 75 power..  This, method allowed  f o r t h e l e n g t h .of a 1000 um p l a t f o r m ;  an approximate a c c u r a c y  i n o t h e r words an a c c u r a c y  i m a t e l y 1.0% and a l l o w i n g f o r a t l e a s t t h r e e s i g n i f i c a n t  of + 5 um of approx-  figures.  Except  f o r F54, t h e samples were measured i n terms of a l l t h e parameters s t u d i e d . Only l e n g t h and t h e number o f d e n t i c l e s were determined f o r F54 as these specimens were p l a c e d on SEM stubs and photographed b e f o r e  i t seemed appro-  p r i a t e t o proceed w i t h t h e more d e t a i l e d a n a l y s i s . O v e r a l l length of platform ( L l ) Perhaps t h e most obvious r e f l e c t i o n o f i n c r e a s i n g m a t u r i t y o f t h e p l a t f o r m i s an i n c r e a s e i n l e n g t h .  More s i g n i f i c a n t  i s the progressive  i n c r e a s e u p s e c t i o n f o r t h e mean v a l u e o f l e n g t h from 816 um t o 905 um ( T a b l e 1 ) . A l t h o u g h t h e v a r i a t i o n i n t h e p o p u l a t i o n samples o v e r l a p s f o r the most p a r t , t h e i n c r e a s e i n t h e mean v a l u e i s e n t i r e l y c o n s i s t e n t upsection  ( F i g . 6) f o r N. i d a h o e n s i s n.subsp. A.  n.subsp. D_ r e p r e s e n t e d  However, N. r o s e n k r a n t z i  i n F96 has a :mean l e n g t h c o n s i d e r a b l y l e s s than t h a t  f o r F48 t o F54 i n d i c a t i n g a r e v e r s a l i n t h e t r e n d .  The l a r g e v a r i a b i l i t y  i n l e n g t h i n F96 r e s u l t s i n a s i g n i f i c a n c e l e v e l , a t b e s t , o f 16% ( T a b l e 2) when s u b j e c t e d represented  to the z - t e s t .  T h i s i s o f even l e s s s i g n i f i c a n c e than t h a t  f o r t h e change o f l e n g t h from F49 to F54 (11.5%).  Despite  this  l a c k of s i g n i f i c a n c e f o r the d i f f e r e n c e , the trend i n d i c a t e d i s consistent w i t h t h e e v o l u t i o n o f N. i d a h o e n s i s  to N. r o s e n k r a n t z i a c c o r d i n g t o the  r e s u l t s o f other workers.(Behnken, 1975; C l a r k and Behnken, 1979). A l t h o u g h t h e t r e n d from F48 t o F54 may p o s s i b l y be e x p l a i n e d by evolutionary processes, stages p r e s e r v e d . of P e n n s y l v a n i a n  i t c o u l d a l s o be t h e r e s u l t o f the o n t o g e n e t i c  M e r r i l l and Powell  (1980) i n d i c a t e d t h a t t h e ontogeny  G o n d o l e l l a proceeded from ramiform o n l y t o p l a t f o r m  only  "ST" • P a r a m e t e r N.  M ± 2a m a 75% range* 95% r a n g e * * M + 2a in a 75% r a n g e * 95% range** M ± 2o m a 75% range* 95% r a n g e * * M ± 2o m a 75% range* 95% range** M ± 2a m a 75% range* 95% range** M ± 2o  ^lpm  L  2)im  lvim  W  Data  Sample —  2um  1pm  V 1  L  l  /  H  0  0  L (W +W )5i 2  1  IO  2  4  nm 2  Table  1.  Statistics of  F53 N =(38-40)  F54 N = (37)  816 ± 134 212 572 - 1060 402 - 1230  847 ± 48 184 635 - 1 0 5 9 479 - 1 2 1 5  847 ± 80 199 618 - 1 0 7 6 449 - 1 2 4 5  888 ± 82 259 590 - 1 1 8 6 370 - 1406  905 ± 84 255 612 - 1198 395 - 1415  297  + 26 40 251 - 343 217 - 377  279  ± 10 42 231 - 327 195 - 363  286 ± 16 38 242 - 330 210 - 3 62  303  181  ± 28 44 1 3 0 - 232 93 - 269  183  ± 1.6 61 113 - 253 61 - 305  200 ± 18 44 149 - 251 112 - 288  209  152  ± 24 38 108 - 196 76 - 228  163  ± 8 31 127 - 199 101 - 225  174 ± 18 47 120 - 228. 8 0 - 268  180  149  164 ± 10 35 124 - 204 94 - 234  166 ± 18 42 118 - 214 82 - 250  172  10.78 + 1.02 1.61 8.15 - 11.85 8.92 7.54 6.78 - 13.22  75% r a n g e * 95% r a n g e * * M i 2a m a 75% range* 95% r a n g e * * M ± 2 m a 75% r a n g e * 95% r a n o e * *  l  F52 N - (24-25)  10.00  a 75% range* 95% r a n g e * * M i 20 m a 75% range* 95% r a n g e * * M ± 2o m  W  F49 N = (58-60)  +20 29 116 - 182 91 - 207  ra  9  F48 N - (9-10)  + 18 55 146 - 272 99 - 319 ± 16 53 119 - 241 74 - 286 + 14 44 121 - 223 84 - 260  11.02 ± .60 1.50 - 12.53 8.58 6.78 - 13.80  10.92 ± .66 2.12 8.85 - 13.46 7.32 - 15.26  80.8  + 7.0 11.1 68.0 - 93.6 58.6 - 103.0  78.3  ± 3.0 11.7 64.8 - 91.8 54.9 - 101.7  77.6  + 3.8 9.5 66.7 - 8 8 . 5 58.6 - 96.6  79.3  4.51 + .22 0.34 4.12 - 4.90 3.83 - 5,19  4.44 ± .14 0.54 3.82 - 5.06 3.36 - 5.52  4.27 + .22 0.55 3.64 - 4.90 3.17 - 5.37  4.23 ± .12 0.40 3.77 - 4.69 3.43 - 5.03  5.30 + .70 1.05 4.09 - 6.51 3.20 - 7.40  5.21 + .22 0.81 4.28 - 6.14 3.59 - 6.83  5.18 ± .30 0.73 4.34 - 6.02 3.72 - 6.64  5.14 + .34 1.05 3.93 - 6.35 3.04 - 7.24  4.99 ± .88 1.40 3.38 - 6.60 2.19 - 7.79  4.79 + .43 1.67 3.05 - 6.89 1.63 - 8.31  5.48 ± .88 2.18 2.97 - 7.99 1.12 - 9.84  6.11 + .86 2.71 2.99 - 9.23 0.69 - 11.53  from d a t a  statistical  10.80 ± .42 1.62 9.07 - 12.64 - 1 4 . 0 2 7.80  ± 16 53 24 2 - 364 197 - 4 09  values  given used.  i n Appendix  I.  + 3.6 11.1 66.5 - 92.1 57.1 - 1 0 1 . 5  See page 44  82.0  ± .60 1.80 - 12.99 - 14.52  + 5.0 15.3 64.4 - 9 9 . 6 51.4 - 1 1 2 . 6  for discussion  -1  1  7 EC  1  1  Jj^  1  I  1  1  9  10  11  1  i 13  1—  12  i  3D  L,/  +-  x10"  2  1——I  • B  SE  3 650  1  1  i j-.'iM.'i  1——f  1-  850  h  1  1050  8  1—-I  1  1  1250  9  1  1  IA  10  -I——I  1450  11 1  1  12 1  1  1650  13  H  ••'•.••••V.>:  l'r-:-'.'  3D  A«BMlSlim^ .-!>W.!'.-. l  •B  •i  H  650  6  •+-  1  —I  8  850 1  1  1  h  10  1050 r-  11  H  h  12  1250 1  r-  H  13  1450  14  H  =3  1  15  1  A  1  16  1650 h  H 17  1F9&  HE  F54 F53 F52  denticles  » i  6 II  F i g u r e 6.  •B 8  9  10  11  12 .9  I  13  •  •  14  <  F49  15  F48  16  G r a p h i c a l r e p r e s e n t a t i o n of p a r t of" T a b l e 1. L i n e s i n f u l l bla_ck show range of M±-2'a^. S t i p p l e d p a t t e r n = M - 1.15a. (.i.e. 75% v a r i a t i o n ) . White = M - 2a ( i . e . 95% v a r i a t i o n )  OX  z-Test f o r L  i  T~ P - v a l u e t t  Samples Compared F49  z-score  z-Test f o r #  'z-Test f o r ' L /#'  E n t i r e Sample  • t In  z-score  1  +  -t  +  z-score  P-value  P-value^  F52  0.01  0.49.6  0.055  0.478  0.29  0.386  F49 -»• F53  0.87  0.192  0.61  0.271  0.43  0.334  F49 + F54  1.20  Q.115  0.39  0.348  1.26  0.104  (F49+F52) -> (F53+F54)  1.37  0.085  0.64  0.261  1.30  0.097  F49  F96  0.51  0.305  1.89  0.029  3.95  0.000  F54 -> F96  0.97  0.166  1.70  0.045  4.35  0.000  T a b l e 2.  Values d e r i v e d  t = -Mi - M2  from z - t e s t s f o r L l , // and'Ll/7/. the z - s c o r e o f t h e d i f f e r e n c e between two independent M = mean  Nl ft=  N  1  a = standard d e v i a t i o n  samples (1, 2)  N = number o f specimens  2  t h e a r e a under a normal curve ( t o t a l a r e a = 1) t o t h e l e f t o r r i g h t o f M ± za which i n d i c a t e s t h e p r o b a b i l i t y t h a t t h e d i f f e r e n c e between samples 1 and 2 c o u l d occur by chance a l o n e (a two s i d e d t e s t i s e q u a l t o t h e t o t a l a r e a t o the l e f t and r i g h t arid i n d i c a t e s t h e combined chance o f g e t t i n g a d e v i a t i o n i n e i t h e r direction  50  apparatuses-.  I f t h i s i s t r u e f o r the Neogondolella.. apparatus  as w e l l ,  the i n c r e a s i n g degree of p l a t f o r m o v e r r e p r e s e n t a t i o n or, r a t h e r ,  then  ramiform  u n d e r r e p r e s e n t a t i o n , c o u l d be r e l a t e d t o i n c r e a s i n g m a t u r i t y and l a r g e r ;,:' s i z e of the p r e s e r v e d sample.  Table 3 l i s t s  the d a t a : f o r element r e p r e -  s e n t a t i o n and demonstrates a f l u c t u a t i o n i n the p l a t f o r m to ramiform Except  f o r the change from F49  the ramiform  to F52  the d a t a show a c l e a r i n c r e a s e i n  u n d e r r e p r e s e n t a t i o n comparing w e l l , w i t h the i n c r e a s e i n  l e n g t h f o r the same i n t e r v a l .  The mean v a l u e f o r l e n g t h i n F52  i s equal t o  t h a t f o r F49 which, a l t h o u g h i t should be a decrease, approximates t r e n d i n element r e p r e s e n t a t i o n . may it  ratio.  be important  the  A l t h o u g h i t seems c l e a r t h a t t h i s  concept  f o r .explaining some of the change i n l e n g t h n e v e r t h e l e s s  i s not c o m p l e t e l y  satisfactory.  Heeding Burma's (1948) p l e a t h a t o n l y s i m i l a r growth stages should s t a t i s t i c a l l y compared the d a t a f o r each sample were d i v i d e d s e t s ( T a b l e 4 ) : the f i r s t  be c o n s i d e r e d a p e r f e c t  ing  Furthermore,  r e l a t i v e age.  the  The number of d e n t i c l e s  i n d i c a t o r of r e l a t i v e age,  however, i t i s  t r u e t h a t t h e i r number i n c r e a s e s d u r i n g the i n f e r r e d ontogeny of the ment.  sub-  w i t h a l l those elements w i t h 10 d e n t i c l e s and  second w i t h a l l the. elements w i t h 11 d e n t i c l e s . cannot  i n t o two  be  ele-  i t i s probably the best t o o l a v a i l a b l e f o r d i s t i n g u i s h The v a l u e s d e r i v e d from t - t e s t s of the r e s u l t i n g  s e t s p r o v i d e some i n t e r e s t i n g r e s u l t s .  sub-  The d i f f e r e n c e between the samples  w i t h 10 d e n t i c l e s i s i n s i g n i f i c a n t whereas the d i f f e r e n c e between elements w i t h 11 d e n t i c l e s i n F49 and graph  F54  i s s i g n i f i c a n t . , even a t the 1% l e v e l .  i n F i g u r e 7 i l l u s t r a t e s t h i s o b s e r v a t i o n w e l l i n t h a t as  number i n c r e a s e s the d a t a p o i n t s f o r the two further apart.  E a r l y s t a g e s cannot  The  denticle  samples p l o t p r o g r e s s i v e l y  be d i s t i n g u i s h e d a t the s u b s p e c i f i c o r  Sample  Number o f Platforms  Number o f Complete Platforms  Percentage of Complete Platforms  Number o f Ramiforms  Number o f P l a t f o r m s : Number o f Ramiforms  F48  28  1Q  36%  12  F49  70.5  60  9%  175-  F52  95  25  26%  48  1.98  F53  174  40  23%  32  5.44 : 1  F54  190  37  19%  25  7.60 : 1  Total  1192  172 :  14%  292  4.08 : 1  T a b l e 3.  Counts and percentages  2.33  :1  4.03 : 1  df p l a t f o r m s and ramiforms o f N e o g o n d o l e l l a  :1  i n F48 - F54.  52 S t a t i s t i c a l D a t a D e r i v e d f r o m Sample S u b s e t s F48  —Sample Parameters.  Data \ s .  h  N = 1, 2  M a  1  h  —  1Q6Q  a a  1  —  1  T a b l e 4.  N = 7, 9  733  752  811  75  44  75  88  863  816  923  984  1Q8  46  105  102  63.0  77.2  a  73.3 4.4  75.2 7.5  81.1 8.8  78.5  74.2  83.9  89.5  9.6  4.2  9.6  9.4  -  5  = 7.81  11 d e n t i c l e s  o  N = 11, 7  772  7  a  N = 6, 5  F54  = 105.3 M  10 d e n t i c l e s  F53  = 78.1 M  11 d e n t i c l e s  F52  N = 1 2 , 12  630  a  10 d e n t i c l e s  F49.  1  M  96.4  a  -  = 9.5?  S t a t i s t i c s from sample subsets (elements w i t h 10 and 11 d e n t i c l e s ) .  t-Test f o r  10  11  Denticles P-value**  t-score*  Samples Compared  a n d 1^/i  t-score*  Denticles P-value**  F49  -*• F52  1.00  0.17  0.84  0.21  F49  •* F53  0.61  0.27:  1.20  0.13  F49  -*- F54  1.05  0.15  2.60  0.009  0.58  0.28  2.75  0.005  CF49+F52) -»• (F53+F54)!  Table 5.  Values d e r i v e d from t - t e s t s on sample A  Mo  s  s i / 1 / % +.'1/N  2  = /  subsets  - CJJ+N2 02  0= standard d e v i a t i o n Efx - M N  Nj_+ N - 2 2  Efx  sum of t h e squares:of values f o r a character **= t h e p r o b a b i l i t y t h a t t h e d i f f e r e n c e between two sample means =  c o u l d occur by chance ( i . e . i f Z-score = 1.2, then P-value = .115, thus t h e r e i s a 11.5% chance t h a t t h e d i f f e r e n c e o c c u r r e d by chance a l o n e ) .  53 E • 15  F i g u r e 7a,b,c. Graphs showing the r e l a t i o n s h i p of the p l a t f o r m l e n g t h to the number of d e n t i c l e s per element f o r F49, F54 and F96.  .14r 1 3 - 1 2  a  i n 0  o  ' 1 0  0»(o/e  9  .  N. rosenkrantzi n.subsp. D ( F 0 6)  ~'*N. idahoensis n.subsp. A (F64) ( T o r  300  500  700 600 length of  2+speclmens)  1 IOC 1 300 platform (^jm)  7 . a  F  54  a n  d  F96  1600  1 «! o E' e1 5  e  .14 CI  i3 a  a  o -1 2  a  7b. Regions d e f i n e d by the c e n t r a l 3/4 of c l o s e l y spaced specimens f o r F49 and F54,  iiir o *10h  N. FS4  a ° E  8  F49  3 c  E  16  •  16  (central  specimens)  6  , '  500  9  700 000 length ol  1100 pisiform  • 1 3  o Z 12 c a> •o 1 1  o io •  of  7  3 00  •  Idahosnais n.subsp. A  0  >  09  9  E  +  .  am  7c.  7 N. i d a h o e n s i s n.subsp. A (F49) (1 or 2 s p e c i m e n s )  6  300  5 00  700  000  length  of  1100 platform  1300 (^im)  1600  F49  1300 (^m)  1600  54  specific level.  A second i n t e r e s t i n g o b s e r v a t i o n  l e n g t h from 10 t o 11 d e n t i c l e s f o r each sample.  i s t h e i n c r e a s e i n mean T h i s i n c r e a s e i s approx-  i m a t e l y 85 um f o r F49 and F52 and about 170 um f o r F53 and F54 ( T a b l e 4 ) . The  importance o f t h i s o b s e r v a t i o n w i l l be d i s c u s s e d  i n the s e c t i o n f o r  length/number o f d e n t i c l e s . Number o f D e n t i c l e s p e r Element (#) The mean number o f d e n t i c l e s i n c r e a s e s from F48 t o F53 but d e c r e a s e s m a r g i n a l l y f o r F54.  I f t h e number of d e n t i c l e s can be used as a "rough"  guide t o r e l a t i v e stages o f development and i f p l a t f o r m i s d i r e c t l y r e l a t e d to ontogenetic parameters should 3, F i g . 6 ) .  roughly  coincide.  then t h e t r e n d s f o r these two  T h i s i s not t h e case, however  (Table  T h i s i n d i c a t e s t h a t e i t h e r l e n g t h i s a more s e n s i t i v e i n d i -  cator of r e l a t i v e maturity explain the observations  or t h a t p l a t f o r m o v e r r e p r e s e n t a t i o n does not  suggesting  ution..:.: The d i f f e r e n c e s r e c o r d e d insignificant  stages,  overrepresentation  t h a t t h e changes a r e r e l a t e d t o e v o l -  i n t h e number o f d e n t i c l e s u p s e c t i o n a r e  ( s i g n i f i c a n c e i s considered  a t 5%) a c c o r d i n g  to z-scores  (Table 2 ) . R a t i o o f l e n g t h t o number o f d e n t i c l e s The  comparison o f two parameters t o g e t h e r  e s t i n g r e s u l t s with regards not be d e c i p h e r e d  (Ll/#)  tends t o produce some i n t e r -  to p o s s i b l e evolutionary i m p l i c a t i o n s that could  through a n a l y s i s o f t h e two parameters a l o n e .  T h i s i s not  s u r p r i s i n g as animals or. t h e i r s k e l e t a l remains a r e not d i f f e r e n t i a t e d one ters  another by t h i s o r t h a t c h a r a c t e r but r a t h e r by t h e sum o f many (Burma, 1948).  from  charac-  The r e s u l t s i n d i c a t e a d e c r e a s e i n t h e r a t i o o f l e n g t h  to number o f d e n t i c l e s f o r F48 through F52 f o l l o w e d by an i n c r e a s e i n samp l e s F53 and F54 ( F i g . 6 ) .  Z - t e s t s f o r t h e e n t i r e sample i n d i c a t e a  fail-  55 u r e o f s i g n i f i c a n c e even a t t h e 10% l e v e l sibility  (Table 2).  As a r e s u l t , t h e pos-  t h a t t h i s v a r i a t i o n could.be r e l a t e d t o a p r o c e s s  p l i n g cannot be d i s c o u n t e d .  F u r t h e r evidence  observed t r e n d may be s i g n i f i c a n t .  suggests,  elements w i t h  11 d e n t i c l e s  F u r t h e r u p s e c t i o n t o F96,. a v e r y sharp decrease i n t h e  Ll/# r a t i o o c c u r s which has a v e r y h i g h s t a t i s t i c a l Obviously,  however, t h a t t h e  The t - t e s t f o r L l / # i n d i c a t e s t h a t t h e  data a r e s i g n i f i c a n t a t t h e 5% l e v e l f o r those from F49 t o F54.  o f random sam-  s i g n i f i c a n c e (Table 5 ) .  t o proceed from,F54 t o F96, a t l e a s t one more p o i n t o f i n f l e c t i o n  i s r e q u i r e d t o obtain, this,>decrease. f l u c t u a t i n g mode o f e v o l u t i o n  What s t a r t s t o become apparent i s a  forthis particular ratio.  Combining  data  from Behnken (1975).with my data f o r l e n g t h v e r s u s number o f d e n t i c l e s , these  t r e n d s may be made g r a p h i c a l l y v i s i b l e  Neogondolella. i d a h o e n s i s N. r o s e n k r a n t z i (F96)  (F54)  ( F i g . 8 ) . The e v o l u t i o n o f  to _N. s e r r a t a and f i n a l l y t o N. p o s t s e r r a t a -  produces a n l i n i t i a l l y l a r g e decrease i n t h e l e n g t h  f o r an element o f g i v e n d e n t i c l e number f o l l o w e d by a s l i g h t  increase.  These major i n f l e c t i o n p o i n t s a r e thus s i g n i f i c a n t a t t h e s p e c i f i c  level.  Perhaps s i m i l a r l y shaped but s m a l l e r p o i n t s o f i n f l e c t i o n l i k e t h a t seen f o r F48 t o F54 a r e s i g n i f i c a n t a t t h e s u b s p e c i f i c l e v e l . Another f a c t o r t h a t becomes apparent from an a n a l y s i s o f t h e data i s t h a t t h e more mature t h e element i s ( i e . t h e l a r g e r and more d e n t i c l e s i t has)  t h e g r e a t e r one's a b i l i t y t o d i s t i n g u i s h between t h e p o p u l a t i o n  In support  o f t h i s statement a r e t - t e s t s f o r L l / # f o r 11 d e n t i c l e s ( T a b l e 5 ) .  As was p o i n t e d out e a r l i e r ,  F49 t o F52 i n c r e a s e s i n l e n g t h by 85 ym between  10 and 11 d e n t i c l e s whereas F53 and F54 i n c r e a s e by 170 ym. v a r i a t i o n s together and  samples.  Grouping  these  and t e s t i n g f o r t h e s i g n i f i c a n c e between F49 p l u s F52  F53 and F54 p r o v i d e d  the c l o s e s t r e s u l t s to s i g n i f i c a n c e f o r z - t e s t s  65*  * 300  1  500  * 700  length  F i g u r e 8.  * 900  • 1 1 00  1300  1 500  of p l a t f o r m (jum)  Graph showing t h e r e l a t i o n s h i p o f t h e p l a t f o r m l e n g t h t o the number o f d e n t i c l e s p e r element of N. i d a h o e n s i s n.subsp. A, _N. serrata,. and N. p o s t s e r r a t a ( i n c h r o n o l o g i c a l o r d e r ) . A f l u c t u a t i n g mode of e v o l u t i o n i s i n d i c a t e d by t h e d e c r e a s e i n l e n g t h f o r g i v e n d e n t i c l e number from _N. i d a h o e n s i s n.subsp. A t o N. s e r r a t a , f o l l o w e d by an i n c r e a s e i n l e n g t h from N. s e r r a t a to N. p o s t s e r r a t a . N_. i d a h o e n s i s n.subsp. A r e g i o n d e f i n e d by F49 t o F54 from E l l e s m e r e I s l a n d whereas N_. s e r r a t a and N_. p o s t s e r r a t a a r e based on d a t a presented i n Behnken, 1975.  57  for this interval  (however, the o n l y s i g n i f i c a n t t e s t s a t the 5% l e v e l were  the t - t e s t s f o r 11 d e n t i c l e s ) .  I t should be!obvious from F i g u r e 7 t h a t  the i n c r e a s e i n mean L l / / / from 11 to 12 d e n t i c l e s i s even more s i g n i f i c a n t than t h a t f o r 10 to 11 between F49  and  F54  as the f i e l d s r e p r e s e n t e d  by  the data p o i n t s become even more d i v e r g e n t . Length from t i p of cusp to f o u r t h d e n t i c l e a n t e r i o r of the cusp The  first  f o u r d e n t i c l e s a n t e r i o r of t h e cusp tend  spaced and more c i r c u l a r pressed  denticles.  to be more c l o s e l y  i n c r o s s s e c t i o n than the remaining  l a t e r a l l y com-  The maximum w i d t h of the element o f t e n o c c u r s at about  t h i s same p o i n t on the p l a t f o r m . analyze  Consequently, i t seemed a p p r o p r i a t e  t h i s parameter as i t c o u l d be v a l u a b l e even f o r fragmental  Except f o r F48  (L2)  (based  on the s m a l l e s t sample) the t r e n d i s one  i n g l e n g t h u p s e c t i o n s i m i l a r to t h e i n c r e a s e f o r L l . ( F i g .  9).  to  specimens.  of i n c r e a s This  simil-  a r i t y of t r e n d s . p o i n t s to the v a l u e of t h i s parameter f o r samples where o n l y fragmental  specimens a r e a v a i l a b l e . Maximum w i d t h  The ( F i g . 9).  (Wl)  data here i n d i c a t e a c o n s i s t e n t i n c r e a s e i n mean w i d t h u p s e c t i o n More d i s c u s s i o n w i l l  f o l l o w i n the s e c t i o n f o r L l / W l .  Width a t p o s t e r i o r end  (W2)  T h i s parameter shows a c o n s i s t e n t i n c r e a s e u p s e c t i o n f o r Wl  ( F i g . 9).  The main reason  f o r t a k i n g the two  s i m i l a r to t h a t  w i d t h measurements  f o r the d e t e r m i n a t i o n , of an a r e a f u n c t i o n i n the p o s t e r i o r 1/3  to 1/2  of  was the  denticle. Height D z i k and  from t i p o f cusp to base o f f l a n g e  (Hi)  Trammer (1980) found t h a t t h i s parameter was  discriminating T r i a s s i c Gondolella species.  The h e i g h t  very u s e f u l f o r  increases  upsection  -r3  2  i  i  i  1  i  i  2  4  u  «  6  5 3D  I  l  V!  B W x10"  *\  1  *  ± 50  1  1  I I  r2  W *10  1A  250  I  H  1-  4  *il  A  2  1 1  2 1  I  1 6  IC  JMhtSJkJ—I A  w. 3 1  1  3 I  h  IB  4 I —I 200  1  5 1  6 h  1 400 D  33E  B  • B  l>:-.v~-».v.'-'.'  Hi  A 50 1  H  1  1  1  250 1  h  H  1 4  4 1  1  5 1  1  6 1  1  7 1  1 8  2  1  °  •  F i g u r e 9.  „m  4  H 9  200 1  I 10  1  1 11  400 1 +-  3 B  L (W,+W )H 2  r-  A  2  jim  1 -j  3 i  •  4 •  -i  5  i_  6  8  9  10  11  • -j i__ G r a p h i c a l r e p r e s e n t a t i o n of p a r t of T a b l e 1. L i n e s i n f u l l b l a c k show range of M + 2 a . S t i p p l e d p a t t e r n = M + 1.15a ( i . e . 75% v a r i a t i o n ) . White = M + 2a (.i.e. 95% v a r i a t i o n ) . M  59 very c o n s i s t e n t l y . t h e two  w i d t h s and  So f a r i t has been e s t a b l i s h e d t h a t the two  the h e i g h t measurements a l l i n c r e a s e , w i t h some minor  f l u c t u a t i o n s , from F48 in  lengths,  to F54.  What i s i n t e r e s t i n g  , however, i s the  trend  the r a t i o s of these v a l u e s as they a l l d e c r e a s e u p s e c t i o n . Ll/Wl  Ratio  D e s p i t e t h e i n c r e a s e s i n b o t h l e n g t h and w i d t h the Ll/Wl r a t i o creases c o n s i s t e n t l y upsection  ( F i g . 9).  Obviously,  t h i s r e s u l t s from a  g r e a t e r i n c r e a s e i n w i d t h r e l a t i v e to the i n c r e a s e i n l e n g t h . be  reflected  elongated  and  dee-  T h i s would  i n a p l a t f o r m becoming more "square" i n shape r a t h e r than narrow.  changes from F48  to F53  Z-test scores f o r t h i s trend i n d i c a t e that and  from F49  (even f o r a two-sided t e s t ) .  to F53  the  a r e s i g n i f i c a n t at the 5%  I t i s impossible  to t e l l whether t h i s  level  trend  f l u c t u a t e s l i k e t h a t f o r Ll/# o r whether t h i s decrease i s t r a n s l a t e d through the e n t i r e s e c t i o n of Permian r o c k s d i s c u s s e d of N .  s e r r a t a ( ? ) suggest a Ll/Wl  r o s e n k r a n t z i v a r i e d between 4.0  ratios for N.  s e r r a t a (3.6), N . b i t t e r i ken',  1979;  C l a r k et a l . , 1979;  such comparisons, few  (3.6)  and  4.7,  averaging  B a s i n of the USA  idahoensis and  4.3.  illus-  similar  (3.9), Ni.. s e r r a t a (3.3), N .  post-  N . r o s e n k r a n t z i ( 4 . 0 ) . (,Cl?ark'-and Behn-  Wardlaw and  C o l l i n s o n , 1979b).  Admittedly,  stand on shaky ground as the measurements a r e based on a  specimens p r e s e l e c t e d by the above authors  similar intermediate  and which do not  to 3.05  take i n t o  I d i d t r y to  se-  to mature specimens from t h e f i g u r e d specimens).  Furthermore, samples from Texas (Behnken, 1975) d e c r e a s e from 4.3  fragments  A l o o k at  suggests a  account the v a r i a t i o n o f Ll/Wl d u r i n g ontogeny ( a l t h o u g h lect  Two  r a t i o : of about 4 w h i l e the r a t i o f o r N .  t r a t i o n s of specimens from the Great p a t t e r n of Ll/Wl  i n t h i s study.  f o r N . idahoensis  suggested a  continuous  to N . r o s e n k r a n t z i .  These  60 _N. r o s e n k r a n t z i a r e more e l o n g a t e Great  Basin.  Nevertheless  Probably  than those  a t the s p e c i f i c l e v e l f o r Ll/Wl  i n suggesting  trend  Ellesmere  Neogondolella..  Here a g a i n a l t h o u g h  Ratio  both L l and HI  L l / H l d e c r e a s e s f o r the same i n t e r v a l . the d i f f e r e n c e between F48 of random sampling.  to F53  Despite  i t i s impossible  i n c r e a s e fro'm F48  i s i n s i g n i f i c a n t and  the  ratio  i n d i c a t e that  c o u l d be the  result  t h i s , the t r e n d i s s t r i k i n g l y c o n s i s t e n t .  to t e l l  i f t h i s trend continues  ever,  the s h o r t e r l e n g t h of N.  n.sp.  .B would i n d i c a t e decreased and  to F53  However, z - t e s t s c o r e s  s e r r a t a ( ? ) and  I n t u i t i v e l y , how-  the h i g h , r o b u s t  L l / H l r a t i o s while  Once  f o r the remainder of  the s e c t i o n , nor a r e t h e r e data, to make i n f e r e n c e s from.  b i t t e r i n.subsp. C  the  a fluctuating  s i m i l a r to t h a t f o r Ll/# i n the  Ll/Hl  again  I s l a n d or  the t r e n d s v a r y w i t h environment and/or geography.  t h e r e seems to be some m e r i t  I s l a n d samples of  from E l l e s m e r e  cusps of  the s h o r t cusps of  N. N.  N. r o s e n k r a n t z i n.subsp. D would i n d i c a t e a subse-  quent i n c r e a s e i n the L l / H l  value.  L2(W1 + W2)% :—7 4 1 Q  „ P o s t e r i o r area :  Data f o r t h i s parameter i n d i c a t e s an i n c r e a s e i n t h i s a r e a v a l u e f o r F48  to F53.  Z-scores.::indicate t h a t the d i f f e r e n c e between F48  and  F53  s i g n i f i c a n t o n l y a t the 7 % l e v e l whereas the d i f f e r e n c e between F49  and  i s s i g n i f i c a n t a t the 2% l e v e l ,  even f o r a two  sided t e s t .  suggest t h a t t h i s a r e a v a l u e would be much l e s s f o r N. er a g a i n f o r N. n.sp. Formation. as w e l l .  is F53  Measurements  s e r r a t a ( ? ) but  great-  B to N. r o s e n k r a n t z i n.subsp. D i n the T r o l d F i o r d  Once a g a i n a f l u c t u a t i n g mode i s suggested f o r t h i s parameter  61  D i s c u s s i o n of E v o l u t i o n a r y Trends and Concepts Evidence f o r t h e L l / # r a t i o from p o p u l a t i o n samples o f N. i d a h o e n s i s n.subsp. A from E l l e s m e r e I s l a n d compared t o d a t a from p o p u l a t i o n of N. s e r r a t a and N. p o s t s e r r a t a  (Behnken,  samples  1975) from western USA i n d i c a t e  a f l u c t u a t i n g mode i n t h e e v o l u t i o n o f t h e s e p l a t f o r m elements w i t h r e s p e c t to t h i s parameter.  Furthermore, t h e r e s u l t s f o r N. i d a h o e n s i s n.subsp. A  suggest t h a t t h i s f l u c t u a t i n g mode may be important a t t h e s u b s p e c i f i c A l t h o u g h n o t backed up by s t a t i s t i c a l d a t a from l a r g e samples, u p s e c t i o n to N. i d a h o e n s i s which a r e c l o s e l y a l l i e d  level.  conodonts  to N. s e r r a t a , N. p o s t -  s e r r a t a and N. r o s e n k r a n t z i i n d i c a t e a s i m i l a r t r e n d to t h a t seen f o r t h e comparison w i t h Behnken's samples  (1975) from western USA.  In a d d i t i o n  s i m i l a r f l u c t u a t i n g t r e n d s seem apparent f o r s e v e r a l o t h e r parameters measured from t h e E l l e s m e r e I s l a n d samples. with a l l  I t t h e r e f o r e seems r e a s o n a b l e ,  t h i s s u p p o r t i n g documentation to suggest t h a t t h i s  fluctuating  tendency i s t h e r u l e r a t h e r than t h e e x c e p t i o n f o r t h e e v o l u t i o n o f Permian neogondolellids. Having proposed a mode f o r t h e e v o l u t i o n of t h e s e conodonts I have opened myself to t h e argument of i n t e r p r e t a t i o n o f t h i s t r e n d and; i n p a r t i c u l a r , am^obliged t o f a c e t h e q u e s t i o n o f p h y l e t i c g r a d u a l i s m (of which many a u t h o r s a r e proponents) v e r s u s punctuated e q u i l i b r i a Gould, 1972; Gould and.Eldredge, 1977).  ( E l d r e d g e and  As I pondered over t h i s  I t r i e d t o t a k e heed of. E l d r e d g e and Gould's warning  problem  (1972) t h a t a l l o b s e r -  v a t i o n i s c o l o u r e d by t h e o r y and e x p e c t a t i o n . My o r i g i n a l i n t e n t i o n f o r t h e s t a t i s t i c a l a n a l y s i s was t o see i f t h e r e was any q u a n t i t a t i v e d i f f e r e n c e between f i v e p o p u l a t i o n samples t h a t I c o u l d not, d i f f e r e n t i a t e  qualitatively  ( e s p e c i a l l y because o f t h e h i g h degree o f v a r i a b i l i t y ) and then determine  62  if  any d i f f e r e n c e s were s i g n i f i c a n t f o r t h e d e t e r m i n a t i o n  of  subspecies.  I t was not u n t i l w e l l i n t o t h e a n a l y s i s t h a t I saw t h e p o s s i b i l i t y of purporting  an e v o l u t i o n a r y  scheme.  I d i d not then, and I do not now w i s h  to get i n t o a deep p h i l o s o p h i c a l d i s c u s s i o n o f t h e m e r i t s other  scheme.  o f one or t h e  Rather I would l i k e to i n d i c a t e my d a t a and suggest an  i n t e r p r e t a t i o n and l e a v e  t h i s i n t e r p r e t a t i o n open t o c r i t i c a l a n a l y s i s by  other workers as p a r t o f t h e c o n t i n u i n g t h e s i s f o r t h i s group o f b i o t a .  p r o c e s s of e v o l u t i o n a r y model syn-  I c e r t a i n l y cannot c l a i m t h a t my d a t a would  ever s o l v e t h e problem of ...phyletic g r a d u a l i s m v e r s u s punctuated mode o f organic The  evolution. concept o f p h y l e t i c g r a d u a l i s m s t a t e s t h a t new s p e c i e s e v o l v e by  the slow and c o n t i n u o u s t r a n s f o r m a t i o n an unbroken g r a d a t i o n  of e n t i r e populations  o f f o s s i l forms ( E l d r e d g e  resulting i n  and Gould, 1972).  These  unbroken g r a d a t i o n a l  s e r i e s a r e r a r e l y ( i f ever) found because o f the "sup-  posed" i m p e r f e c t i o n s  i n the geologic  record.  e q u i l i b r i a " s t a t e s t h a t new s p e c i e s e v o l v e i s o l a t e d l o c a l populations i n the f o s s i l r e c o r d i t s ancestors  (allopatric  o f "punctuated  r a p i d l y from s m a l l , p e r i p h e r a l l y  s p e c i a t i o n ) r e s u l t i n g i n many breaks  s i n c e t h e new s p e c i e s e v o l v e  (Eldredge  and Gould, 1972).  t h e r e f o r e , n o t one o f s t a t e l y u n f o l d i n g , i b r i a , disturbed  The theory  i n an area remote from  The h i s t o r y of e v o l u t i o n i s , but a s t o r y o f homeostatic  equil-  o n l y r a r e l y by r a p i d and e p i s o d i c events o f s p e c i a t i o n  (ibid.). Since  1972 when E l d r e d g e and Gould f i r s t  published  their  "punctuated  e q u i l i b r i a " i n t e r p r e t a t i o n of e v o l u t i o n a few workers have come out t o support i t w h i l e many have come out i n o p p o s i t i o n , and contend t h a t research  data i n d i c a t e s p h y l e t i c gradualism.  their  T h i s prompted Gould and E l -  63 dredge  t o w r i t e a second paper  the success o f t h e i r model.  i n 1977 t o r e f u t e these c l a i m s and i n d i c a t e  I n o r d e r t o r e f u t e t h e i r punctuated  equili-  b r i a model a r e s e a r c h e r must show g r a d a t i o n a l s p e c i e s l e v e l l i n e a g e s w e l l p r e s e r v e d over t h e f u l l  span of an e x t e n s i v e geographic and temporal  range.  As Gould and E l d r e d g e (1977) a d e q u a t e l y p o i n t out most o f t h e c l a i m s f o r p h y l e t i c g r a d u a l i s m a r e based on l o c a l s e c t i o n s ( n o t t h e i r f u l l  geographic  range) and s h o r t d u r a t i o n (too s m a l l a s c a l e ) and a r e t h e r e f o r e i n v a l i d as t h e data a r e i n s u f f i c i e n t  f o r a test'.  I n defence o f t h e i r openmindedness  Gould and E l d r e d g e (1977) do accept one case o f g r a d u a l i s m as b e i n g v e r y impressive.  T h i s case i l l u s t r a t e d  t h e i n c r e a s e i n p r o l o c u l a r diameter o f a  v e r b e e k i n o i d f o r a m i n i f e r i n 34 r e l a t i v e l y l a r g e samples spanning t h e Middle to  Upper Permian, from southeast. A s i a , southern China and Japan.  As Gould  and E l d r e d g e (1977) s t a t e , "We.:are d e l i g h t e d w i t h these r e s u l t s as we expect some c o u n t e r c a s e s , e s p e c i a l l y among predominantly a s e x u a l forms".  Their  d i s c u s s i o n o f another g r a d u a l i s t i c case, t h a t o f G i n g e r i c h (1976) f o r E a r l y Eocene mammals i n n o r t h e r n Wyoming, i s o f p a r t i c u l a r  interest  t o t h i s work.  G i n g e r i c h (1976) c l a i m e d t h a t s p e c i e s o f t h e Eocene mammal Hyopsodus evolved i n a manner conforming  to Cope's r u l e  ( i n c r e a s i n g s i z e through'time)" based  on t h e i n c r e a s e u p s e c t i o n o f t h e l o g a r i t h m o f l e n g t h times w i d t h o f t h e first  lower molar.  However, t h i s o v e r a l l i n c r e a s e was a c h i e v e d o n l y a f t e r  a number o f f l u c t u a t i o n s  (of n i n e descendant  s m a l l e r s i z e and o n l y f o u r t o l a r g e r ) .  species, f i v e evolve  toward  Gould and E l d r e d g e (1977) c o u n t e r  t h a t G i n g e r i c h ' s s p l i t t i n g o f l i n e a g e s f i t s t h e i r model o f punctuated i b r i a better.  They found l o n g segments o f apparent  p o s e d l y g r a d u a l i s t i c sequences.  Furthermore,  s t a s i s w i t h i n h i s sup-  they s t a t e t h e f l u c t u a t i n g  p a t t e r n towards i n c r e a s e i n t o o t h s i z e c o n f i r m s t h e most important c a t i o n o f punctuated  equilibria,  equil-  impli-  t h a t s p e c i a t i o n i s e s s e n t i a l l y random w i t h  64 respect  to the d i r e c t i o n of a m a c r o e v o l u t i o n a r y t r e n d  Wright, 1967).  Stanley  (Wright's r u l e ,  (1975) wrote t h a t m a c r o e v o l u t i o n a r y t r e n d s a r e  a r e s u l t of g r a d u a l i s t i c o r t h o s e l e c t i o n , but a r i s e from a "higher  not  level  s e l e c t i o n " o f c e r t a i n morphologies from a random pool, o f s p e c i a t i o n events produced by punctuated e q u i l i b r i a .  According  to.Gould and  E l d r e d g e (1977)  the phylogeny of Hyopsodus af f i r m s :;Wright' s r u l e where s i z e i n c r e a s e i n the e n t i r e c l a d e a r o s e from the d i f f e r e n t i a l , success random subset  of c l a d i s t i c  events.  Neogondolella  i d a h o e n s i s n.subsp. A  I t i s my  o p i n i o n t h a t the phylogeny of  to _N.. r o s e n k r a n t z i n.subsp. JJ e q u a l l y  supports the model of punctuated e q u i l i b r i a and data d i s p l a y e d on F i g u r e 8 i s based on N. West Texas (from Behnken, 1975) ern E l l e s m e r e  Island.  and  of l a r g e r s p e c i e s i n a  a f f i r m s Wright's r u l e .  The  s e r r a t a and _N. p o s t s e r r a t a from  on _N.. i d a h o e n s i s  The m a t e r i a l thus c o v e r s  n.subsp. A from n o r t h -  a l a r g e geographic a r e a  a s i g n i f i c a n t p o r t i o n of the temporal range of the N e o g o n d o l e l l a  serrata  complex, t h a t of the l a t e Lower Permian through M i d d l e Permian (about m i l l i o n y e a r s ) , both n e c e s s a r y p r e r e q u i s i t e s f o r an adequate t e s t . data  i n d i c a t e s an o v e r a l l i n c r e a s e i n number of d e n t i c l e s per u n i t  upsection  but o n l y a f t e r a f l u c t u a t i n g p a t h where N.  The length  postserrata.  I s l a n d f a l l s w i t h i n the same  field  represented  ever,  a l l of t h e s e members of the c l a d e have a l a r g e r v a l u e  by data p o i n t s f o r N. p o s t s e r r a t a from West Texas.  number per u n i t l e n g t h than t h a t f o r the a n c e s t r a l . f o r m , subsp. A.  Having accepted  15  s e r r a t a has more  d e n t i c l e s per.:unit l e n g t h than the r e s u l t i n g descendent N. N. r o s e n k r a n t z i n.subsp. D from E l l e s m e r e  and  N.  How-  for denticle idahoensis  a punctuated e q u i l i b r i a mode of e v o l u t i o n to  n. ex-  p l a i n the f l u c t u a t i n g p a t t e r n of d e n t i c l e number per. u n i t l e n g t h I must a l s o accept  some of the other  f e a t u r e s of the model; namely, t h a t between  t h e s e s p e c i a t i o n events the forms d i d not change, t h a t i s they underwent a  65  p e r i o d of s t a s i s . sistent tively  However, a t f i r s t  g l a n c e I cannot do  t r e n d s were apparent i n the samples F48 in detail  Gould and  (Table  to F54  t h i s , as some constudied quantita-  1).  Eldredge  (1977) i n d i c a t e t h a t the norm f o r a s p e c i e s d u r i n g  the heyday of i t s e x i s t e n c e as a l a r g e p o p u l a t i o n i s m o r p h o l o g i c a l minor n o n - d i r e c t i o n a l f l u c t u a t i o n b e a r i n g no r e l a t i o n s h i p species.  The  stasis,  i n form, or minor d i r e c t i o n a l change  to pathways of a l t e r a t i o n i n subsequent daughter  n a t u r e o f degree of t h i s minor change can be best  understood  by r e a l i z i n g t h a t the r a p i d i t y o f s p e c i a t i o n i n such a model does not q u i r e the i n t e r m e d i a t e My  stage o f a r e c o g n i z a b l e s u b s p e c i e s  naming the s t u d i e d p o p u l a t i o n s r e p r e s e n t e d  s p e c i e s of N e o g o n d o l e l l a  idahoensis  by F48  n i f i c a n t a t the s u b s p e c i f i c l e v e l . (15 m of a 200  m s e c t i o n ) may  The  and  time r e p r e s e n t e d  as a new  time s c a l e s f o r the Permian).  for  N.  t h a t t h i s may  t h a t these a r e from F48  (assuming c o n t i n u o u s  C l a r k and  to  h i g h by t h i s a u t h o r ) .  average d u r a t i o n d f 2 3 . 3 m i l l i o n y e a r s f o r N. ed between F48  and  F54  extended to the presumed f u l l  sedimenta-  those  Even i f we  figure  assume the  is insignificant,  In  i t could,  range of the s p e c i e s , become s i g n i f i c a n t .  between samples were i n s i g n i f i c a n t . into just  (the l a t t e r  million  i d a h o e n s i s , the time r e p r e s e n t -  to F54  Z - t e s t s f o r the parameters L l , #, and  ated  F54  i s . o n l y a s m a l l f r a c t i o n of t h i s temporal range.  o t h e r words even i f the change from F48 if  sig-  Behnken (1979) i n d i c a t e  v a r y from 2 to 10 m i l l i o n years  idahoensis - considered  sub-  recognizable  t h a t the average s p e c i e s d u r a t i o n f o r Permian n e o g o n d o l e l l i d s i s 3.3 y e a r s but  1977).  r e p r e s e n t as much as one m i l l i o n y e a r s but i s  p r o b a b l y more on the o r d e r of 500,000 y e a r s t i o n and  to F54  i n d i c a t e s that I f e e l  changes do occur between the r a p i d s p e c i a t i o n events  (Stebbins,  re-  elements w i t h  Ll/// i n d i c a t e d that d i f f e r e n c e s  However, when the samples were 11 d e n t i c l e s a t - t e s t  separ-  indicated that  66 d i f f e r e n c e s between F49 and F54 were v e r y s i g n i f i c a n t .  This s i g n i f i c a n c e  a l o n e c o u l d not be used, t o d i f f e r e n t i a t e between s u b s p e c i e s but may c a t e t h a t d i f f e r e n c e s between e n t i r e samples, temporal range, c o u l d become s i g n i f i c a n t .  i f extended  indi-  through g r e a t e r  A l t h o u g h Gould and E l d r e d g e (1977)  do a l l o w f o r some minor changes i t i s w i t h the amount t h a t many a u t h o r s seem to be a t odds w i t h them.  I would p r e f e r to b e l i e v e that between these  r a p i d s p e c i a t i o n events some g r a d u a l i s m does o c c u r . i n d i c a t e d t h a t s p u r i o u s " p h y l e t i c change" may s u c c e s s i v e immigration of normal l o c a l environments.  However, Newell  (1956)  a r i s e i n l o c a l s e c t i o n s by  geographic v a r i a n t s r e s p o n d i n g to changing  I f t h e environment  was  changing i n a p r o g r e s s i v e man-  ner (eg. s h a l l o w i n g d u r i n g r e g r e s s i o n ) then samples u p s e c t i o n would change i n one d i r e c t i o n w i t h r e s p e c t to a parameter.  Given the g e n e t i c and p h y s i o -  l o g i c a l c o m p l e x i t y of any. p o p u l a t i o n of organisms, many d i f f e r e n t ways o f a d j u s t i n g to a new  f a c t o r o f the environment  a r e p o s s i b l e ( S t e b b i n s , 1977),  s u g g e s t i n g t h a t any change i n a l o c a l s e c t i o n w i t h r e s p e c t t o  environment  need not be accompanied by a s i m i l a r c o n s i s t e n t g r a d u a l change i n b i o t i c response.  I f the immigrants  (normal geographic v a r i a n t s ) respond  i n many  ways to e n v i r o n m e n t a l change then the r e s u l t s a r e u n l i k e l y t o be c o n s i s t e n t u p s e c t i o n and u n i d i r e c t i o n a l .  A form of s e l e c t i o n d i f f e r e n t  from n o n - d i r e c -  t i o n a l ( s t a b i l i z i n g ) o r d i r e c t i o n a l i s t h a t of d i v e r s i f y i n g s e l e c t i o n where, if  environmental h e t e r o g e n e i t y i s i n c r e a s i n g over time, the response o f the  p o p u l a t i o n w i l l be to become more heterogeneous  withrrespect to-various par-  ameters r e s u l t i n g i n a once homogeneous p o p u l a t i o n b r e a k i n g up i n t o d i f f e r e n t l y adapted  s u b u n i t s ( S t e b b i n s ; 197.7).  p o p u l a t i o n s from F48 to F54  several  The v a r i a b i l i t y of those  seems to be i n c r e a s i n g as evidenced by the a l -  most c o n s i s t e n t l y i n c r e a s i n g standard d e v i a t i o n of the samples u p s e c t i o n . How  t h e s e two  types of s e l e c t i o n , d i r e c t i o n a l and d i v e r s i f y i n g ;  interact  67  and whether t h e r e s u l t i n g g r a d a t i o n u p s e c t i o n can s t i l l s p u r i o u s r a t h e r than r e a l ,  isdifficult  many c o m p l i c a t i n g . f a c t o r s , o b v i o u s l y .  be regarded as  t o say u n e q u i v o c a l l y . Perhaps  There a r e  e a s i e r t o demonstrate  would  be whether o r n o t t h e minor change i n form bears a r e l a t i o n s h i p t o t h e e v o l u t i o n o f subsequent  daughter  species.  I f one c o n s i d e r s t h e i n t e r v a l r e p r e s e n t e d between N. i d a h o e n s i s and N. s e r r a t a t h e expected and  t r e n d s would be towards o v e r a l l d e c r e a s i n g s i z e  i n c r e a s i n g d e n t i c l e number.  The r e s u l t s f o r F48 t o F54, which r e p r e -  s e n t s p a r t o f t h i s j u s t mentioned  i n t e r v a l , depict  i n c r e a s i n g l e n g t h , width,  h e i g h t and p o s t e r i o r a r e a u p s e c t i o n ( e x a c t l y o p p o s i t e t h e e x p e c t a t i o n ) and o v e r a l l i n c r e a s e but f l u c t u a t i n g d e n t i c l e number (approximates t h e expected trends).  These b e t w e e n - s p e c i a t i o n t r e n d s c o u l d be regarded as random o r  chance events i n the developmental  pathway because o n l y one of t h e two  t r e n d s bears any . r e l a t i o n s h i p t o t h e s p e c i a t i o n t r e n d s . I t seems a p p r o p r i a t e a t t h i s time t o compare t h e r e s u l t s o f a study by D z i k and Trammer (1980) which i n many r e s p e c t s i s s i m i l a r to t h i s one. T h e i r a n a l y s i s i s t h e r e s u l t o f study o f 25 samples over about  23 metres  of s e c t i o n from t h e Holy Cross Mountains  o f Poland which c o n t a i n T r i a s s i c  n e o g o n d o l e l l i d s ( g o n d o l e l l i d s t o them).  Their r e s u l t s indicate a general  d e c r e a s e i n d e n t i c l e number and l e n g t h but a l o n g a f l u c t u a t i n g p a t h .  They  i n t e r p r e t t h e i r r e s u l t s as t h e r e s u l t o f p h y l e t i c g r a d u a l i s m and not o f punctuated e q u i l i b r i a .  However, t h e s e r e s u l t s f a i l  i n p r o v i d i n g an ade- -. 1  quate t e s t s i n c e they r e p r e s e n t a l o c a l s e c t i o n o f s h o r t d u r a t i o n .  I n such  a s e c t i o n one would not expect t o see t r e n d s t h a t a r e t h e r e s u l t of punctuated e q u i l i b r i a .  Furthermore,  t h e f l u c t u a t i n g p a t h ( i f i t were over a  l a r g e r time frame) c o u l d be b e t t e r i n t e r p r e t e d  i n a punctuated  equilibria  68 model.  These r e s u l t s c o u l d be i n t e r p r e t e d as " s p u r i o u s " d i r e c t i o n a l  t i c change because they a r e from a l o c a l s e c t i o n .  Furthermore, a compli--  c a t i n g f a c t o r p r e v i o u s l y d i s c u s s e d , t h a t of i n c r e a s i n g v a r i a b i l i t y , not seem t o be t h e case h e r e .  does  They i n d i c a t e , however, t h a t p l a t f o r m cono-  donts i n the uppermost Muschelkalk Holy  phyle-  of Germany ( s l i g h t l y younger than  Cross Mountain specimens) r e p r e s e n t i n morphology a p r o g r e s s  the t r e n d d i r e c t i o n r e c o g n i z e d i n the conodonts from the Holy a i n s s u g g e s t i n g t h a t t h e i r t r e n d s were not " s p u r i o u s " and  the  along  Cross Mount-  t h a t the  popular,  t i o n s of G o n d o l e l l a i n h a b i t i n g the C e n t r a l European b a s i n were e v o l v i n g s i m u l t a n e o u s l y and  r e g a r d l e s s of l o c a l f a c i e s changes.  D e s p i t e the  t h a t the time frame r e p r e s e n t s a l a r g e f r a c t i o n of the Middle the demonstrated e v o l u t i o n i s of t h r e e temporal subspecies, a r e r e p r e s e n t e d the uppermost and  i n the l o c a l Holy  lowermost samples.  nomic  Two  of  these  e a r l i e r the t r e n d s a t  These r e s u l t s g i v e me  b i a s t h a t the changes from F48  s u b s p e c i f i c rank and  Triassic,  Cross Mountain s e c t i o n - i n  As i n d i c a t e d  t h i s taxonomic l e v e l should be more g r a d u a l . reason to a c c e p t my  subspecies.  fact  to F54  some  are r e a l at  t h a t g r a d u a l changes should be expected  at t h i s  the taxo-  level. In summary, the r e s u l t s f o r the N e o g o n d o l e l l a  the A s s i s t a n c e and Arctic  T r o l d F i o r d Formations  s p e c i e s d e s c r i b e d from  of n o r t h e r n E l l e s m e r e I s l a n d ,  Canada compared to s p e c i e s from the Great B a s i n of the western  a r e b e s t i n t e r p r e t e d as the r e s u l t of e v o l u t i o n c o n s i s t e n t w i t h a e q u i l i b r i a model.  The  punctuated  r e s u l t s f o r a s m a l l f r a c t i o n of t h i s i n t e r v a l seem  to i n d i c a t e t h a t d i r e c t i o n a l and/or d i v e r s i f y i n g s e l e c t i o n r e s u l t changes a t the s u b s p e c i f i c l e v e l . t i o n to the punctuated  USA,  i n gradual  T h i s should not. be c o n s i d e r e d i n o p p o s i -  e q u i l i b r i a model but regarded  as a f e a t u r e t h a t  en-  69  hances t h e r e s u l t a n t changes d u r i n g t h e r a p i d but ..punctuated s p e c i a t i o n events.  J u s t as Gould and E l d r e d g e  gradual  (1977) r e g a r d  s t a s i s as r e a l so should  change between punctuated s p e c i a t i o n events be regarded  at t h e s u b s p e c i f i c l e v e l .  as a r e a l i t y  T h i s does not seem to. be an u n r e a s o n a b l e s t a t e -  ment when one c o n s i d e r s t h e many v a r y i n g e v o l u t i o n a r y s t y l e s demonstrated by d i f f e r e n t b i o t i c  forms. SYSTEMATIC PALEONTOLOGY  Introduction There a r e a number o f problems w i t h t h e d e s i g n a t i o n o f t h e genus Neogondolella  t h a t make i t s concept u n c l e a r .  I t was o r i g i n a l l y  erected  f o r forms t h a t developed from t h e genus Spathognathodus (now Neospathodus) i n t h e Lower T r i a s s i c . Lower T r i a s s i c  Subsequently, Upper C a r b o n i f e r o u s ,  s p e c i e s have.been a s s i g n e d  i n c l u d i n g a l l American a u t h o r s . f o r a l l o f these forms. one  to N e o g o n d o l e l l a  Permian and by many  Kozur (1968) r e t a i n s t h e genus G o n d o l e l l a  The p r e s e n t  controversy  over t h e d e s i g n a t i o n i s  o f the opposing views between t h e European and North American The  revised diagnosis  authors  "schools".  (Sweet, 1970; f i d e . Z i e g l e r , 1973) i n c l u d e s  conodont s p e c i e s i n which t h e s k e l e t a l apparatus comprises elements of a s i n g l e morphologic type. i n d i v i d u a l l y asymmetrical,  " These elements, which a r e e l o n g a t e , have a t e r m i n a l o r s u b t e r m i n a l  p a i r e d , and  p o s t e r i o r cusp;  a median nodose o r d e n t i c u l a t e c a r i n a ; and f i n e l y t o c o a r s e l y p i t t e d , l a r g e l y unornamented, p l a t f o r m l i k e l a t e r a l e x t e n s i o n s ,  which a r e j o i n e d  p o s t e r i o r l y i n most s p e c i e s by a more o r l e s s w e l l developed brim encloses  1  the p o s t e r i o r end of t h e c a r i n a .  that  Underside of elements marked  by a l o n g i t u d i n a l l y grooved k e e l t h a t widens p o s t e r i o r l y t o e n c l o s e .a p i t beneath t h e cusp ( Z i e g l e r ,  1973, p. 127-128)."  Kovacs and Kozur (1980)  70 suggest t h a t t h i s d i a g n o s i s of N e o g o n d o l e l l a insufficient 1932.  to s e p a r a t e  Bender and  Stoppel,  t h i s genus from G o n d o l e l l a S t a u f f e r and  is  Plummer,  I t appears t h a t many of these f e a t u r e s ( p o s t e r i o r brim, ornamenta-  t i o n ) a r e v a r i a b l e w i t h i n the phylogeny of t h i s group and members (eg. smooth and and  1965  p i t t e d p l a t f o r m of N.  p i t t e d p l a t f o r m of N.  of N. p o s t s e r r a t a ) .  s e r r a t a and  Kovacs and  f o r s e p a r a t i o n of the two  ontogeny of i t s  idahoensis versus  serrated  l a c k of s e r r a t i o n i n e a r l y ontogeny  Kozur (1980) i n d i c a t e t h a t the main argument  genera i s the assumption t h a t N e o g o n d o l e l l a  has  a s i n g l e element apparatus ( p l a t f o r m o n l y ) whereas G o n d o l e l l a has a m u l t i element a p p a r a t u s ( p l a t f o r m p l u s r a m i f o r m s ) .  Because of t h e  conflicting  o p i n i o n of a number of s e n i o r workers i t i s not c l e a r which argument greater merit.  Von  B i t t e r and  M e r r i l l ' s ' (19/7) s u g g e s t i o n  had  a multielement  and  One  of the reasons f o r t h i s c o n f l i c t i n g o p i n i o n i s the  phenomenon of p l a t f o r m o v e r r e p r e s e n t a t i o n ; i n any  although  r e j e c t e d by  Pennsyl-  vanian Neogondolella Behnken (1979).  apparatus was  that a  has  the frequency  l o c a l s e c t i o n , i t g e n e r a l l y i n c r e a s e s from C a r b o n i f e r o u s  M e r r i l l and  Powell  in Missourian  (1980) have shown t h a t t h i s p l a t f o r m  Gondolella i s probably  of o n l y ramiform apparatuses, comprising t h i s low  platform only.  Clark  varies  to  Triassic.  overrepresentation  the r e s u l t . o f an o n t o g e n e t i c  series  to.ramiform p l u s p l a t f o r m , to apparatuses  I n o t h e r words, they suggest a mechanism f o r  index of mutual o c c u r r e n c e  which i n d i c a t e s - t h a t a pure  statistical  a n a l y s i s c o u l d l e a d to s e r i o u s m i s t a k e s i n the combination of conodont apparatuses.  They agree w i t h Kovacs and Koziir's (1980) o p i n i o n of a m u l t i -  element apparatus f o r both of these genera but a t the same time r e t a i n d e s i g n a t i o n of N e o g o n d o l e l l a .  Von  Bitter  (1976) suggests t h a t the  l a t e d m i c r o s t r u c t u r e i s d i s t i n c t i v e ; c o v e r i n g much more of the o r a l  the  reticusurface  71  in. N e o g o n d o l e l l a . elements  Sweet (1970) i n d i c a t e d t h a t i f N e o g o n d o l e l l a c o n t a i n e d  o f a s i n g l e type then N e o g o n d o l e l l a was fundamentally  from G o n d o l e l l a .  different  E v i d e n c e seems t o i n d i c a t e t h a t t h e two genera a r e not  fundamentally d i f f e r e n t but t h a t minor d i f f e r e n c e s do e x i s t  (reticulation,  v a r i a b l e but much l e s s r i b b e d o r s e r r a t e d ornament) t o t h e extent t h a t many a u t h o r s ( i f n o t most: Kozur t h e n o t a b l e e x c e p t i o n ) r e t a i n t h e genus Neogondolella . Furthermore,  t h e o r i g i n and phylogeny  o f t h e genus remains  clouded.  The o r i g i n a l N e o g o n d o l e l l a was e r e c t e d a s a form d e r i v e d from Neospathodiis but Mosher (1968) ..indicates t h a t N. mombergensis (the type s p e c i e s ) i s not r e l a t e d t o Neospathodus making t h e o r i g i n a l d i a g n o s i s i m p r a c t i c a l . ski  Szaniaw-  and Malkowski (1979) i n d i c a t e t h a t t h e e v o l u t i o n a r y development o f t h e  Permian n e o g o n d o l e l l i d s shows c l e a r l y that, they r e p r e s e n t one p h y l o g e n e t i c branch and t h a t t h e r e c o g n i z e d ontogeny bears no r e l a t i o n s h i p t o Neospathodus.  C l a r k and Behnken (1979) i n d i c a t e t h a t N. b i s s e l l i ,  the ancestor of  a l l Permian and younger s p e c i e s , o c c u r s s e v e r a l hundred f e e t above e l l a b e l l a w i t h which i t bears l i t t l e morphologic et  a l . (1979, f i d e . Kovacs and Kozur,  similarity.  Gondol-  Movshovich  1980) i n d i c a t e t h a t "G". p r a e b i s s e l l i  i s i n t e r m e d i a t e between G. b e l l a and "G". b i s s e l l i .  There appears  at pre-  sent no c l e a r s o l u t i o n f o r a student a t t e m p t i n g t o d e c i p h e r these r e p o r t s i n terms o f an adequate phylogeny.  I t does seem c l e a r t h a t more work i s  n e c e s s a r y from an u n b i a s e d p o i n t o f view t o determine d i f f e r e n c e s between N e o g o n d o l e l l a and G o n d o l e l l a .  the s i g n i f i c a n c e of  Only a t t h i s p o i n t c o u l d  the e v o l u t i o n o f t h e s e forms be d e c i p h e r e d . U n t i l a r e v i s e d d i a g n o s i s i s p r e s e n t e d t h a t i s w i d e l y acknowledged and r e f u t e s t h e genus N e o g o n d o l e l l a , t h i s author r e f e r s h i s g o n d o l e l l i f o r m e l e -  72 merits t o s p e c i e s o f N e o g o n d o l e l l a as diagnosed  i n Ziegler  (1973).  Using  t h i s d i a g n o s i s i n d i c a t e s t h a t d e s c r i p t i o n s f o r t h e s p e c i e s a t hand assume a s i n g l e element-type  apparatus.  For t h i s r e a s o n the p l a t f o r m elements o f  N e o g o n d o l e l l a w i l l be d i s c u s s e d s e p a r a t e l y from t h e ramiforms. t h a t t h e ramiform  elements  c o u l d belong  apparatus a d u a l taxonomic nomenclature follows:  to multielement i s adopted  M u l t i e l e m e n t s p e c i e s - Form s p e c i e s .  i s somewhat u n s a t i s f a c t o r y but i t does r e f l e c t  Recognizing  "Neogondolella"  f o r these elements as  T h i s method o f d e s c r i p t i o n t h e p r e s e n t c o n t r o v e r s y and  the d e s i r e by t h e author t h a t t h e q u e s t i o n be r e s o l v e d i n t h e near Should t h e genus be found t o have a multielement  future.  apparatus then t h e form'  s p e c i e s name should be p l a c e d i n synonymy w h i l e i f t h e genus i s found t o have.a s i n g l e element apparatus then t h e m u l t i e l e m e n t be dropped.  D e s c r i p t i o n o f t h e s e ramiform  elements  s p e c i e s name should  w i l l f o l l o w that of the  platforms of Neogondolella. As t h e p l a t f o r m elements  e v o l v e d r a p i d l y and a r e c o n s e q u e n t l y  strati-  g r a p h i c a l l y v e r y important they a r e d e s c r i b e d i n d e t a i l , whereas t h e ramiform elements  e v o l v e d s l o w l y and are, as a r e s u l t , o f l i t t l e v a l u e  1  strati-  g r a p h i c a l l y and thus a r e not d e s c r i b e d i n d e t a i l . The p l a t f o r m elements o l o g i c a l order.  of Neogondolella species a r e described i n chron-  Two o t h e r genera,  Neostreptognathodus  i n c l u d i n g Anchignathodus  p r a y i are discussed f i r s t  minutus and  because of t h e i r  association  w i t h t h e o l d e s t N e o g o n d o l e l l a p l a t f o r m s p e c i e s (N. i d a h o e n s i s ) .  These two  genera a r e o n l y b r i e f l y d e s c r i b e d as they a r e unimportant - i n t h e a r e a owing to t h e i r slow e v o l u t i o n a r y r a t e s and v e r y r a r e o c c u r r e n c e ,  respectively.  In summary, t h e o r d e r o f appearance w i l l be Anchignathodus Neostreptognathodus' p r a y i , p l a t f o r m elements c i e s , and f i n a l l y t h e ramiform  elements  .  minutus,  o f v a r i o u s N e o g o n d o l e l l a spe^  73  Systematics Genus  .ANGHIGNATHODUS. Sweet, 1971  Type s p e c i e s  Anchignathodus minutus ( E l l i s o n ) , 1941 ASGHIGNATHODUS MINUTUS ( E l l i s o n ) , 1941 P I . 1, f i g s . 3-6. Spathodus minutus E l l i s o n , 1941  Occurrence: F49,  Lower A s s i s t a n c e Formation,  Hamilton P e n i n s u l a s e c t i o n (F48,  F52, F53, F54, F63 arid F 7 5 ) .  Description:  T h i s element possesses  s l i g h t l y curved  a short, t h i n , l a t e r a l l y s t r a i g h t to  b l a d e about t h r e e times as l o n g as wide and w i t h s i x t o  n i n e l a t e r a l l y compressed, subequal,  p a r t l y fused d e n t i c l e s p o s t e r i o r t o .  the cusp and zero t o t h r e e s h o r t d e n t i c l e s a n t e r i o r t o t h e cusp.  The den-  t i c l e s a r e o f f s e t a b r u p t l y t o t h e cusp which i s l a r g e and t r i a n g u l a r i n outline.  The b a s a l c a v i t y o f t h e b l a d e  i s broadly f l a r e d  e s p e c i a l l y under t h e d e n t i c l e s p o s t e r i o r t o t h e cusp.  i n t h e mid r e g i o n ,  The c a v i t y  reduces  to a narrow groove a t both t h e a n t e r i o r and p o s t e r i o r ends o f t h e a b o r a l surface. ond  The deepest p o i n t o f t h i s b a s a l c a v i t y i s below t h e f i r s t  or s e c -  d e n t i c l e p o s t e r i o r t o t h e main cusp.  Discussion:  Representatives  o f t h i s s p e c i e s range from C h e s t e r i a n  (Late  M i s s i s s i p p i a n ) t o Roadian ( e a r l y M e d i a l Permian) i n age (Behnken, 1975; Ziegler,  1973).  by t h e abrupt  They a r e d i f f e r e n t i a t e d from t h e younger A. t y p i c a l i s  offset  i n l a t e r a l p r o f i l e p o s t e r i o r to t h e cusp as opposed  to a g r a d u a l d i m i n u t i o n of t h e l a t e r a l Genus  NEOSTREPTOGNATHODUS'- C l a r k , 1972  Type s p e c i e s 1951)  profile.  Streptognathodus s u l c o p l i c a t u s (Youngquist,  Hawley and M i l l e r ,  74  NEOSTREPTOGNATHODUS PRAYI Behnken, Pl. Occurrence: Description:  1, f i g s .  1,2.  A s s i s t a n c e Formation, McKinley Bay T h i s element  1975  section  c o n s i s t s of a subsymmetrical,  (F100). posteriorly  p o i n t e d p l a t f o r m w i t h c l o s e l y spaced, s u b p a r a l l e l , t r a n s v e r s e r i d g e s on the o r a l s u r f a c e which extend almost to c o m p l e t e l y a c r o s s the m e d i a l Discussion: appear  No a n t e r i o r f r e e b l a d e s were observed.  fragments  to r e p r e s e n t forms i n t e r m e d i a t e i n ontogeny, f o l l o w i n g  by Behnken (1975).  occurrence of t h i s species i s Latest  NEOGONDOLELLA" Bender and S t o p p e l ,  Type s p e c i e s  descriptions  A c c o r d i n g to Wardlaw and C o l l i n s o n (1979a) and C l a r k  et a l . (1979) the youngest Genus  The two  groove.  G o n d o l e l l a mombergensis  1965  (Tatge)  NEOGONDOLELLA IDAHOENSIS (Youngquist,-Hawley "- .  Leonardian.  P l . 1,. f i g s .  and M i l l e r ,  1951)  subsp.  indet.  7-13.  G o n d o l e l l a i d a h o e n s i s Youngquist, Hawley and M i l l e r ,  1951  G o n d o l e l l a p h o s p h o r i e n s i s Youngquist, Hawley and M i l l e r , Occurrence:  A s s i s t a n c e Formation, McKinley Bay  section  Discussion:  These specimens a r e v e r y s i m i l a r to N.  1951  (F100).  i d a h o e n s i s n.subsp. A  except t h a t no s e r r a t i o n s or d i s t i n c t v a r i e t i e s were r e c o g n i z e d .  This i s  p a r t l y owing to the low number of specimens and poor p r e s e r v a t i o n ( r e c r y s t a l l i z e d ) . . For these reasons and because tognathodus  of i t s o c c u r r e n c e w i t h N e o s t r e p -  p r a y i ( s u g g e s t i n g a s l i g h t l y o l d e r age from N e o g o n d o l e l l a i d a h o -  e n s i s n.subsp. A) a s u b s p e c i f i c d e t e r m i n a t i o n was t i o n i s s i m i l a r t o t h a t of N. t i e s ) which f o l l o w s .  not made.  Their  descrip-  i d a h o e n s i s n.subsp. A (except f o r the varied",  NEOGONDOLELLA IDAHOENSIS. n.subsp. A PI. 2, f i g s . Occurrence: F49,  9-19; P i . 3; Pi.' 4; P i . 5.  Lower A s s i s t a n c e Formation, Hamilton P e n i n s u l a  F52, F53, F54).;-  Diagnosis:  T h i s subsymmetrical u n i t has a c a r i n a composed of 6 t o 16  d e n t i c l e s (average 10 t o 11)  which a r e d i s c r e t e i n e a r l y growth  becoming p a r t i a l l y fused and f i n a l l y c o m p l e t e l y The  s e c t i o n (F48,  p l a t f o r m elements a r e v e r y v a r i a b l e , r a n g i n g  fused  stages,  i n g e r o n t i c forms.  from l o n g and s l e n d e r  forms  to robust and s e r r a t e d . . The maximum w i d t h t y p i c a l l y o c c u r s a t some p o i n t i n t h e p o s t e r i o r h a l f , a f t e r which t h e element t a p e r s g r a d u a l l y and f i n a l l y r a p i d l y i n the a n t e r i o r h a l f .  Except f o r t h e a n t e r i o r t i p t h e p l a t f o r m  margin i s r e t i c u l a t e d i n a l l examples.  This r e t i c u l a t i o n although  absent from t h e w e l l developed furrows and c a r i n a , i s p r e s e n t some mature r o b u s t  forms.  During  generally  on these i n  ontogeny t h e k e e l on t h e a b o r a l s i d e  changes from narrow and h i g h , t e r m i n a t i n g i n an o v a l l o o p t o wide and low with a terminal t r i a n g u l a r loop. The  cusp i s l a r g e , e r e c t and i n c l i n e d p o s t e r i o r l y w h i l e t h e f o u r den-  t i c l e s a n t e r i o r t o i t a r e low, n o d e - l i k e , and c l o s e l y spaced.  The remain-  der o f t h e d e n t i c l e s i n c r e a s e i n h e i g h t and a r e p r o g r e s s i v e l y compressed anteriorly. D e s c r i p t i o n : A. J u v e n i l e - The element i s s m a l l  ( l e n g t h = 350 t o 800 um;  maximum w i d t h = 100 t o 180 um), subsymmetrical, and s l i g h t l y a r c h e d .  The  t h i n p l a t f o r m has i t s l a t e r a l margins upturned more i n t h e c e n t r a l p o r t i o n s of t h e element than a t e i t h e r end.  The p o s i t i o n o f t h e maximum width,;:thoug  v a r i a b l e , i s g e n e r a l l y s l i g h t l y a n t e r i o r of t h e . p o s t e r i o r t i p . The l a t e r a l margins a r e s u b p a r a l l e l t o o n l y s l i g h t l y t a p e r i n g a n t e r i o r l y f o r much o f  t h e i r length corresponding ment.  The  t e r n and  remaining  1/3  to the p o s i t i o n of the r e t i c u l a t e d  of the l e n g t h bears l i t t l e or no r e t i c u l a t e  t a p e r s much more r a p i d l y than the p o s t e r i o r 2/3.  ornament i s r e s t r i c t e d  micro-orna-  The  pat-  reticulate  to the o r a l s u r f a c e on the edges of the p l a t f o r m  i s absent on both the c a r i n a and  the furrows l a t e r a l to i t . The  carina  g e n e r a l l y c o n s i s t s of 6 to 9 l a t e r a l l y compressed t r i a n g u l a r and  nodose  denticles.  The  p o s t e r i o r cusp i s h i g h e r  than a l l o t h e r d e n t i c l e s , c i r c u -  l a r but more commonly s l i g h t l y o v a l i n c r o s s n s e c t i o n , i n c l i n e d but w i t h a s l i g h t , a n t e r i o r l y d i r e c t e d c u r v a t u r e , and h i n d the p o s t e r i o r margin of the p l a t f o r m . p o s t e r i o r f r e e blade  and  The  i n the e a r l i e s t r e c o g n i z e d  posteriorly  s i t u a t e d v a r i a b l y be-  cusp thus extends as a stages but  this feature i s  q u i c k l y l o s t as the p l a t f o r m extends to the p o s t e r i o r edge of the cusp f i n a l l y forms a brim p o s t e r i o r to t h e cusp i n l a t e r o n t o g e n e t i c  and  stages.  The next t h r e e or f o u r d e n t i c l e s a n t e r i o r to the cusp a r e s m a l l and  slight-  l y compressed but a r e more g e n e r a l l y nodose compared to the r e s t of  the  carina.  T h i s f e a t u r e remains i n a l l ontogenetic, stages and  be of major g e n e t i c s i g n i f i c a n c e and  important  maining d e n t i c l e s i n c r e a s e i n s i z e r a p i d l y and  would seem to  to the d i a g n o s i s .  The  re-  become compressed,  inclined  p o s t e r i o r l y and  t r i a n g u l a r i n o u t l i n e as the a n t e r i o r t i p i s approached.  In the e a r l i e s t  stages  the p l a t f o r m does not r e a c h the a n t e r i o r p o r t i o n  where the c a r i n a extends as a f r e e b l a d e , but to encompass t h i s f r e e b l a d e .  The  i n l a t e r stages  it  a b o r a l s u r f a c e bears a narrow and  h i g h k e e l t e r m i n a t i n g p o s t e r i o r l y as an e l e v a t e d o v a l l o o p .  The  groove i s v e r y narrow, extends the e n t i r e l e n g t h of. the k e e l and p o s t e r i o r l y as an elongated, about 3/4 oral  narrow and.curved p i t .  of the p l a t f o r m w i d t h , and  surface.  lengthens  The  very  basal terminates  crimp i s v e r y wide,  smooth In c o n t r a s t to the ornamented  B.  Intermediate  - As the element i n c r e a s e s i n l e n g t h the p l a t f o r m becomes  t h i c k e r and w i d e r . donts  s  As i s the r u l e r a t h e r than the e x c e p t i o n f o r these  the v a l u e s and  variable.  r a t i o s f o r the.  measured parameters a r e  T h i s v a r i a b i l i t y i s e s p e c i a l l y l a r g e f o r L l / H l and  In g e n e r a l , however, the l e n g t h of the i n t e r m e d i a t e 800  and  1100  um and  the w i d t h between 180  imum w i d t h i s g e n e r a l l y 1/3 The  to 1/4  and  220  extremely Ll/Wl  ratios.  element ranges between  um.  The p o s i t i o n of max-  the l e n g t h from the cusp to t h e a n t e r i o r .  p l a t f o r m extends from the p o s t e r i o r edge of the cusp to. the a n t e r i o r  t i p : no  f r e e blades  ties start  e x i s t at t h i s stage.  I t i s at t h i s stage t h a t v a r i e -  to become apparent but. . t h e i r d i f f e r e n t i a t i o n becomes even  more c l e a r i n mature forms where i t w i l l be d i s c u s s e d , i n d e t a i l .  Generally,  t a p e r of the p l a t f o r m i s g e n t l e towards the a n t e r i o r f o r much of the but  cono-  i n c r e a s e s f o r the a n t e r i o r 1/3  to the a n t e r i o r t i p but the platform.  The  is. s t i l l  to 1/4.  The  restricted  r e t i c u l a t e p a t t e r n extends  to the t h i c k e n e d margins of  c a r i n a c o n s i s t s of 10 to 11. l a t e r a l l y compressed,  g u l a r , nodose d e n t i c l e s .  The next 3 to 5 d e n t i c l e s  a n t e r i o r to t h e cusp ( i n most specimens i t i s 4 d e n t i c l e s ) a r e low, i n c r o s s s e c t i o n and n o d e - l i k e .  i n h e i g h t and  The  remaining  a r e p r o g r e s s i v e l y compressed a n t e r i o r l y .  e n t i r e l y d i s c r e t e at t h i s s t a g e . . The  roughly  denticles increase A l l d e n t i c l e s are  a b o r a l s u r f a c e bears a wider and  er k e e l (as compared, to j u v e n i l e stages)  which t e r m i n a t e s  an .elevated o v a l to s l i g h t l y square shaped l o o p .  The  to.3/4 of .the p l a t f o r m w i d t h , and  posteriorly  lowas  b a s a l groove i s s i m i -  l a r to t h a t i n j u v e n i l e s t a g e s , but a l i t t l e less, narrow. narrower, 2/3  trian-  The;posterior,.cusp.(is s i m i l a r to t h a t d e s c r i b e d  f o r the j u v e n i l e . s t a g e except t h a t i t i s l a r g e r .  circular  length  The  crimp i s  smooth.  C. Mature - The v a r i a b i l i t y of form f o r these conodonts i s v e r y h i g h , as mentioned above, and  becomes a c c e n t u a t e d  i n the mature s t a g e s .  It i s at  was  78 t h e s e stages t h a t d i f f e r e n t v a r i e t i e s can. be d i s t i n g u i s h e d . v a r i e t i e s a r e so d i s t i n c t i v e . t h a t , c r i b e d as d i f f e r e n t  Some of  the  i f found a l o n e , they might w e l l be  s p e c i e s j however, i t seems t h a t the v a r i a b i l i t y  t u a l l y that w i t h i n a s i n g l e subspecies.  The g r a p h i c a l evidence  i s ac-  (Figs.  14) f a i l s i n a l l cases t o i s o l a t e these, v a r i e t i e s from each o t h e r . g r a p h i c a l p l o t s t h a t t o a c e r t a i n degree s e p a r a t e the two  des-  10-  Those  extreme v a r i e t i e s ,  namely the p o s t e r i o r a r e a v e r s u s l e n g t h / d e n t i c l e number r a t i o and  the p o s t -  e r i o r a r e a v e r s u s l e n g t h ( F i g s . 12-14), do so w i t h some o v e r l a p between themselves  and a l a r g e number of i n t e r m e d i a t e forms.  I t seems apparent  that  such a f e a t u r e i s the r e s u l t of the normal d i s t r i b u t i o n of v a r i a b i l i t y i n a s i n g l e gene p o o l .  An anisometr.ic type of growth, i n d i c a t e d by the  graph  of a r e a v e r s u s l e n g t h ( F i g . 12), demonstrates q u a n t i t a t i v e l y t h a t shape of the element changes d u r i n g ontogeny: a f a c t a l l u d e d to throughout  the  des-  c r i p t i o n of t h i s s p e c i e s . The two v e r y d i f f e r e n t v a r i e t i e s p r e s e n t f o r t h i s s u b s p e c i e s w i l l d e s c r i b e d as the g r a c i l i s v a r i e t y and forms t h a t cannot  the robustus v a r i e t y .  q u a l i t a t i v e l y be separated  v a r i e t y , so-named because of i t s v e r y l o n g and s i m i l a r t o the s p e c i e s N e o g o n d o l e l l a The v a r i e t y r o b u s t u s resembles  A number of  i n t o one o r another  v a r i e t i e s w i l l be r e f e r r e d to as the i n t e r m e d i a t u s v a r i e t y .  be  The  of  these  gracilis  slender platform, i s very  g r a c i l i s C l a r k and  Ethington,  1962.  the s p e c i e s N e o g o n d o l e l l a s e r r a t a C l a r k and  E t h i n g t o n , 1962.except f o r the much, l a r g e r l e n g t h per d e n t i c l e number and the l a c k of r i d g e s a s s o c i a t e d w i t h the s e r r a t e margin. v a r i e t y most resembles Hawley and M i l l e r ,  1951  the s p e c i e s N e o g o n d o l e l l a l e a d i n g to the new  The  idahoensis  intermediatus Youngquist,  s u b s p e c i e s b e i n g r e f e r r e d to N.  i d a h o e n s i s r a t h e r than N. g r a c i l i s or N. s e r r a t a . r e f e r r i n g the p o p u l a t i o n s i n q u e s t i o n to a new  I t was  considered that  s p e c i f i c rank r e q u i r e d unsub-  •g  •r  •r  •r.  g  N. idahoensis n.subsp.  i  400  j 600  •  one  specimen  •  two  specimens  i  i  i  '  800  1000  1200  1400  Length  F i g u r e 10.  A(F49)  -  L1  Graph showing t h e r e l a t i o n s h i p o f Length/Height t o Length o f t h e p l a t f o r m f o r F49. r= v a r . r o b u s t u s g= v a r . g r a c i l i s , t h e remainder a r e v a r . i n t e r m e d i a t u s  +r  N. idahoensis n.subsp. A(F49) • one specimen • two specimens 400  600  800  1000  -L  1200  1400  Length-L1  F i g u r e 11.  Graph showing the r e l a t i o n s h i p of Length/Width to Length of t h e p l a t f o r m f o r F49. r= v a r . r o b u s t u s g= v a r . g r a c i l i s , the remainder a r e v a r . i n t e r m e d i a t u s .  81 14  13r  12  11  10k  <  LU CC <  /  tr o E  LU H CO  /  2*  i  v / /a  A *  1  '  /  /  /  3  4  -I  5  L.  6  7  8  9  LENGTH x * f  Figure  -1  10  11  l_  12  13  14  15  16  2  12. -.Graph showing the r e l a t i o n s h i p of P o s t e r i o r Area to Length of the p l a t f o r m f o r F53. r=~var. r o b u s t u s g= v a r . g r a c i l i s , the remainder a r e v a r . i n t e r m e d i a t u s  82  / / / I  1 I I i  i  I  I  I  I I  -r  /  /•  /  •  /  /  /  •/. .  /  /  / /  /  '  ''  /  :9 / /  . *.-r  ... /  •  / -I  2  3  ••  / »/.g  -g  -g  y  /•  /  9 /  '  /  /  a  /  / 1  1  1  1  1  1  4  5  6  7  8  . 9  1  10  I  11  12  13  14  15  16  LENGTH x 10"  2  F i g u r e 13.  Graph showing the r e l a t i o n s h i p of P o s t e r i o r Area to Length of the p l a t f o r m f o r F49. r= v a r . r o b u s t u s g= v a r . g r a c i l i s , the remainder a r e v a r . i n t e r m e d i a t u s .  tg-  a  •g  • N. idahoensis n.subsp. A(F49) — i  0  60  i  i  I  i  '  70  80  90  100  110  Length/number  F i g u r e 14.  of d e n t i c l e s - L 1 / *  Graph showing the r e l a t i o n s h i p o f P o s t e r i o r Area to Length/number of d e n t i c l e s f o r F49. r= v a r . ;robustus g= v a r . g r a c i l i s , the remainder a r e v a r . i n t e r m e d i a t u s .  84 s t a n t i a t e d taxonomic. s p l i t t i n g .  I f q u a n t i t a t i v e methods i n d i c a t e d a  separ-  a t i o n of t h i s form from o l d e r N.. i d a h o e n s i s . s i m i l a r , to the s e p a r a t i o n f o r _N. s e r r a t a and t a as N.  N. p o s t s e r r a t a (Behnken,.1975 - he r e f e r r e d to N.  s e r r a t a p o s t s e r r a t a which was  rank) then perhaps t h i s new rank.  subspecies  subsequently  separated  l a t e s c o u l d , f o l l o w i n g s p e c i a t i o n and m i g r a t i o n ,  l e a d to p o p u l a t i o n s more  " d e s c r i b e d as _N. g r a c i l i s . o r _N. s e r r a t a .  or. u n u s u a l forms, a r e mentioned below.  l a r l y advantageous; a t l e a s t one, t h a t of the T r i a s s i c N.  Wl  = 160  s p e c i e s N.  as mutations, t h a t were not p a r t i c u -  however, has a p l a t f o r m shape s i m i l a r  i d a h o e n s i s n.subsp. A'Var.  ym),  subsymmetrical, and  tends the f u l l l e n g t h of the element and of t h e cusp.  The  The  slender  ( L l = 8 5 0 to 1200  gently arched.  The  ym;  platform  ex-  i s f l u s h w i t h the p o s t e r i o r edge especially  l a t e r a l margins of t h e p o s t e r i o r h a l f o f  p l a t f o r m a r e s u b p a r a l l e l to p a r a l l e l and The  gracilis  l a t e r a l edges of the p l a t f o r m a r e upturned,  i n the c e n t r a l r e g i o n s .  to  constricta.  p l a t f o r m s a r e t y p i c a l l y l o n g and to 220  "experiments"  Perhaps, as t h e s e u n u s u a l forms a r e  i n number, they c o u l d be c o n s i d e r e d  The  specific  as.peripheral iso-  As w e l l as these t h r e e named morphologies a. number o f  few  specific  c o u l d be l a t e r e l e v a t e d to  I t i s p o s s i b l e t h a t these v a r i e t i e s . i f  reliably  e l e v a t e d to  postserra-  the  t a p e r g e n t l y i n the a n t e r i o r h a l f .  p o s t e r i o r margin i s g e n t l y rounded and w i t h the r e t i c u l a t e d .portion  t a p e r i n g to the cusp.  The  r e t i c u l a t i o n ornament t a p e r s on the  margins i n the same f a s h i o n as the p l a t f o r m i t s e l f the most a n t e r i o r p o r t i o n s . d e n t i c l e s and couple  furrows.  The  t a p e r s but  lateral i s absent i n  r e t i c u l a t e p a t t e r n i s absent on b o t h  the  Some f a i n t r i d g e s were observed on the cusp o f a  of specimens - these may  represent  the v e r y e a r l y f o r m a t i o n o f  reti-  85 culae.  The  c a r i n a t y p i c a l l y comprises 12 or 1 3 . d i s c r e t e  a n t e r i o r , f o u r of which a r e low a noticeable on  and  c r e a s e or m i d - l i n e .  denticles,  n o d i f o r m , a l l of which a r e connected No  by  a n t e r i o r s e r r a t i o n s have been noted  this variety. N. As  i d a h o e n s i s n.subsp. A v a r .  robustus  the name i n d i c a t e s t h i s v a r i e t y i s t h i c k e r , wider and  more s t o u t  or r o b u s t compared to v a r .  is generally  between 800  and  1250  um  gracilis. and  The  the w i d t h between 200  T h i s r e s u l t s i n p o s t e r i o r a r e a s a v e r a g i n g 6.2 a b l y more t h a n the 4.4 extends the f u l l  u n i t average for. the  l e n g t h o f the  p o s t e r i o r brim but  and  (um  X 10 )',  gracilis variety.  and  consider-  The  The  p a r a l l e l s the p l a t f o r m  margin, and  The  half:  p o s t e r i o r margins The  of gerontic  The  distinct  furrows (PI. 5,  anterior  r e t i c u l a t e pattern  the d e n t i c l e t i p s i n many specimens.  individuals.  The  ridges  on the o u t e r edge, and  f i g . 1). . D e n t i c l e s  f a d i n g and  (generally  s e r r a t i o n s a r e v a r i a b l y developed but  the  i t s range,  e l o n g a t e towards  11 to 13)except f o r  some f u s i o n of the p o s t e r i o r . o n e s , c o n f o r m to e a r l i e r d e s c r i p t i o n s . o f specimens e x h i b i t a s e r r a t e p l a t f o r m  This  feature  that c o n s t i t u t e  meshes of the r e t i c u l a t i o n are t y p i c a l l y sharp i n the m i d d l e of f l a t t e n e d but  reticu^  i s m i s s i n g on the  seems to be r e l a t e d to e a r l y f u s i o n of the d e n t i c l e s - a  representative  platform  increase  f i n a l l y t a p e r over the a n t e r i o r  more r a p i d l y c l o s e to the t i p .  m i g r a t e s onto the furrows and  m.  u  element, i n some c a s e s forming a minor  -most edges where the margin t a p e r s most r a p i d l y .  feature  270  4  a r e q u i t e square as opposed to the rounded g r a c i l i s v a r i e t y . lated pattern  and  a r e s u b p a r a l l e l f o r the p o s t e r i o r 3/10,  i n w i d t h over the next 2/10, at f i r s t  units  element  n o r m a l l y f l u s h w i t h the r e a r edge of the cusp.  margins of the p l a t f o r m  slowly  generally  l e n g t h of the  2  the  the  margin i n the a n t e r i o r  1/3.  g e n e r a l l y weaker than those  A number The described  86 by.Clark  and Behnken (197.9) f o r N. s e r r a t a and younger Permian conodonts  and  without  The  k e e l i s wide and .low and t e r m i n a t e s  The  b a s a l groove i s wider than i n more j u v e n i l e forms and t e r m i n a t e s  basal p i t .  t h e r i d g e s accompanying t h e s e r r a t i o n s as. i n N. s e r r a t a . i n an e x t e n s i v e t r i a n g u l a r l o o p .  The smooth crimp t y p i c a l l y c o v e r s  i n the  6/10 t o 2/3 o f t h e a b o r a l  surface. !N. i d a h o e n s i s n.subsp. A v a r . The  intermediatus  form o f t h e c a r i n a , p o s i t i o n o f t h e r e t i c u l a e , and development  of t h e a b o r a l s u r f a c e a r e s i m i l a r t o those d e s c r i b e d f o r t h e o t h e r v a r i e d , ties.  The p l a t f o r m shape i s i n t e r m e d i a t e between t h a t f o r t h e g r a c i l i s and  robustus  v a r i e t i e s w i t h t h e maximum w i d t h g e n e r a l l y c o r r e s p o n d i n g  p o s i t i o n o f t h e f o u r t h d e n t i c l e a n t e r i o r t o t h e cusp. i o r r e g i o n average about 5.0 u n i t s .  Areas i n t h e p o s t e r -  The p l a t f o r m t a p e r s more or l e s s  d u a l l y from t h e maximum w i d t h t o t h e a n t e r i o r t i p . present  to the  gra-  Anterior serrations are  but l e s s .common ' than i n t h e r o b u s t u s v a r i e t y .  The main . d i f f e r e n c e s i n p l a t f o r m shape a r e t h e average maximum w i d t h , and  t h e manner, p o s i t i o n , and degree o f t a p e r .  described  These f e a t u r e s have been  i n d e t a i l f o r t h e extreme v a r i e t i e s , whereas i t seems adequate  to s t a t e t h a t t h e r e p r e s e n t a t i v e s o f v a r . i n t e r m e d i a t u s p a r t o f a continuous  form t h e c e n t r a l  g r a d a t i o n and t y p i f y t h e p o p u l a t i o n means.  I n t r o d u c t i o n t o t h e u n u s u a l v a r i e t i e s o f N. i d a h o e n s i s n.subsp. A Those forms t h a t a r e u n u s u a l in.terms generally recognized  o f t h e i r p l a t f o r m shape a r e  a t t h e mature t o g e r o n t i c stages  do n o t show o b s e r v a b l e  ontogeny.  These forms comprise  of development and only a small  frac-  t i o n o f any p o p u l a t i o m : sample. N. i d a h o e n s i s n.subsp. A v a r . c o n s t r i c t u s T h i s v a r i e t y has a c o n s t r i c t e d p l a t f o r m margin i n t h e p o s t e r i o r o f t h e  element.  The  platform  i s rounded i n the p o s t e r i o r and  a n t e r i o r l y f o r about 1/4 widest p o i n t about 3/8  has  fused  of i t l e n g t h . The  From t h i s p o i n t the p l a t f o r m  illustrated  p o s t e r i o r d e n t i c l e s ( P i . 4,  a l s o been r e c o g n i z e d  slightly  of i t s l e n g t h where i t widens r a p i d l y to i t s  n o r m a l l y f o r the s u b s p e c i e s . and  tapers  specimen i s a g e r o n t i c  f i g . 7).  Var.  form  constrictus  i n specimens w i t h a l l d e n t i c l e s d i s c r e t e .  form shape i s c h a r a c t e r i s t i c of N.  tapers  has  This p l a t -  c o n s t r i c t a from the T r i a s s i c  i n terms of  the c o n s t r i c t e d p o s t e r i o r margin. N. T h i s form has except.;for  idahoensis  a v e r y wide and  the p o s t e r i o r end,  of the p l a t f o r m  N. The  rosenkrantzi.  lobe  the  The.furrows are wider-and s l i g h t l y deeper than t h o s e i n . s i m i l a r -  :  :  ate  node-like  as  i n height  a n t e r i o r l y and  i s g e n e r a l l y c h a r a c t e r i s t i c f o r the  cusp i s a l s o more compressed then normal and  but  r a t h e r s t r a i g h t upwards.  Discussion:  J u v e n i l e forms of N.  time by S z a n i a w s k i and of S p i t s b e r g e n . to t h a t f o r N.  The  S z a n i a w s k i and  idahoensis  not  a r e not  idahoensis  subspecies.  directed posteriorly  were d e s c r i b e d  f o r the  first  n.subsp. A,  by these a u t h o r s i s v e r y s i m i l a r  except t h a t the j u v e n i l e c a r i n a  has  than 8 to 10 f o r the Kapp S t a r o s t i n specimens.  Malkowski (1979) s t a t e d t h a t the ontogeny f o r N.  s i m i l a r to t h a t of N.  Behnken (1971) and  dis-  Malkowski (1979) from the Kapp S t a r o s t i n Formation  ontogeny d e s c r i b e d  6 to 9 d e n t i c l e s r a t h e r  welli•  n o t - d i r e c t e d p o s f e e r o - l a t e r a l l y as  d e n t i c l e s increase only s l i g h t l y  very  p o s t e r i o r margin  s u r r o u n d i n g the cusp and"separated  The  was  The  l o b e s forming the p o s t e r o - l a t e r a l margins of  r o s e n k r a n t z i n.subsp. B and  t i n c t and  lobatus  t h i c k platform o v e r a l l that i s s i m i l a r ,  to N.  i s t r i - l o b e d : one  by furrows from the two element.  n.subsp. A v a r .  bitteri  (Kozur) as d e s c r i b e d  concluded t h a t these two  by  idahoensis Clark  s p e c i e s bore a c l o s e  and  relation-  88 The j u v e n i l e forms, of N . i d a h o e n s i s n.subsp. A a r e v e r y s i m i l a r to  ship.  those of N . r o s e n k r a n t z i n.subsp. D...., Furthermore j u v e n i l e forms of N . p o s t s e r r a t a as i l l u s t r a t e d  i n Behnken (1975) a r e a l s o v e r y s i m i l a r to those o f  the o t h e r s p e c i e s j u s t mentioned.  There seems to be good r e a s o n to b e l i e v e re.-,  t h a t the c l o s e s i m i l a r i t y o f the j u v e n i l e forms d e s c r i b e d above i s the s u i t of c l o s e phylogenetic a f f i n i t y f o r a l l s e r r a t a complex sp. A.  ( C l a r k and Behnken,  those s p e c i e s b e l o n g i n g to the  1979) and i n c l u d i n g N . i d a h o e n s i s n.sub-  C l e a r l y , s p e c i e s cannot be determined on the b a s i s of j u v e n i l e  ma^  The d i f f e r e n c e s between members of N . i d a h o e n s i s n.subsp. A  t e r i a l alone.  from F49 t o F54 a r e d i r e c t l y p r o p o r t i o n a l to the number of d e n t i c l e s  on  the element which i s more or l e s s d i r e c t l y r e l a t e d to s i z e and o n t o g e n e t i c development and i l l u s t r a t e s t h e need f o r i n t e r m e d i a t e and mature forms bef o r e d i f f e r e n t i a t i o n o f s u b s p e c i e s and s p e c i e s can be attempted. Mature specimens d i f f e r l i t t l e N.  idahoensis.  from specimens p r e v i o u s l y r e f e r r e d t o  Most workers a t t e s t to a h i g h v a r i a b i l i t y  i n shape and i n  l e n g t h to w i d t h r a t i o a l t h o u g h none have attempted to q u a n t i f y t h i s . bility  Varia-  i s a c h a r a c t e r i s t i c of N . i d a h o e n s i s n.subsp. A ( F i g s . 10, 11).  Perhaps the most c l o s e l y comparable specimens t h a t I have seen a r e t h o s e .'. i l l u s t r a t e d and d e s c r i b e d by S z a n i a w s k i and Malkowski (1979) from S p i t s b e r gen. N.  As w e l l as _N. i d a h o e n s i s (137 fragments) t h e s e a u t h o r s d e s c r i b e d _ N . c f .  gracilis  (3 fragments) and IT. sp. A (1 specimen).  I n - t h e l i g h t of my  faunas and t h e i r v a r i a b i l i t y . a l l of t h e s e fragments would be i n c l u d e d i n a single species.  The N . c f . N . g r a c i l i s  i s s i m i l a r to t h e E l l e s m e r e I s l a n d  v a r . g r a c i l i s whereas t h e N . sp. A i s s i m i l a r to V a r . l o b a t u s . imens d i f f e r  from H.  These s p e c -  i d a h o e n s i s n.subsp. A i n terms of t h e l a c k of a n t e r i o r  s e r r a t i o n s , t h e number of d e n t i c l e s  (8 to 13 d e n t i c l e s compared  and i n t h e manner of t a p e r i n g (widest p o i n t c l o s e r t o p o s t e r i o r  to 6 t o 16), tip).  89 Their  specimens appear c l o s e r to t h e type specimens f o r the s p e c i e s  are probably  s l i g h t l y o l d e r than N.  C l a r k and and N.  and  i d a h o e n s i s n.subsp. A.  Mosher (1966)' regarded  N. p h o s p h o r i e n s i s Youngquist, Hawley  M i l l e r w i t h i t s p o s t e r i o r r i d g e - l i k e c a r i n a , as a l a t e r growth stage of i d a h o e n s i s where the p o s t e r i o r c a r i n a has become f u s e d .  mere c o l l e c t i o n s t h a t form i s r e s t r i c t e d to mature and  In the  Elles-  g e r o n t i c forms o n l y .  As w e l l , the f u s i o n of t h e c a r i n a i s shown to be g r a d a t i o n a l ( P l . 4, 5, 6,  10).  These o b s e r v a t i o n s most c e r t a i n l y l e n d f u r t h e r support  and Mosher's c o n c l u s i o n t h a t N. p h o s p h o r i e n s i s N.  figs.  to C l a r k  be p l a c e d i n synonymy w i t h  idahoensis. The b a s a l l o o p shows a development from a h i g h e l o n g a t e o v a l to rounded  shape i n j u v e n i l e and i n mature and  i n t e r m e d i a t e forms to a low,  g e r o n t i c forms.  specimens ( P l . 5, f i g s .  9,  10)  Observation  l a r g e , t r i a n g u l a r shape  of the growth l a m e l l a e i n a  few  shows t h a t some of the l a t e r a l ' t r a c e s of  l a m e l l a e a r e t r u n c a t e d at the p o s t e r i o r of the l o o p .  the  The ^ t r u n c a t i o n i s  a p p a r e n t l y caused by r e s o r p t i o n as d e s c r i b e d by M u l l e r and Nogami (1972) which, a c c o r d i n g to these a u t h o r s , iformes.  i s a common phenomenon'for the Conodont-  I t r e s u l t e d i n s q u a r i n g - ' o f f the p o s t e r i o r of the l o o p such t h a t  subsequent r e g e n e r a t i o n of the l o o p l e d to a t r i a n g u l a r shape. then,  seems to be an important  odont. having  M u l l e r and  phenomenon w i t h i n the ontogeny of t h i s  Nogami (1972) conclude  the f u n c t i o n to support  Resorption,  t h a t the conodont element,  a t i s s u e , may  conbesides  a l s o have served as an organ  f o r the temporary d e p o s i t i o n of p h o s p h a t i c , s u b s t a n c e ,  which might l a t e r  utilized  One  to form another  element i n the same a n i m a l .  s p e c u l a t e whether t h i s r e s o r p t i o n phenomenon observed ment of N e o g o n d o l e l l a  also occurred  cannot h e l p  i n the p l a t f o r m  i n the ramiforms ( i f a  be  but ele-  multielement  90 apparatus indeed  e x i s t e d ) r e s u l t i n g i n a p l a t f o r m - o n l y apparatus (as sug-  gested by M e r r i l l and P o w e l l ,  1980): the excess  form t h e l a r g e , t h i c k - m a r g i n e d  phosphate b e i n g used t o  mature and ger.ontic p l a t f o r m elements.  NEOGONDOLELLA SERRATA(?) ( C l a r k and E t h i n g t o n , 1962) P I . 6, f i g s . 7-9. G o n d o l e l l a s e r r a t a C l a r k and E t h i n g t o n Gondolella nankingensis Occurrence:  Upper A s s i s t a n c e Formation,  Ching, 1960  Hamilton P e n i n s u l a s e c t i o n (F63,  F73) . Description:  T h i s d e s i g n a t i o n i s based only on fragmental  specimens.  Es-  t i m a t e s o f l e n g t h were made by t a k i n g i n t o c o n s i d e r a t i o n the g e n t l e t a p e r . Specimens t h a t would be regarded  as i n t e r m e d i a t e  i n ontogeny have an e s t i -  mated l e n g t h o f 400 t o 580 ym and a maximum w i d t h o f 100 ym. a n t e r i o r d e n t i c l e s were n o t observed,  t h e t o t a l number o f d e n t i c l e s f o r  these i n t e r m e d i a t e forms i s on t h e o r d e r o f 11. to t h e cusp a r e s h a r p distinct  s  Although t h e  The f o u r d e n t i c l e s a n t e r i o r  f u s e d a t t h e i r bases, l a t e r a l l y compressed and not  from t h e o t h e r d e n t i c l e s as i n N. i d a h o e n s i s n.subsp. A.  i s high'and l a t e r a l l y compressed, e s p e c i a l l y t h e a n t e r i o r h a l f . form i s g e n t l y t a p e r i n g , l a t e r a l l y upturned, arched e r i o r end. and  The p l a t -  and rounded a t t h e p o s t -  The lower s u r f a c e o f t h e p l a t f o r m i s smooth and bears a h i g h  r e l a t i v e l y narrow k e e l .  sharp,  The cusp  The r e t i c u l a t e " p a t t e r n on t h e upper s u r f a c e i s  i r r e g u l a r i n shape and r e s t r i c t e d t o t h e l a t e r a l margins as i t fades  q u i c k l y towards t h e c a r i n a , r e a c h i n g t h e l a t t e r o n l y a t t h e p o s i t i o n o f t h e cusp. Discussion:  These specimens d i f f e r from t h e o l d e r specimens o f N.  idahoen-  s i s n.subsp. A i n t h e i r s m a l l e r s i z e and they do n o t have the f o u r  nodiform  denticles .distinct  from t h e o t h e r s a n t e r i o r t o t h e cusp.  Reduction  i n size  91  a t comparable growth stages has been o b s e r v e d . i n the e v o l u t i o n o f N. h o e n s i s to N'. s e r r a t a i n o t h e r r e g i o n s . r a t a that i s present mediate  Another c h a r a c t e r i s t i c of N.  ser-  i n these specimens i s the f u s i o n of d e n t i c l e s i n i n t e r -  r a t h e r than g e r o n t i c stages of growth.  of the p l a t f o r m were not observed  S i n c e a n t e r i o r fragments  i t i s i m p o s s i b l e to a s s e s s whether the  a n t e r i o r s e r r a t i o n s , d i a g n o s t i c of the s p e c i e s and p r o v i d i n g the of the name, a r e  ida-  derivation  present.  A c c o r d i n g to the f i g u r e s mentioned above, the v a l u e s f o r L l and  Wl  i n d i v i d u a l l y average about two  standard d e v i a t i o n s s m a l l e r than the mean  f o r N.  The r a t i o Ll/number of d e n t i c l e s i s more  i d a h o e n s i s n.subsp. A.-  than two  standard d e v i a t i o n s l e s s than the means.for N.  A and almost subsp. _D.  two  i d a h o e n s i s n.subsp.  standard d e v i a t i o n s l e s s than t h a t f o r N. r o s e n k r a n t z i n.  D e s p i t e the f a c t t h a t these f i g u r e s a r e based on o n l y v e r y  few  specimens t h e i r s i g n i f i c a n t d e p a r t u r e from the means f o r specimens lower i n the s e c t i o n suggest  t h a t they a r e indeed a d i f f e r e n t  species.  Furthermore,  s i n c e they share some of the f e a t u r e s of N. s e r r a t a they a r e a s s i g n e d to ; t h a t taxon but a r e more p r o b a b l y A and  N.  serrata.  The  i n t e r m e d i a t e between N.  specimens a l s o p l o t  i n or near the f i e l d of''data  p o i n t s f o r L l v e r s u s number of d e n t i c l e s of N. ken  (1975).  The  p r e s e r v a t i o n and The  i d a h o e n s i s n.subsp.  s e r r a t a as d e f i n e d by Behn-  assignment i s l i s t e d as i n d e f i n i t e because of the poor p a u c i t y of specimens.  i r r e g u l a r r e t i c u l a t e p a t t e r n on the upper p l a t f o r m s u r f a c e a l s o  seems worthy of f u r t h e r d i s c u s s i o n .  Behnken (1975, P l . 2, f i g s . 35,  36)  i l l u s t r a t e s the r e t i c u l a t e m i c r o s t r u c t u r e of an i n t e r m e d i a t e and a mature N. p o s t s e r r a t a .  The m i c r o s t r u c t u r e i s v e r y r e g u l a r l y shaped and  i n l i n e a r rows (ordered)  arranged  i n the i n t e r m e d i a t e form whereas m i c r o s t r u c t u r e of  92 the mature form i s i r r e g u l a r and arranged i n a roughly ordered) . served  S i m i l a r ontogenetic  sinuous manner  v a r i a t i o n , a l t h o u g h n o t as marked, was ob-  i n specimens o f N. i d a h o e n s i s n.subsp. A.  i n d i c a t i v e of r e l a t i v e maturity.  Perhaps t h e f e a t u r e i s  T h i s would suggest t h a t t h e s m a l l s p e c i -  mens r e f e r r e d here t o _N. s e r r a t a ( ? ) a r e approaching m a t u r i t y exhibit disordered  (dis-  s i n c e they  reticulation.  A purely q u a l i t a t i v e observation  of the platforms  r e f e r r e d t o N. s e r -  r a t a ( ? ) i s t h e i r g e n e r a l degenerate appearance, l a c k i n g t h e r o b u s t n e s s o f the o l d e r f a u n a s .  There appears t o be e v i d e n c e t h a t t h e l a c k o f conodonts  f o r a s i g n i f i c a n t p a r t o f t h e s e c t i o n above _N. s e r r a t a ( ? ) i s t h e r e s u l t o f a biologic crisis.  Perhaps t h e degenerate appearance o f t h e s e specimens o f  N.. s e r r a t a ( ? ) r e f l e c t barren  the i n i t i a t i o n of t h i s c r i s i s . .  The faunas above t h e  i n t e r v a l a r e e q u a l l y sparse:' a f e a t u r e common t o any c r i s i s o r near  extinction.  N. r o s e n k r a n t z i marks t h e reappearance o f robust  and abundant  specimens. . NEOGONDOLELLA n.sp.  B  PI. 6, f i g s v 1-4. Occurrence: section  Peninsula  (F36, F 8 3 ) .  Diagnosis: very  Lower p a r t of t h e T r o l d F i o r d Formation, H a m i l t o n  This very  symmetrical p l a t f o r m element i s d i s t i n q u i s h e d by i t s  l a r g e cusp which i s round i n c r o s s s e c t i o n and d i r e c t e d s t r a i g h t up-  wards.  Other d i a g n o s t i c f e a t u r e s i n c l u d e a w e l l developed, f a i n t l y  l o b e d brim p o s t e r i o r t o t h e cusp i n mature elements.  tri-  T h i s brim bears a  c o a r s e l y s t r i a t e t o f a i n t l y r e t i c u l a t e ornament on t h e o r a l s u r f a c e .  Fur-  thermore, t h e r e t i c u l a t i o n on t h e p l a t f o r m margin ends v e r y a b r u p t l y a t t h e f u r r o w margin.  93  Description:  T h i s new  s p e c i e s i s based only, on the p o s t e r i o r r e g i o n s of  p l a t f o r m owing to f r a g m e n t a t i o n present  of the elements.  on the p o s t e r i o r h a l f a r e v e r y d i s t i n c t i v e .  form the p l a t f o r m margins a r e s u b p a r a l l e l (how r e g i o n i s unknown) and cusp.  The  In the  the  features  intermediate  they t a p e r . i n the a n t e r i o r  rounded on the p o s t e r i o r end where i t meets the  cusp i s l a r g e , rounded i n c r o s s s e c t i o n , d i r e c t e d almost  upward, and low,  Nevertheless  shows l i t t l e  to no l a t e r a l compression.  l a t e r a l l y compressed, and  are r e t i c u l a t e d  The  f u s e d a t t h e i r bases.  i n a regular fashion.  The  straight  d e n t i c l e s are a l l  The  p l a t f o r m margins  r e t i c u l a t i o n does not r e a c h  rounded p a r t s of the p o s t e r i o r of the p l a t f o r m margin.  The  furrows  d e n t i c l e s are.smooth up t o / t h e p o i n t where the r e t i c u l a t i o n begins  p o s t e r i o r margin i s f a i n t l y t r i - l o b e d o t h e r s can be v e r y narrow.  The  abruptly.  i n o u t l i n e on most specimens but  d i r e c t e d s t r a i g h t upwards.  reaches the p o s t e r i o r margin and to a c o a r s e  This in  cusp i s g e n e r a l l y v e r y l a r g e , c i r c u l a r i n  s i m i l a r to t h a t i n the i n t e r m e d i a t e  way  the  and  In mature forms a p l a t f o r m margin forms p o s t e r i o r to the cusp.  c r o s s s e c t i o n , and  the  The  r e t i c u l a t e ornament i s  form except t h a t the  the cusp.  The  reticulation  r e t i c u l a t i o n generally gives  s t r i a t e d ornament on the brim p o s t e r i o r to the cusp.  The  d e n t i c l e s a r e smooth, l a t e r a l l y compressed (some can be n o d e - l i k e ) ,  and  fused up  bears  a low,  to h a l f of t h e i r h e i g h t .  wide k e e l which t e r m i n a t e s  The  narrow b a s a l groove t e r m i n a t e s  The  crimp o c c u p i e s  Discussion:  about 2/3  The a b o r a l s u r f a c e i s smooth and i n t o an e q u a l l y low,  i n a s l i g h t l y curved,  loop.  elongated.oval  pit  of the a b o r a l w i d t h .  S u p e r f i c i a l l y , the intermediate  h o e n s i s n.subsp. A.  rounded b a s a l  However, N. n.sp.  B_ has  form l o o k s s i m i l a r to N_. a l a r g e r , l e s s compressed  t h a t i s d i r e c t e d upwards u n l i k e t h a t f o r the o l d e r s p e c i e s .  The  idacusp  reticulate  94  ornament fades  towards the furrows ending i n f a i n t , l i n e a r r i d g e s  perpendi-  c u l a r to the l e n g t h i n N_. i d a h o e n s i s n.subsp. A whereas the same ornament ends a b r u p t l y and  l a c k s any  l i n e a r r i d g e s i n N. n.sp.. B_.  Other d i f f e r e n -  t i a t i n g f e a t u r e s a r e the p a r t i a l f u s i o n of d e n t i c l e s .at t h e i r base and l a c k of r e t i c u l a t i o n on the p o s t e r i o r " s h o u l d e r s " of the p l a t f o r m . more, the element of N. n.sp.  B_„is much more symmetrical  s p e c i e s owing to the p o s i t i o n of the cusp and p o s t e r i o r of the p l a t f o r m .  Further-  than i n o l d e r  the p a r a l l e l margin i n the  These forms l o o k v e r y s i m i l a r to a younger  s p e c i e s i d e n t i f i e d by C l a r k and Behnken (1979) as N. w i l c o x i a l t h o u g h cusp i n N. n.sp.  of the w e l l developed p o s t e r i o r brim striations".  i d a h o e n s i s n.subsp. A i n terms  surrounding  T h i s p o s t e r i o r brim  i s present  i n younger s p e c i e s  and  2,  21), however, the ornament i s r e t i c u l a t e d r a t h e r than of  14,  striations. more  N. w i l c o x i i n p a r t i c u l a r ;  the cusp t h a t i s ornamented  (N_. b a b c o c k i figs.  the  B_ i s g e n e r a l l y l a r g e r .  Mature forms a r e d i s t i n g u i s h e d from N.  with coarse  the  N. n.sp.  see C l a r k and  Behnken, 1979,  PI.  coarse  B_ d i f f e r s from N. b i t t e r i n.subsp. C_ i n b e i n g much  symmetrical. The  a b o r a l s u r f a c e i s s i m i l a r to t h a t i n younger and  The measurements f o r L2 which range between 240 t h a t f o r N.  and  310  older species.  um a r e s i m i l a r  to  i d a h o e n s i s n.subsp. A whereas the measurements for.maximum w i d t h  which range between 140 and  280  um,  tend to be a l i t t l e wider on average.  NEOGONDOLELLA POSTSERRATA(.?) (Behnken,.. 1975) PI.'6, f i g s . 5, 6. Neogondolella Occurrence : Description:  s e r r a t a p o s t s e r r a t a Behnken,  1975  M i d d l e T r o l d F i o r d Formation, Hamilton P e n i n s u l a T h i s i d e n t i f i c a t i o n i s based on o n l y a v e r y few  t h a t a r e d i s t i n c t l y u n l i k e any  others  section  (F87).  fragments  seen through the e n t i r e sequence.  95  The  gently  arched t o almost f l a t p l a t f o r m  margins and an a b r u p t l y  possesses . s u b p a r a l l e l  squared-off p o s t e r i o r .  form a r e l a r g e , c i r c u l a r i n c r o s s  The d e n t i c l e s i n t h e mature  s e c t i o n and d i s t i n c t .  I n a s m a l l mid-  platform  fragment, t h e d e n t i c l e s a r e l a t e r a l l y compressed and almost e n t i r e -  l y fused  s u g g e s t i n g t h i s may be a g e r o n t i c  rectangular  shaped node p r o j e c t i n g  form.  The cusp i s a c t u a l l y a  to one s i d e of t h e p l a t f o r m .  A small  r i d g e marks t h e p o s t e r i o r border on t h e s i d e o p p o s i t e t o t h e cusp.  A  f a i r l y r e g u l a r l y arranged r e t i c u l a t e ornament on t h e margins of t h e p l a t form reaches t h e p o s t e r i o r margin and ends a b r u p t l y e r a l to the c a r i n a begin. be  Very f a i n t r i d g e s  where t h e furrows  perpendicular  lat-  t o t h e l e n g t h can  seen on t h e o t h e r w i s e smooth furrows o f t h e m i d - p l a t f o r m fragment but '.:  t h e r e a r e no s e r r a t i o n s on t h e p l a t f o r m  margins.  Measurements f o r Wl (mean = 200 um) and L2 (mean = 260 ym) a r e v e r y s i m i l a r t o those f o r N. i d a h o e n s i s n.subsp. A. Discussion;  These specimens a r e r e f e r r e d t o N_. p o s t s e r r a t a ( ? )  because o f  t h e i r s t r a t i g r a p h i c p o s i t i o n , t h e i r uniqueness compared t o o t h e r  species  i n t h e same s e c t i o n and because o f t h e i r s i m i l a r i t y t o some o f t h e squareended specimens f i g u r e d by C l a r k and Behnken (1979, P l . 1, f i g . 17). The  shape o f t h e cusp and t h e squared p o s t e r i o r end a r e the main d i s -  tinguishing features. can  However, Behnken (1975) i n d i c a t e s t h a t N.  have both rounded and squared p o s t e r i o r margins, so t h a t  distinguishing features  should n o t be c o n s i d e r e d e x c l u s i v e  postserrata  t h e above  to the species.  NEOGONDOLELLA BITTERI n.subsp. _C P l . 7, f i g s . Occurrence: Diagnosis:  1-8.  Upper T r o l d F i o r d Formation, Hamilton P e n i n s u l a s e c t i o n A species  cusp o f c i r c u l a r c r o s s  characterized  by a t h i c k p l a t f o r m  (F96).  w i t h a low, wide  s e c t i o n surrounded by a brim w i t h r e t i c u l a t e " o r n a -  96  merit arid low  d e n t i c l e s on the c a r i n a .  The  subspecies i s characterized  a d i s t i n c t asymmetry r e s u l t i n g from the o f f - c e n t r e p o s i t i o n of the posterior  This designation  i s based on a number of p o s t e r i o r end  ments which have v e r y d i s t i n c t i v e  features.  i s lobed  and  d i s t i n c t l y asymmetrical.  extends beyond the cusp as a well, developed brim. i s formed by a l a r g e l o b e o c c u p y i n g one The  cusp i s p o s i t i o n e d  the. other.:as w e l l .  The  low,  the c a r i n a a r e low  tially  a t t h e i r bases.  fused  p l a t f o r m s are r e t i c u l a t e d on  es the c a r i n a .  The  and  p o s t e r i o r margin  posterior  be  asymmetry platform  to one  a c i r c u l a r cross  side  section.  smooth.  furrows a d j a c e n t to the  The The  t h i c k and  or The  parcarina  l a t e r a l l y upturned  r e t i c u l a t e ornament extends  i n t o l i n e a r r i d g e s as the ornament approach-  r e t i c u l a t i o n extends around t h e . e n t i r e  posterior  brim  well. Measurements f o r the w i d t h of the p l a t f o r m  ym  the  rounded, forming nodes t h a t a r e  the margin.  v e r y c l o s e to the c a r i n a f a d i n g  The  near the median but may  Longitudinal  a r e r e l a t i v e l y deep, narrow and  The  s i d e of the o t h e r of the  wide cusp has  d e n t i c l e s on  as  frag-  p o s t e r o - l a t e r a l margins a r e p a r a l l e l to s u b - p a r a l l e l w h i l e  p o s t e r i o r end  centre.  larger  lobe.  Description:  The  by  whereas those f o r L2 range from 260  a straight oval  to 300  range between 180  ym.  The  aboral  the o u t l i n e of the p o s t e r i o r  end.  The  a n t e r i o r l y as a narrow groove bordered by a*low, wide k e e l .  g i o n and  o c c u p i e s 6/10  7/10  Discussion:  220  surface  has  shaped p i t surrounded by a r o u g h l y t r i a n g u l a r shaped l o o p  which r o u g h l y f o l l o w s  smooth and  and  of the a b o r a l  The  crimp i s  s u r f a c e w i d t h i n the p o s t e r i o r  i n the medial, to a n t e r i o r A c c o r d i n g to Wardlaw and  p i t extends  re-  parts.  Collinson  (1979b) N.  b i t t e r i i s char-  97  a c t e r i z e d by a p l a t f o r m t h a t a b r u p t l y narrows i n the a n t e r i o r t h i r d f o u r t h of i t s l e n g t h , a low c l e s on the c a r i n a .  cusp of c i r c u l a r c r o s s s e c t i o n , and  They d i s t i n g u i s h t h i s s p e c i e s from _N.  which i s c h a r a c t e r i z e d by a wide p l a t f o r m t h a t has and  t h a t commonly g r a d u a l l y t a p e r s a n t e r i o r l y and  gate-oval cross s e c t i o n . determinations  Unfortunately,  f o r the same m a t e r i a l .  et a l . (1979) i n c l u d e forms Wardlaw and w i t h i n N. r o s e n k r a n t z i and Designation  of any  the p a l e o n t o l o g i s t .  N.  species.  rosenkrantzi  by a l a r g e cusp of differ  in their  Behnken (1971) and  is difficult  i c forms.  s p e c i e s i s a s u b j e c t i v e and  He j u s t i f i e s , t h i s procedure by  This i s impossible  a r b i t r a r y procedure by separating h i s  to a c c o m p l i s h  w i t h conodonts.  Further-  biot-  s p e c i e s open to s u b j e c t i v e and However, the success  and  arbitrary validity  model i s determined by the ease w i t h which another  of t h e s e conodonts can a p p l y  arid C o l l i n s o n ' S - d i a g n o s e s  the model to h i s m a t e r i a l .  Wardlaw  seem more a p p r o p r i a t e l y to f i t the m a t e r i a l from  Island. specimens here r e f e r r e d to N. b i t t e r i a r e i d e n t i f i e d as such  cause of t h e i r v e r y and  1980),  T h i s l e a v e s the i n t e r p r e t a t i o n of importance of v a r i o u s morphol-  of any m o r p h o l o g i c a l  The  species  e x h i b i t e d by d i f f e r e n t but r e l a t e d  procedure w i t h o u t means of r e s o l u t i o n .  Ellesmere  bitteri  to i n t e r p r e t taxonomic problems by comparison to o t h e r  ogic features, for. d i f f e r e n t i a t i n g  student  Clark  C o l l i n s o n (1979b) r e f e r to N.  more, w i t h the l a c k of a f u n c t i o n a l model f o r conodonts (Bengtson, it  elon-  babcocki.  on comparable morphologic v a r i a b i l i t y extant  low d e n t i - r  a b l u n t p o s t e r i o r end  o t h e r authors  C l a r k and  or  t h i c k platforms,  bee.:  t h e i r cusp of c i r c u l a r c r o s s s e c t i o n ,  t h e i r low. n o d i f o r m d e n t i c l e s on the c a r i n a .  Wardlaw and  Collinson(1979b)  f i g u r e specimens of N. b i t t e r i from the G e r s t e r Limestone i n Nevada and. tUtah t  98 and  from the R e t o r t  P h o s p h a t i c s h a l e Member o f the P h o s p h o r i a Formation,.  Wyoming.  There seem to be minor d i f f e r e n c e s between these two c o l l e c t i o n s  which may  be of s i g n i f i c a n c e at the s u b s p e c i f i c l e v e l .  the R e t o r t  The  specimens from  Member appear to have s l i g h t l y rounder p o s t e r i o r margins  t a p e r more g r a d u a l l y then those from the G e r s t e r  Limestone.  The  and  Retort  Member i n c l u d e s asymmetric forms-owing to the . o f f - c e n t r e p o s i t i o n of l a r g e r lobe  (Wardlaw and  C o l l i n s o n (1979b) P I . 1, f i g s .  specimens a r e "very simiiar:,--indeed  11,  12).  the  These  'almost-'identical,.to - those from  Elles-  mere I s l a n d . These specimens d i f f e r from N. n.sp. symmetry, g e n e r a l l y s m a l l e r form brim as opposed to  cusp, and  The  the o t h e r  Ellesmere  are s i m i l a r to those f o r N.  idahoensis  f e a t u r e s make the s e p a r a t i o n c l e a r .  samples do not  t h a t can be p r e s e n t  r e t i c u l a t e microornament on the p l a t -  striations.  The measurements f o r L2 and Wl n.subsp. A but  B_ i n terms of t h e i r d i f f e r e n t  e x h i b i t any  i n N. b i t t e r i ,  postero-lateral denticles  but Wardlaw and  C o l l i n s o n (1979b)  indi-  c a t e t h a t t h o s e t h a t do*are. r a r e v a r i a n t s . NEOGONDOLELLA ROSENKRANTZI n.subsp. D PI. 7, Occurrence :  f i g s . 9-12;  This species  almost b l u n t  i s c h a r a c t e r i z e d by a t h i c k , ^ w i d e  by a w e l l developed brim, and  platform  (F96). with  anterior-  cusp of o v a l c r o s s s e c t i o n surrounded  by a narrow but  directed postero-laterally  The  section  to s l i g h t l y rounded p o s t e r i o r which g r a d u a l l y t a p e r s  l y , by a prominent, o f t e n m o d i f i e d ,  margin.  8.  Upper T r o l d F i o r d Formation, H a m i l t o n P e n i n s u l a  Diagnosis:  c a r i n a and  PI.  s h a l l o w furrow l a t e r a l  towards the. c o r n e r s  s u b s p e c i e s i s based on the e n l a r g e d  of the  to:the  posterior  posterdr-lateral platform  99 margins, the g r a d u a l  t a p e r throughout the e n t i r e l e n g t h , and  rounded p o s t e r i o r , a s opposed to a s t r a i g h t and Description: arched and  A.  J u v e n i l e - The  by the  b l u n t margin.  element i s s m a l l , subsymmetrical,  upturned on i t s margins.  The  slightly  degree of u p t u r n i n g  slightly  i s greatest  i n the a n t e r i o r h a l f whereas the p o s t e r i o r h a l f i s b a r e l y upturned at a l l . The  c a r i n a has  at l e a s t 9 d e n t i c l e s i n c l u d i n g the cusp which, a s . t h e p l a t -  form extends o n l y to the a n t e r i o r t i p of the cusp forms a f r e e b l a d e posteriorly.  The  cusp i s h i g h , e l o n g a t e  o u t l i n e , l a t e r a l l y compressed, and  oval i n cross section, t r i a n g u l a r i n  directed slightly posteriorly.  t i c l e s a n t e r i o r to the cusp a r e p o i n t e d , and  increase s l i g h t l y The  p l a t f o r m t a p e r s g r a d u a l l y towards the a n t e r i o r and  The  a b o r a l s u r f a c e i s smooth and  terminates  p o s t e r i o r l y i n a high elongate  bears a  reti-  i t s margins.  the a n t e r i o r 1/3  the p o s t e r i o r 1/3  1  bears a h i g h narrow k e e l which  o v a l b a s a l loop or  flange.  element i s subsymmetrical, s l i g h t l y a r c h e d ,  s c a r c e l y upturned on and  compressed  to t h e margins over t h e . e n t i r e l e n g t h o f the  platform.  I n t e r m e d i a t e - The  den-  i n height a n t e r i o r l y .  c u l a t e ornament r e s t r i c t e d  B.  distinct, laterally  The  This s l i g h t upturning  and  i s greatest  in  whereas the m i d d l e p a r t i s f l a t .  The  p l a t f o r m margins a r e s u b p a r a l l e l to g r a d u a l l y t a p e r i n g over the  poster-  ior  2/3  platform  whereas the a n t e r i o r 1/3  t a p e r s a l i t t l e more r a p i d l y .  extends the e n t i r e l e n g t h of the element arid i s a d j a c e n t s m a l l brim, behind, the cusp. The  The  t o , or forms a ..  p o s t e r i o r margin i s rounded i n o u t l i n e .  r e t i c u l a t e microornament, which i s r e s t r i c t e d to the margins,  throughout the e n t i r e l e n g t h of the element. and  The  The  a b o r a l s u r f a c e i s smooth  b e a r s a r e l a t i v e l y h i g h , wide k e e l which t e r m i n a t e s  s l i g h t l y squared-off  basal  loop.  occurs  i n a rounded to  TOO The cusp i s h i g h , o v a l i n x r o s s upwards i f not b a r e l y a n t e r i o r l y . which t h e r e a r e 11 or 12, a r e fused at t h e i r bases.  s e c t i o n , p o i n t e d , and  The  d e n t i c l e s a n t e r i o r to the cusp, of  i n c r e a s e i n height, and The  first  slightly distinct  t i p s of the d e n t i c l e s a r e f l a t  p o s t e r i o r h a l f and  compression a n t e r i o r l y  and  four d e n t i c l e s are barely p e r c e p t i b l y  more c l o s e l y spaced, of equal h e i g h t and the c a r i n a . The  directed straight  from the  others-on  to s l i g h t l y p o i n t e d i n the  p o i n t e d i n the a n t e r i o r h a l f .  C. Mature to G e r o n t i c - I t i s at t h i s stage t h a t the many d i a g n o s t i c f e a ~ . „. t u r e s of the s p e c i e s and  s u b s p e c i e s become apparent.  e x i s t even between t h i s s p e c i e s and mediate and  N.  Only minor d i f f e r e n c e s  i d a h o e n s i s n.subsp. A a t the  inter-  e s p e c i a l l y a t the j u v e n i l e growth s t a g e .  The p l a t f o r m element has a r o u g h l y  elongated  t r i a n g u l a r o u t l i n e and  a v e r y r o b u s t appearance owing to i t s w i d t h (Wl = 200 t h i c k margins.  to 260  ym)  and  very  A r c h i n g of .:the element i s g r e a t e r than t h a t observed  i n t e r m e d i a t e forms whereas the degree of u p t u r n i n g specimens a c t u a l l y show a degree of downturning. s t r a i g h t to s l i g h t l y rounded and  f o r the  i s imperceptible.  Some  The p o s t e r i o r margin i s  sometimes l o b e d but not w i t h the same  asymmetry as N. b i t t e r i n.subsp. C.  The p o s t e r o - l a t e r a l margins a r e en^-  l a r g e d to the a n t e r i o r end of the cusp,where the l a t e r a l margins remain p a r a l l e l u n t i l they b e g i n to t a p e r more r a p i d l y Whereas.an extensive' brim form  i t i s absent  from the a n t e r i o r t i p , where the l a s t 2 to 3 almost The margins of the p l a t f o r m  bear a r e t i c u l a t e microornament which i s widest  does not  1/3.  i s formed about the cusp on the p o s t e r i o r of. t h e  t o t a l l y fused d e n t i c l e s form a f r e e b l a d e .  extends over  . i n the a n t e r i o r  sub-  the e n t i r e l e n g t h .  The  near the m i d - l e n g t h  i n t e r i o r border  of the  and  reticulation  end a b r u p t l y but r a t h e r fades i n t o f a i n t r i d g e s d i r e c t e d towards  plat-  101  the c a r i n a .  On  some specimens t h i s r e t i c u l a t i o n can  c l e s and  cusp but  smooth.  The  s h a l l o w and  cusp i s g e n e r a l l y  low,  the median of the p l a t f o r m  narrow and  p a r a l l e l the c a r i n a up  elongate oval and  directed  specimens i t i s fused  an e l o n g a t e r i d g e d i r e c t e d  i n o u t l i n e and  crease again.  whereas i n a  w i t h a c o u p l e of a d j a c e n t d e n t i c l e s  towards a p o s t e r o - l a t e r a l c o r n e r .  to about 1/2  to 2/3  The  forming  majority  are f l a t - t o p p e d ,  of t h e i r h e i g h t and  The  the d e n t i c l e s f i r s t  smallest  increase  of  f o u r d e n t i c l e s a n t e r i o r t o t h e cusp s m a l l e r  equal  then  aboral  surface  a t e s i n a h i g h e r and  i s smooth and  bears a low,  roughly t r i a n g u l a r  loop i s quite v a r i a b l e .  The  i n a s t r a i g h t elongate oval p i t .  fused, In  Rare specimens have  the  and more c l o s e l y spaced  basal  narrow b a s a l The  keel  wide k e e l which t e r m i n -  loop.  The  shape of t h i s  tip.  The  crimp c o v e r s r o u g h l y 6/10  platform.  ba-  groove t e r m i n a t e s p o s t e r i o r l y  increases,.;in h e i g h t and  narrows  i n a b l a d e - l i k e r i d g e which connects w i t h the o r a l f r e e b l a d e at the  o f the  de-  others.  The  ior  lat-  f r e e of the p l a t f o r m .  o t h e r specimens the d e n t i c l e s a r e more n o d e - l i k e .  than the  i n h e i g h t and  a n t e r i o r d e n t i c l e s a r e almost t o t a l l y  forming a c h a r a c t e r i s t i c b l a d e which i s i n p a r t  sal  a t about  In some specimens  to the m i d l i n e  10)  pos-  From t h i s p o i n t , which r o u g h l y c o i n c i d e s w i t h the more r a p i d l y t a p -  e r i n g of the p l a t f o r m ,  first  The  corners.  positioned  s t r a i g h t upwards.  of the d e n t i c l e s a n t e r i o r to the cusp (as many as e r a l l y compressed, fused  In  to the  the p o s t e r o - l a t e r a l  the cusp i s d i r e c t e d e i t h e r d e x t r a l or s i n i s t r a l  height.  furrow i s •-.  almost s t r i a t e d i n appearance.  i t i o n of the cusp where they d i v e r g e towards  of- the  denti-  r e t i c u l a t i o n extends around the brim i n most specimens.  smooth furrows a r e  few  found on the  even i n these specimens a t l e a s t p a r t of the  some, t h i s r e t i c u l a t i o n i s f a i n t and  The  be  of the w i d t h i n the p o s t e r i o r  anterparts  102 Discussion:  The  e a r l y growth stages  n.subsp. A.  S z a n i a w s k i and  a r e v e r y s i m i l a r even to N_.  Malkowski (1979) p o i n t e d  out  the  idahoensis  similarity  of e a r l y growth stages between N t idaho ens i s - ahdff N. --h i t t er i . • - T-his s i m i l a r i t y of e a r l y growth stages  throughout the phylogeny of N e o g o n d o l e l l a  to the c l o s e r e l a t i o n s h i p of a l l the s p e c i e s . hoensis  points  Some mature forms of N.  n.subsp. A a c t u a l l y mimic N_. r o s e n k r a n t z i .  ida-  Mature specimens of  N.  r o s e n k r a n t z i n.subsp. D _ d i f f e r i n terms of a more t r i a n g u l a r o u t l i n e , l a r ger number of d e n t i c l e s , shape and the a n t e r i o r b l a d e (a f e a t u r e not and  t h i c k e r p l a t f o r m margins.  arrangement of the d e n t i c l e s i n c l u d i n g seen i n N.. i d a h o e n s i s ) , s h a l l o w e r  furrows,  However, f o r r e l i a b l e i d e n t i f i c a t i o n  this  mimicry p o i n t s to the need, not o n l y f o r mature forms, but a l s o f o r enough r e p r e s e n t a t i v e s to i n c l u d e a l l of the wide v a r i a b i l i t y of form so  charac-  t e r i s t i c of t h e s e s p e c i e s . As p o i n t e d out  i n the d i s c u s s i o n f o r N. b i t t e r i n.subsp. C. the main  f e a t u r e s d i s t i n g u i s h i n g N.. r o s e n k r a n t z i . from N.. b i t t e r i a r e the s l i g h t l y straight  to almost b l u n t p o s t e r i o r ends, and  oval cross section. different  As w e l l the e n l a r g e d  a l a r g e cusp of  elongate-  p o s t e r o - l a t e r a l margins  symmetry d i s t i n g u i s h N. r o s e n k r a n t z i n.subsp. I) from N.  and bitteri  n.subsp. C^. Wardlaw and  C o l l i n s o n (1979b) f i g u r e . s p e c i m e n s  of N. r o s e n k r a n t z i from  the R e t o r t P h o s p h a t i c Shale Member of the Phosphoria Formation and  from  the G e r s t e r Limestone..  and  The  l a t t e r have a v e r y b l u n t p o s t e r i o r end  pear more "advanced" than the E l l e s m e r e  specimens, which more c l o s e l y r e -  semble the R e t o r t Member specimens i n p o s t e r i o r o u t l i n e and manner and g r e e of t a p e r i n g .  ap-  There seem to be grounds f o r s u g g e s t i n g  f e r e n c e s a r e s i g n i f i c a n t a t the i n f r a s p e c i f i c l e v e l  that these  (subspecies).  To  dedifcall  a t t e n t i o n to some of t h e s e d i f f e r e n c e s , t h e r e f o r e , I have r e f e r r e d t h e s e  103 specimens t o a d i s t i n c t The  subspecies.  specimens d i f f e r  from o t h e r s p e c i e s i n t h e T r o l d F i o r d  Formation  i n t h e i r d e n t i c l e shape and c o n f i g u r a t i o n , t h e shape and p o s i t i o n o f t h e cusp,  the g r e a t e r t h i c k n e s s o f t h e p l a t f o r m margins and t h e p o s i t i o n and  c o n f i g u r a t i o n "of r e t i c u l a t e ornament. Ramif orm.. Elements NEOGONDOLELLA IDAHOENSIS n.subsp. A - XANIOGNATHUS PI. i, Ozarkodina Occurrence:  T h i s blade-shaped  tortilis  Tatge, 1956 Hamilton  Peninsula section.  element has a l o n g a n t e r i o r p r o c e s s w i t h as  many as 8 sharp, p o i n t e d , subequal inclined posteriorly.  (Tatge)  f i g s . 1-4.  Lower A s s i s t a n c e Formation,  Description:  TORTILIS  and l a t e r a l l y compressed d e n t i c l e s , a l l  The cusp i s h i g h , sharp and l a t e r a l l y  compressed.  At l e a s t t h r e e d e n t i c l e s a r e p r e s e n t on t h e s h o r t p o s t e r i o r p r o c e s s , which i s t w i s t e d t o one s i d e o r t h e o t h e r . i s grooved and t e r m i n a t e s cusp.  The u n d e r s u r f a c e  o f both  processes  i n a pronounced b a s a l p i t d i r e c t l y below t h e main  The d e n t i c l e s a l l bear a v e r y f i n e s t r i a t e microornament..  Discussion:  T h i s s p e c i e s i s d i s t i n g u i s h e d from t h e o l d e r X. a b s t r a c t u s  by t h e l e s s r o b u s t b l a d e and t h e t w i s t e d (as opposed t o s t r a i g h t ) p o s t e r i o r process.  A c c o r d i n g t o Behnken (1975) t h e range o f X. t o r t i l i s  about t h e end o f t h a t o f N. s e r r a t a .  T h i s would i n d i c a t e a L a t e Roadian  age but t h i s s p e c i e s o c c u r s w i t h N e o g o n d o l e l l a age.  begins a t  p l a t f o r m s o f E a r l y Roadian  These specimens may be i n t e r m e d i a t e between X. t o r t i l i s and t h e o l d e r  X. a b s t r a c t u s as t h e base o f t h e d e n t i c l e s appear more r o b u s t than of  X. t o r t i l i s  f i g u r e d by Behnken. ( 1 9 7 5 )  to  X. t o r t i l i s  i n most r e s p e c t s .  ;  specimens  n e v e r t h e l e s s they--do Appear  closer  104 NEOGONDOLELLA.IDAHOENSIS n.subsp. A - ELLISONIA EXCAVATA Behnken,  1975  P I . 1, f i g . 15. Occurrence:  Lower A s s i s t a n c e Formation,  Description:  The  Hamilton P e n i n s u l a s e c t i o n .  specimens have a v a r i a b l e h i n d e o d e l l i f o r m morphology  w i t h , beneath the- cusp,- a s m a l l conical'basa'l- p i t compressed and at l e a s t  is  laterally  The p o s t e r i o r bar i s l o n g and  bears  10 to 12 d i s c r e t e p o i n t e d d e n t i c l e s t h a t a r e of v a r i a b l e s i z e .  a l o n g the bar. anterior  inclined posteriorly.  which  3 d e n t i c l e s a r e p r e s e n t on the downward p r o j e c t i n g s h o r t  bar.  Discussion:  The. f i g u r e d .spec imehif. probably: r e p r e s e n t s the LB.: element of a  multielement  s p e c i e s which i n c l u d e s U, LA,  observed.  and  LF elements which were not  Behnken (1975) i n d i c a t e s t h a t t h i s s p e c i e s o c c u r s w i t h N.  e n s i s i n West Texas., and would thus have a L e o n a r d i a n  age.  NEOGONDOLELLA IDAHOENSIS n.subsp. A - ELLISONIA TRIBULOSA ( C l a r k and ton,  Occurrence:  and  ( f i g s . 5, 8) elements a r e p r e s e n t  LC elements LAI  and  ( f i g s . 6, 7) and  LC  described. l o h c h o d i n i f o r m element w i t h a  p o s t e r i o r l y i n c l i n e d cusp.  The p o s t e r i o r  3 to 4 d e n t i c l e s whereas t h e s h o r t a n t e r i o r bar bears  but l a r g e r d e n t i c l e s . expansion  1962)  - T h i s i s a s l i g h t l y asymmetrical  h i g h , l a t e r a l l y compressed and bar bears  Ething-  PI. 2, f i g s . 5-8 Apatognathus t r i b u l o s u s C l a r k and E t h i n g t o n , 1962 Lower A s s i s t a n c e Formation, Hamilton P e n i n s u l a  D e s c r i p t i o n : Of t h e U, LAI, LA2  LAI  idaho-  2 pointed  The b a s a l p i t below the cusp i s formed by v e r y  of a groove which i s p r e s e n t over  slight  the e n t i r e l e n g t h of the a b o r a l  surface. LC - T h i s i s an e n a n t i o g n a t h i f o r m  element w i t h a l o n g p o s t e r i o r bar  p r o j e c t i n g downward from the main cusp and  bearing 5 d e n t i c l e s that are  105  d i s c r e t e and s u b p a r a l l e l f o r much o f t h e i r l e n g t h .  The main cusp i s t r i a n -  g u l a r i n c r o s s s e c t i o n and.the c o r n e r s a r e extended i n t o sharp r i d g e s .  The  s h o r t downward p r o j e c t i n g a n t e r i o r bar b e a r s one t o two d e n t i c l e s and p r o j e c t s a t an a n g l e  o f about 60 degrees t o t h e p o s t e r i o r b a r .  Below t h e cusp  i s a large t r i a n g u l a r basal p i t . Discussion:  This species occurs  postserrata indicating sence o f b o t h E.  a  Roadian t o E a r l y Wordian age.  excavata and E..' / t f i b u l o s a t o g e t h e r  t h e i r ranges i n t h e E a r l y Roadian age  assigned  through t h e range o f N. s e r r a t a and N.  which f i t s  Perhaps t h e p r e -  i n d i c a t e s o v e r l a p of  i n w e l l w i t h t h e E a r l y Roadian  t o N. i d a h o e n s i s n.subsp. A because o f i t s i n t e r m e d i a t e  t i o n between N. i d a h o e n s i s  posi-v'.  and N. s e r r a t a .  NEOGONDOLELLA IDAHOENSIS n.subsp. A - PRIONIODELLA DEGRESCENS Pl.  Tatge, 1956  1, f i g . 14.  O c c u r r e n c e : Lower A s s i s t a n c e Formation, H a m i l t o n P e n i n s u l a  section.  D e s c r i p t i o n : This- specimen' I s a s h o r t , s t r a i g h t and d e n t i c u l a t e "element w i t h o u t a main cusp. in  The s i x d i s c r e t e and p o i n t e d  d e n t i c l e s a r e subequal  height.  Discussion:  Behnken (1975) i n c l u d e s t h i s s p e c i e s w i t h i n t h e range of N.  postserrata,  i n o t h e r words, E a r l y Wordian.  much g r e a t e r r a n g e s . species are fragmental Addendum:  However, s i m i l a r forms have  Other elements which may be a s s i g n e d  to Prioniodella  and u n i d e n t i f i e d .  A paper, i n p r e p a r a t i o n by t h e author d e s c r i b e s  Neogondolella  i d a h o e n s i s n.subsp. A, N_. n s p . By N. b i t t e r i n.subsp. C and N. r o s e n k r a n t z i n.subsp. D a s N. i d a h o e n s i s p r a e s e r r a t a n.subsp., N.. p e r r y i n.sp., N.  bitteri  a r c t i c a n.subsp. and N_. r o s e n k r a n t z i e l l e s m e r e n s i s n.subsp., r e s p e c t i v e l y .  106 REFERENCES Bamber, E.W. and Waterhouse, J.B. 1971: C a r b o n i f e r o u s and Permian s t r a t i g r a p h y . a n d p a l e o n t o l o g y , n o r t h e r n Yukon T e r r i t o r y , Canada; B u l l , of Can. P e t r o l . 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Upper P a l e o z o i c and Mesozoic s t r a t i g r a p h y i n the Y e l v e r t o n Pass r e g i o n , E l l e s m e r e I s l a n d , D i s t r i c t of F r a n k l i n ; G e o l . Surv. Can., Paper 68-31.  Nassichuk, 1965:  W.W., F u r n i s h , W.M. and G l e n i s t e r , B.F.. The Permian ammonoids of A r c t i c Canada; Bull. 131.J  Nassichuk, 1970:  W.W. and Spinosa, C H e l i c o p r i o n sp., a Permian elasmobranch,from E l l e s m e r e I s l a n d , Canadian A r c t i c ; J . P a l e o n t o l . , 44, pp. 1130-1132.  Newell, N.D. 1956: F o s s i l populations; pp. 63-82.  G e o l . Surv.  Can.  Systematics A s s o c i a t i o n P u b l i c a t i o n ,  #2,  N i c o l l , R.S. 1975: The e f f e c t of L a t e C a r b o n i f e r o u s - E a r l y Permian g l a c i a t i o n on the d i s t r i b u t i o n of conodonts i n A u s t r a l i a ; The G e o l o g i c a l ' . A s s o c i a t i o n of Canada S p e c i a l Paper, 15, pp. 273-278. Raup, D.M. 1971:  Stanley, 1975:  1979:  and S t a n l e y , S.M. P r i n c i p l e s of P a l e o n t o l o g y ; c i s c o , 388p.  W.H.  Freeman and  Company, San  S.M. A t h e o r y of e v o l u t i o n above the s p e c i e s l e v e l ; Acad. S c i . , 72, pp. 646-650.J/ Macroevolution: P a t t e r n and P r o c e s s ; San F r a n c i s c o , 332p.  W.H.  Proc.  Fran-  Natl.  Freeman and  Company,  S t a p l i n , F. 1969: Sedimentary o r g a n i c m a t t e r , o r g a n i c metamorphism, and o i l and gas o c c u r r e n c e ; B u l l . Can. P e t r o l . Geol., 17, pp. 47-66. 1975:  I n t e r p r e t a t i o n of thermal h i s t o r y from c o l o u r of o r g a n i c matter; P a l y n o l o g y , 1.  particulate  S t a u f f e r , C R . arid Plummer, H.J. 1932: Texas P e n n s y l v a n i a n conodonts and t h e i r s t r a t i g r a p h i c Texas Univ.:.Bull., 3201, pp. 13-50. Stebbins, 1977:  G.L. Processes, of o r g a n i c e v o l u t i o n ; C l i f f s . , New J e r s e y , 269p.  relations;  P r e n t i c e H a l l Inc., Englewood  Sweet, W.C I:' 1970: Uppermost Permian and Lower T r i a s s i c conodonts of the S a l t Range and Trans-Indus Ranges, West P a k i s t a n ; Univ. Kansas Dept. G e o l . Spec. P u b l . #4, pp. 207-275.  Ill  S y l v e s t e r - B r a d l e y , P.C. 1951: The s u b s p e c i e s i n p a l e o n t o l o g y ; 88-102.  G e o l o g i c a l Magazine, 88, pp.  S z a n i a w s k i , H". and Malkowski, K. 1979: Conodonts form t h e Kapp S t a r o s t i n Formation (Permian) o f S p i t s bergen; A c t a P a l a e o n t o l o g i c a P o l o n i c a , 24, pp. 231-264. Tatge, U. 1956;  Conodonten aus dem germanischer Muschelkalk; 30, pp. 108-127.  Palaont.  Zeitschr,  T h o r s t e i n s s o n , R.. = 1974": CaTboniferous-;;and_>Permian s t r a t i g r a p h y o f A x e l H e i b e r g I s l a n d and western E l l e s m e r e I s l a n d , Canadian A r c t i c A r c h i p e l a g o ; Geol. Surv. Can. B u l l . , 224, 115p. Tozer, E.T. and T h o r s t e i n s s o n , R, 1964: Western Queen E l i z a b e t h I s l a n d s , A r c t i c A r c h i p e l a g o ; Can. Mem. 332. V a l e n t i n e , J.W. 1973: E v o l u t i o n a r y P a l e o e c o l o g y o f t h e Marine B i o s p h e r e ; H a l l Inc., Englewood C l i f f s , New Jersey, 511p.  Geol.  Surv.  Prentice-  :  Von B i t t e r , P.H. 1976: The apparatus of G o n d o l e l l a s u b l a n c e o l a t a G u n n e l l (Conodontop h o r i d a , Upper Pennsylvanian) and, i t s r e l a t i o n s h i p t o I l l i n e l l a t y p i c a Rhodes; L i f e S c i e n c e s .Contrs. Royal O n t a r i o Museum, 109, 44p. Von B i t t e r , P.H. and M e r r i l l , G.K. 1977: N e o g o n d o l e l l i f o r m conodonts o f E a r l y and M i d d l e P e n n s y l v a n i a n age; L i f e S c i e n c e s O c c a s i o n a l Paper Royal O n t a r i o Museum, 29, lOp. Wardlaw, B.R. and C o l l i n s o n , J.W. 1978: S t r a t i g r a p h i c r e l a t i o n s o f Park C i t y Group (Permian) i n e a s t e r n Nevada and western Utah; Am. Assoc. P e t r o l . G e o l o g i s t s B u l l l , 62, pp. 1171-1184. 1979a: 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 t h e Park C i t y Group; Geol. Surv., P r o f . Pap. 1163D, pp.. 17-22.  United States  1979b: Youngest Permian conodont.faunas, from t h e Great B a s i n and Rocky Mountains; Brigham Young Univ. Geology S t u d . , 2 6 , pp. 151-164. Waterhouse, J.B. 1973: Communal h i e r a r c h y and s i g n i f i c a n c e of. environmental parameters f o r B r a c h i o p o d s : t h e New Zealand. Permian model;.'Life S c i e n c e s C o n t r s . Royal O n t a r i o Museum, 92, pp. 1-49,  112  1976:  World C o r r e l a t i o n s f o r Permian Marine :Faunas; . U n i v e r s i t y of Queensland Papers, Dept. o f Geology, 7, 232p.  Waterhouse, J.B., Waddington, J . and A r c h b o l d , N. 1978: The e v o l u t i o n o f t h e M i d d l e C a r b o n i f e r o u s to L a t e Permian b r a c h i o p o d s u b f a m i l y S p i r i f e r e l l i n a e , Waterhouse; I n Western and A r c t i c Canadian B i o s t r a t i g r a p h y , C R . S t e l c k and B.D.E. C h a t t e r t o n ( E d s . ) , G e o l . A s s o c . o f Can. Spec. Paper, 18, pp. 415-443. Weimer, R.J. 1970: Rates o f d e l t a i c s e d i m e n t a t i o n and i n t r a b a s i n d e f o r m a t i o n , Upper Cretaceous o f Rocky Mountain Region; I n D e l t a i c Sedi m e n t a t i o n : Modern and A n c i e n t , J . P i Morgan ( E d . ) , S o c i e t y o f Economic P a l e o n t o l o g i s t s and M i n e r a l o g i s t s Spec. P u b l . #15, pp. 270-292. W i l s o n , L.R, 1962: P l a n t M i c r o f o s s i l s , Flowerpot Formation;'. Oklahoma G e o l . Surv. C i r c u l a r , 49, 50p. Wright, S. 1967: Comments on.the p r e l i m i n a r y working papers, d f Eden and Waddington; I n Mathematical. C h a l l e n g e s to t h e Neo-Darwinian Theory of E v o l u t i o n , P.S. Moorehead. and M.M'. Kaplan ( E d s . ) , The W i s t a r I n s t . Symposia Monograph #5, P h i l a d e l p h i a , pp. 117-120. Y o u n g q u i s t , W., Hawley, R.W. and M i l l e r , A.K. 1951: P h o s p h o r i a conodonts from S.E. Idaho; 356-364.  J . P a l e o n t o l . , 25, pp.  Z i e g l e r , W. (Ed.) 1973: Catalogue o f Conodonts; -E". S c h w e i z e r b a r t ' sche Verlagsbuchhandl u n g , v o l . 1, 504p.  113 Explanation f o r P l a t e 1 A l l f i g u r e s are Scanning E l e c t r o n Micrographs. F i g s . 1, 2  Neostreptognathodus p r a y i  1. O b l i q u e l a t e r a l t o o r a l view. loc.TlOO.  Page Note t h e r e c r y s t a l l i z e d  73  texture.  (X250).  2. O b l i q u e l a t e r a l t o o r a l view o f p l a t f o r m p o s t e r i o r . F 1 0 0 . (X125). F i g s . 3-6 Anchignathodiis minutus Page 3. L a t e r a l view. Note d e n t i t i o n a n t e r i o r t o cusp. F49*. (X100) .  73  4. O r a l v i e w . F 4 9 . ( X I 0 0 ) . 5. L a t e r a l view.  Note l a c k o f d e n t i t i o n a n t e r i o r t o cusp.F49.(X100).  6. C l o s e - u p of d e n t i t i o n on F i g . 5.  Note t h e smooth  surface  free  o f ornament.F49. (X400). F i g s . 7-13  N e o g o n d o l e l l a i d a h o e n s i s subsp. i n d e t .  Page  74  7. O r a l view o f i n t e r m e d i a t e to mature form.F100. (X125). 8. L a t e r a l v i e w o f a n t e r i o r p a r t o f element showing d i s c r e t e  denti-  c l e s . F100. (X125). 9. L a t e r a l v i e w o f p o s t e r i o r o f i n t e r m e d i a t e  form.F100.(X125).  10. O r a l v i e w o f i n t e r m e d i a t e form.FlOO. (X125). 11. O r a l v i e w o f mature form.F100. (X250). 12. C l o s e - u p o f r e c r y s t a l l i z e d  (apatite) texture.F100.(X500).  13. A b o r a l v i e w of p o s t e r i o r of i n t e r m e d i a t e Fig.  14  form.F100.(X125).  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A - P r i o n i o d e l l a d e c r e s c e n s  14. L a t e r a l view.F49. (X100). Fig.  15  Page 105  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A - E l l i s o n i a e x c a v a t a  15. L a t e r a l view..F49. (X100) .  P  a  8  e  1  0  4  115 Explanation for Plate 2 All  f i g u r e s a r e Scanning E l e c t r o n M i c r o g r a p h s .  F i g s . 1-4  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A - Xaniognathus  1. Close-up o f d e n t i c l e . Note t h e f i n e s t r i a t e d  tortilis  surface texture.F49.  (X400).  P  ^  a  1  0  3  2. Close-up of d e n t i c l e . F 4 9 . (X400). 3. L a t e r a l view. F49. (X100). 4. L a t e r a l view. F49. (X100) . F i g s . 5-8  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A - E l l i s o n i a  5. LC element.  tribulosa  Note r i d g e o r k e e l on main cusp.F49. (X100). Page 104  6. LAI element.F49. (X100). 7. LAI element.F54. (X150). 8. LC element.F49. ( X I 0 0 ) . Figs.  9-19  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A  9. O r a l view o f v a r . g r a c i l i s . to  Page  Main cusp and p o s t e r i o r end  the bottom o f t h e page.F49. (X100).  pointing P  10. O r a l v i e w o f v a r . i n t e r m e d i a t u s . F 4 9 . (X100). 11. O r a l view o f v a r .  75  a  g  e  8  Page  4  86  intermediatus.F54.(X90).  12. O r a l view of v a r . g r a c i l i s .  Note the d i s t i n c t  four  denticles  a n t e r i o r t o the cusp.F49.(X100). 13. O r a l view of v a r .  intermediatus.F49.(X75).  14. O r a l v i e w o f v a r . r o b u s t u s .  Note the s e r r a t i o n s on a n t e r i o r  F49.(X100).  P  a  g  e  1/3. 8  5  15. L a t e r a l view of p o s t e r i o r showing l a r g e cusp and l a c k of b r i m p o s t e r i o r to c u s p : i n t e r m e d i a t e to mature.F49.(X100). 16. L a t e r a l view o f j u v e n i l e . compressed,  triangular outlined  Note t h e l a c k of p l a t f o r m on  laterally  cusp.F49. (X150).  17. O b l i q u e l a t e r a l view of j u v e n i l e . F 4 9 . (X150). 18. O b l i q u e l a t e r a l view of i n t e r m e d i a t e . F 4 9 . (X100). 19. O b l i q u e l a t e r a l view o f j u v e n i l e .  Note the g r a d u a l i n c r e a s e i n  d e n t i c l e h e i g h t a n t e r i o r l y and t h e u p t u r n i n g on the p l a t f o r m margin.F49.(XI00).  117  Explanation All  for Plate 3  f i g u r e s a r e Scanning E l e c t r o n M i c r o g r a p h s .  Figs.  1-17  Neogondolella idahoensis  n.subsp. A  1. O r a l view v a r . i n t e r m e d i a t u s .  Note  Page 75  four d i s t i n c t d e n t i c l e s  anter-  i o r to cusp. F49.(X80). 2. O r a l view v a r . i n t e r m e d i a t u s . F49. (X10Q).  Page 86  3. O r a l v i e w v a r . g r a c i l i s . F 4 9 . ( X 7 5 ) .  Page 84  4. O r a l view v a r . g r a c i l i s . F 4 9 - ( X 7 5 ) . 5. O r a l view v a r . g r a c i l i s . F 4 9 . ( X I 0 0 ) . 6. O r a l view v a r .  gracilis.F49.(X100).  7. O r a l view v a r . r o b u s t u s . F 4 9 . ( X I 0 0 ) . 8. O r a l view showing h i g h ,  Page 85  d i s c r e t e d e n t i c l e s at a n t e r i o r  end.F54.(X150).  9. O r a l view v a r . r o b u s t u s . F 4 9 . ( X 1 0 0 ) .  that  10. A b o r a l  view.F54.(X125).  11. A b o r a l  view.  Note d i f f e r e n t o u t l i n e of p o s t e r i o r m a r g i n from  i n F i g . 10.F54.(X125). 12. O r a l view o f t h i c k p l a t f o r m margins of mature 13. L a t e r a l v i e w o f i n t e r m e d i a t e  form.F49.(X100).  14. L a t e r a l v i e w o f i n t e r m e d i a t e  to mature  15. L a t e r a l , view of mature form.  form.F54.(X85).  form.F49.(X80).  Note t h a t t h e p l a t f o r m  reaches the  p o s t e r i o r o f t h e cusp ( u n l i k e t h a t i n F i g s . 13, 14). F54.(X80). 16. L a t e r a l view o f v a r . g r a c i l i s . F 4 9 . ( X l O O ) . 17. L a t e r a l v i e w o f v a r . g r a c i l i s . F 4 9 . ( X l O O ) .  119  Explanation  for Plate 4  A l l f i g u r e s are F i g s . 1-10 1. N.  Neogondolella idahoensis  n.subsp. A  Page  O r a l v i e w o f mature to g e r o n t i c form of v a r . l o b a t u s  rosenkrantzi 2.  Scanning E l e c t r o n Micrographs.  i n shape.  t h a t mimics  Note d i s c r e t e a n t e r i o r d e n t i c l e s . F49.(X75).  O r a l v i e w of v a r .  3. O r a l v i e w of v a r .  75  intermediatus•F49.(X100). intermediatus  the l a c k o f r e t i c u l a t e d ornament on  Page  with anterior serrations.  the a n t e r i o r - m o s t  m a r g i n s of  86  Note  platform.  F54.(X75). 4.  C l o s e - u p of a n t e r i o r p a r t of F i g . 3. F54.(X150).  5.  O r a l v i e w of mature to g e r o n t i c form.F54. (X125).  6.  O r a l v i e w of mature to g e r o n t i c  posterior  denticles.F54.(X150).  7.  O r a l v i e w of v a r .  8.  O r a l v i e w of v a r . l o b a t u s .  d i s c r e t e , very 9.  form showing p a r t i a l f u s i o n of  constrictus.F49.(X100).  Page  Note the t h i c k p l a t f o r m m a r g i n  l a t e r a l l y compressed d e n t i c l e s . F 5 4 . (X150).  O r a l v i e w of i n t e r m e d i a t e  to mature v a r .  the l a c k of r e t i c u l a t e d ornament at the a n t e r i o r end 10.  of the  and  Page  intermediatus.  86  87  Note  platform.F54.(X150).  O r a l v i e w of g e r o n t i c form w i t h complete f u s i o n of p o s t e r i o r  denticles. d i s p l a y e d by  Note the g r a d u a l F i g s . 5,  p h o r i e n s i s w i t h N.  6 and  and 10.  progressive  f u s i o n of p o s t e r i o r d e n t i c l e s  T h i s demonstrates the  idahoensis.F54.(X180).  synonymy of N.  phos-  Plate 4  120  121  Explanation f o r Plate 5 A l l f i g u r e s a r e Scanning E l e c t r o n M i c r o g r a p h s . F i g s . 1-10  N e o g o n d o l e l l a i d a h o e n s i s n.subsp. A  1. Close-up o f r e t i c u l a t e d ornament. rounded  Page  Note the d i s t i n c t  edges o f r i d g e s on the o u t s i d e margin  (left),  75  but  the d i s t i n c t  and  sharp r i d g e s on the m i d - p a r t o f t h e p l a t f o r m margin, and f a d i n g and  elon-  gate r i d g e s near t h e furrow ( r i g h t ) . F49.(X1500). 2. Close-up o f P l . 2, F i g . 14 showing  the presence o f  reticulated  ornament on the d e n t i c l e t i p . F 4 9 . ( X 4 0 0 ) . 3. Close-up o f P l . 2, F i g . 14 showing  the p r e s e n c e o f r e t i c u l a t e d  ornament on the cusp and f u s e d p o s t e r i o r d e n t i c l e s but absence on t h e furrows.F49.(X400). 4. A b o r a l v i e w o f j u v e n i l e form showing  t h e h i g h , narrow  keel  and  e l o n g a t e - o v a l loop. F49.(XI00). 5. A b o r a l  view.F49.(X100).  6. A b o r a l v i e w of i n t e r m e d i a t e form.F49. (X100). 7. A b o r a l view of i n t e r m e d i a t e t o mature form showing k e e l and  slightly triangular  low, wide  loop.F49.(X100).  8. A b o r a l v i e w of mature form showing v e r y low and wide k e e l triangular loop.  Note the p r o g r e s s i v e changes  of a b o r a l f e a t u r e s  and from  F i g . 5 ( j u v e n i l e ) t o F i g . 8 (mature). F49.(XI00). 9. A b o r a l view of p o s t e r i o r end of mature 10. Close-up o f l o o p i n F i g . 9.  form.F49.(X150).  Note the t r u n c a t i o n o f growth  lamellae  a t p o s t e r i o r o f l o o p ( e s p e c i a l l y e v i d e n t on r i g h t hand s i d e ) which accomp a n i e s t h e t r a n s i t i o n from e l o n g a t e o v a l to t r i a n g u l a r shape o f l o o p . F 4 9 . (X600).  Plate 5  122  123  Explanation f o r Plate 6 All  f i g u r e s a r e Scanning E l e c t r o n M i c r o g r a p h s .  F i g s . 1-4  N e o g o n d o l e l l a n.sp. B  1. O r a l view o f i n t e r m e d i a t e form.  Page Note t h e v e r y symmetric  92  shape.  F83.(X250). 2. O r a l view o f mature form showing l a r g e c i r c u l a r cusp d i r e c t e d s t r a i g h t upwards and w i t h c o a r s e s t r i a t e ornament on brim.F83.(X300). 3. O r a l view o f symmetric mature form.F36.(X250). 4. O r a l view o f g e r o n t i c form w i t h l a r g e , c i r c u l a r ( i n c r o s s  section)  cusp w i t h narrow p o s t e r i o r p l a t f o r m margins and f u s e d d e n t i c l e s . F36.(X250). F i g s . 5, 6  Neogondolella postserrata(?)  Page  94  5. O r a l view.F87.(X125). 6. Close-up of F i g . 5 showing b l u n t p o s t e r i o r margin and  rectangular  n o d i f o r m cusp.F87. (X250). F i g s . 7-9  Neogondolella serrata(?)  7. O r a l view o f i n t e r m e d i a t e t o mature form.  Page Note t h e d e g e n e r a t e  o v e r a l l appearance.F73. (X300). 8. Close-up o f p o s t e r i o r end o f F i g . 7. F73.(X600). 9. O b l i q u e - l a t e r a l t o o r a l view o f i n t e r m e d i a t e form. s h a r p n e s s of a l l t h e d e n t i c l e s . F 6 3 . ( X 3 0 0 ) .  Note t h e  90  125  Explanation f o r Plate 7 A l l f i g u r e s a r e Scanning E l e c t r o n M i c r o g r a p h s . F i g s . 1-8  N e o g o n d o l e l l a b i t t e r i n.subsp. C  Page  95  1. A b o r a l v i e w o f m i d - p l a t f o r m fragment.F47. (X125). 2. A b o r a l v i e w o f p o s t e r i o r end.F47. (X125). 3. O r a l v i e w o f mature form w i t h asymmetric  p o s t e r i o r end and  large  4. O r a l v i e w o f mature form w i t h asymmetric p o s t e r i o r end and  large  brim.F47. (X250).  brim.F47.(X250). 5. O r a l view.  Note t h e l a r g e , c i r c u l a r cusp s i m i l a r t o N. n.sp. IS  but a l s o t h e d i s t i n c t asymmetry. F47. (X250). 6. O r a l v i e w . F 9 6 . ( X l O O ) . 7. O r a l v i e w showing p a r t i a l f u s i o n o f cusp and p o s t e r i o r - m o s t and r e t i c u l a t e ornament  denticle  on t h e c a r i n a . F96.(XlOO).  8. O r a l v i e w of. mature form showing a s l i g h t l y rounded and  asymmetric  p o s t e r i o r end t h a t i s s i m i l a r to the more b l u n t ended N. r o s e n k r a n t z i . F 9 6 . ( X 1 5 0 ) F i g s . 9-12  N e o g o n d o l e l l a r o s e n k r a n t z i n.subsp. D  Page  9. O r a l v i e w o f mature form showing the s t r a i g h t , asymmetric ornament  p o s t e r i o r end as compared  to _N. b i t t e r i .  less  98  distinctly  Note t h e r e t i c u l a t e d  on the c a r i n a . F 9 6 . ( X l O O ) .  10. O r a l view o f mature  form w i t h a v e r y t h i c k p l a t f o r m ,  elongate-  o v a l cusp d i r e c t e d p o s t e r o - l a t e r a l l y and w i t h f u r r o w s and c a r i n a t h a t a r e almost e n t i r e l y c o v e r e d w i t h r e t i c u l a t e  microornament.F96.(XlOO).  11. A b o r a l view of p l a t f o r m showing a wide t r i a n g u l a r , and  asymmetric  loop.F96.(X85). 12. O r a l v i e w o f mature t o g e r o n t i c form showing t h e p o s t e r o - l a t e r a l l y d i r e c t e d f u r r o w s and s w o l l e n p o s t e r i o r p l a t f o r m margins.F96.(X140) .  Plate 7  126  127  Explanation f o r Plate 8 All  f i g u r e s a r e Scanning E l e c t r o n M i c r o g r a p h s .  F i g s . 1-13  N e o g o n d o l e l l a r o s e n k r a n t z i n.subsp. _D  Page  1. O r a l v i e w of i n t e r m e d i a t e form t h a t l o o k s v e r y s i m i l a r  to N.  98 ida-  h o e n s i s ^ F96.(X85). 2. L a t e r a l v i e w of p o s t e r i o r end of a mature t o g e r o n t i c l a r g e brim, f u s e d c a r i n a and downturned  form showing  l a t e r a l margins.F96.(X100) .  3. O r a l v i e w of mature form showing t h e p o s t e r o - l a t e r a l l y furrows and a s l i g h t t w i s t o f t h e p l a t f o r m a t t h e a n t e r i o r  directed  end.F96-(X85).  4. O r a l v i e w o f mature form w i t h narrow b r i m and wide but s h a l l o w furrows.F96.(X100). 5. L a t e r a l view o f i n t e r m e d i a t e form.F96.(X85). 6. L a t e r a l view o f a n t e r i o r end showing t h e k e e l - l i k e c a r i n a  (owing  to f u s i o n o f d e n t i c l e s ) and l a c k of p l a t f o r m a t a n t e r i o r - m o s t end: both f e a t u r e s a r e c h a r a c t e r i s t i c f o r mature  to g e r o n t i c i n d i v i d u a l s o f t h i s  subspecies.F96.(X85). 7. L a t e r a l view o f i n t e r m e d i a t e form.F96. (.X85) . 8. L a t e r a l view o f F i g . 3 showing k e e l - l i k e a n t e r i o r 9. O b l i q u e l a t e r a l view of j u v e n i l e form.  carina.F96.(X75).  Note t h e s i m i l a r i t y to  j u v e n i l e s o f N. i d a h o e n s i s n. subsp. A ( P l . 2, F i g . 16).F96.(X225). 10. O r a l view o f g e r o n t i c d e n t i c l e ( s ) f u s e d and d i r e c t e d  i n d i v i d u a l w i t h cusp and  posterior-most  postero-laterally.F96(X85).  11. C l o s e - u p o f o r d e r e d r e t i c u l a t e d mieroprnament  and  flat-topped,  smooth d e n t i c l e . F 9 6 . ( X 9 0 0 ) . 12. C l o s e - u p o f g e r o n t i c o r a l s u r f a c e showing the almost.complete l a c k o f furrows.' surface.  R e t i c u l a t e d microornament c o v e r s almost t h e e n t i r e  Note t h e e l o n g a t e form o f r e t i c u l a t e d ornament  are normally positioned.F96.(X200) . 13. A b o r a l s u r f a c e of i n t e r m e d i a t e  form.F96.(X100).  oral  where the f u r r o w s  APPENDIX I Schematic of morphological terminology and measured parameters f o r Neogondolella and measurements (Lj_, L , H i , Wi, W : 2  2  a l l i n ym) o f specimens  from F48 - F54..  aboral view Posterior Parameter Measurements for Sample F48.  ecimen  L  l  L  2  H  V*l  *1  W  2 1Q  4  1  io6a  28Q  17Q  6.24- 12  88.33  4.42  240  180  5.88  2  69.0  38Q  150  4.6Q  8  86.25  4.31  160  140  5.70  3  10.20. 320  140  7.29  11  92.73  5.10  200  170  5.92  4  110Q  30Q  220.  5.QQ  11 100.00  4.58  240  220  6.90  5  630  260  110  5.73  10  63. Q0  4.85  130  120  3.25  6  600  24 Q  140  4.29  8  75.00  4.62  130  110  2.88  7  720  330  120  6.00  9  80.00  4.80  150  130  4.62  8  580  250  150.  3.87  9  64.44  4.14  140  100  3.00  9  1100  300  160  6.88  13  84.62  4.58  240  200  6.60  10  660  310  140  4.71  9  73.33  3.67  180  150  5.12  130  Parameter Measurements f o r Sample F49  ecimen  L  l  L  2  H  V 1 W  W  l  W  2 .  h\  (w+w 1  io  4  1  800  24 Q  140.  5.71  11  72.73  4.Q0  20Q  150  2  1400  380  280  5.00  13 107.69  4.83  29.0  280 :  3  780  250  170.  4.59  12  65.00  3.90  20Q  200  5.00  4  700  24 Q  160  4.38  10  70.00  4.12  170  130  3.60  5  1040  340  160  6.50  9. 115.56  4.00  260  200  7.82  6  700  240  140  5.00  9  77.78  3.89  180  140  3.84  7  8 60  280  180  4.78  12  71.67  4.53  190  190  5.32  8  600  230  150  4.00  9  66.67  4.00  150  110  2.99  9  760  260  190  4.00  10  76.00  4.47  170  140  4.03  10  740  310  190  3.89  10  74.00  4.63  160  150  4.81  11  1020  310  180  5.67  9 113.33  4.43  230  220  6.98  12  1220  340  240  5.08  15  81.33  4.57  270  240  8.67  13  840  240  160  5.25  11  76.36  5.25  160  130  3.48  14  460  220  120  3.83  8  140  120  2.86  15  900  240  160  5. 63  12  75.00  4.74  190  170  4.32  16  760  260  160  4.75  9  84.44  4.22  180  160  4.42  17  760  240. 160  4.75  12  63.33  4.00  190  150  4.08  18  800  230  140  5.71  11  72.73  4.00  200  180  4.37  19  1000  340  200  5.00  12  83.33  4.76  210  160 ?  6.29  20  860  320  200  4.30  10  86.00  4.53  190  160  5.60  21  860  210  10Q- .8.60  13  66.15  5.06  170  140  3.26  22  820  32Q  170  4.82  10  82.00  4.10  200  180  6.08  23  640  300  140  4.57  9. 71.11  4.00  160  140  4.50  24  1060  360  220  4.82  12  88.33  4i42  240  210  8.10  25  1Q2Q  320  14Q  7.29  12  84.00.  4.25  240  220'  7.36  26  1180  28Q  19Q  6.21.  13  90.77  5.36  22Q  180  5.60  980  350  23Q  4.26  11  89.. 09. 4.45  220  200:;  7.35  1100  290  200.  5.50  13  84.62  5.00  220  20.0  6.09  29:  800  330  220.  3.64  10.  80.00  4.Q0  2Q0  170  6.11  30  600  280  140  4.29  9  66.67  4.29.  140  120:  3.64  27: 28  47.40./ 3.29.  4.20 10.83  131  Parameter Measurements f o r Sample F49 (cont.)  lecimen  L  l  L  2  H  V1 W  W  l  W  2  % L (Wj+V 1  ID  4  31  980  340  190  5.16  12  81.67  5.44  180  170  5.95  32  820  280  18Q  4.56  10  82.00  4.32  190  170  5.04  33  960  27Q  160.  6.QQ  12  80.00  4.36  220  170.  5.27  34  1060  300  19Q  5.58  12  88.33  4.42  240  190  6.45  35  720  240  160  4.50  10  72. 00  5.14  140  120  3.12  36  820  310  150  5.45  10  82.00  4.10  200  160  5.58  37  780  280  160  4.88  12  65.00  4.88  160  15Q  4.34  38  900  270  140  6.43  10  90.00  5.00  180  160  4.59  39  920  240  16Q  5.75  13  70.77  5.11  180  170 ;  4.20  40  1000  260  180  5.56  12  83.33  5.56  180  160  4.42  41  880  260  170  5.18  11  80.00  4.63  190  180  4.81  42  940  240  190.  4.95  12  78.33  4.70  200  170  4.44  43  760  240  120  6.33  11  69.09  4.00  190  160  4.20  44  880  280  190  4.63  12  73.33  4.40  200  180  5.32  45  520  250  140  3.71  8  65.00  3.47  150  120  3.38  46  680  280  140  4.86  8  85.00  4.25  160  130  4.06  47  980  260  170  5.76  12  81.67  3.92  250  200  5.85  48  660  230  140  4.71  9  73.33  4.40  150  140  3.34  49  600  240  120  5.00  10  60.00  4.62  130  110  2.88  50  520  220  70  7.43  9  57.78  3.47  150  120  2.97  51  720  280  120  6.00  10  72.00  4.24  170  140  4.34  52  900  300  150  6.00  11  81.82  4.50  200  160.  5.40  53  720  260  130  5.54  11  65.45  4.50  160  140  3.90  54  460  290  1Q0.  4. 60  7  65.71  3.29  140  100  3.48  55  820  250  140  5.86  11  74.55  4.82  170  140  3.88  56  780  260  120  6.50. 11  7Q.9.1  4.33  180  140  4.16  57  1100  340.  18Q  6.11  100.00.  5.24  210  180  6.63  58  980  320  100  9.80. . 11  89.0.9  5.16  • 190 160  5.60  59  7 60  300  140  5.43  9  84.44  4.22  180  130  4.65  320  180.  6.22  14  80.00' 5.09  220  19.0  6.56  60  1120  11  132  Parameter Measurements f o r Sample F52  Specimen  L  l  L  2  H  V*l  W  l  F  J2L (W + , 1  1  2  1  io  4  1  1120  28Q  19.0  5.89  12  93.33  4.31  260  230  6.86  2  1190  300  190  6.26  12  99.17  5.95  200  180  5.70  3  860  290  160.  5.38  11  78.18  4.10  210  180  5.66  4  600  240  110  5.45  9  66.67  3.75  160  140  3.60  5  720  260  170  4.24  10  72.00  3.79  190  180  4.81  6  760  220  100  7.60.  11  69.09  4.00  190  170  3.96  7  690  280  150  4.60  10  69.00  4.93  140  130  3.78  8  570  270  140  4.07  8  71.25  4.07  140  110  3.38  9  700  280  150  4.67  9.  77.78  4.38  160  120  3.92  10  880  300  200  4.40. 11  80.00  4.19  210  170  5.70  11  800  240  150  5.33  11  72.73  5.71  140  120  3.12  12 .  740  29.0  140  5.29  10  74.00  4.11  180  160  4.93  13  700  270  150  4.67  10  70.00  3.68  190  160  4.73  14  1000  270  140  7.14  12  83.33  4.55  220 ; 180  5.40  15  1100  380  240  4.58  13  84.62  3.93  280  240  9.88  16  610  260  120  5.08  9  67.78  3.59  170  150.>  4.16  17  800  290  140  5.71  10  80.00  4.21  190  140  4.79  18  780  280  140  5.57  11  70.91  4.59  170  140  4.34  19  960  260  160  6.00  13  73.85  4.57  210  190  5.20  20  820  250  160  5.13  12  68.33  4.32  190  170  4.50  21  560  270  110  5.09  8  70.00  3.73  150  130  3.78  22  750  230  150  5.0Q  10  75.00  3.95  190  180  5.18  23  1200  410  300  4.Q0  13  92.31  4.00  300  290  12.10  24  1100  340  200  5.5Q  12  91.67  3.79  290  260  9.35  25  1170  330  220.  5.32  13  90.  4.50  260  240  8.25  oa  133  Paramet er  2Cimen  L  l  L  2  H.  y H  .ts f o r Sample F53  1  760  220  160  4.75  11  69.09  4 .22  180  150-  3.63  2  1180  340  250  4.72  13  9Q.77  4 .07  290  280  9.69  3  1160  320  220  5.27  . 15  77.33  4 .46  260  220  7.69  4 .  1270  300  210  6.05  16  79.38  5 .29  240  200 .  6.60  5  1310  320  190  6.89  14  93.57  5 .04  260  260  8.32  6  1120  320. 180  6.22  13  86.15  4 .31  260  240  8.00  7  980  340  190  5.16  11  89.09  4 .45  220  180  6.80  8  580  280  130  4.46  9  64.44  3 .63  160  130  4.06  9  720  320  180  4.00  10  72.00  4 .00  180  160  5.44  10  700  300  160  4.38  9  77.78  3 .68  190  180  5.55  11  980  280  140  7.00  13  75.38  4 .45  220  180  5.60  12  1500  380  17Q  8.82  16  93.75  3 .95  380  310  13.11  13  960  29Q  180  5.33  11  87.27  4 .00  240  200  6.38  14  860  320  160  5.38  10  86.00  4 .30  200  160  5.76  15  640  260  130  4.92  9  71.11  3 .56  180  140  4.16  16  1100  400  170  6.47  11  100.00  4 .23  260  200  9.20  17  920  350  180  5.11  10  92.00  4 .18  220  180  7.00  18  620  280  140  4.43  9  68.89  3 .65  170  140  4.34  19  670  280  160  4.19  9  74.44  4 .47  150  130  3.92  20  740  240  150  4.93  10  74.00  4 .63  160  140  3.60  21  700  240  130  5.38  10  70.00  4 .38  160  140  3.60  22  500  260  120  4.17  8  62.50  3 .57  140  120  3.38  23  800  300  200  4.0Q  10  80.00  3 .64  220  200  6.30  24  12Q0  40Q  190  6.32  12  100.00  4 .14  290  280  11.40  25  840  29.0. 160  5.25  11  76.36  4 .67  180  140  4.64  26  700  280  130.  5.38  10  70.00.  4 .12  17Q  120  4.06  27  660  280  120 ., 5.50.  10.  66.00. 3 .67  180  160  4.76  28  980  360  180  5.44  11  89. Q9  4 .45  220  180  7.20  29  580  240  120  4.83  9. '.. 64.. 44 4 .14  140  120  3.12  30  760  340  150. 5.07  160  120  4 .76  9  84.44  4 .75  134  Parameter Measurements f o r Sample F53  Specimen  L  L  2  H  I^/II  //  1^/// L /W  \  (cont.)  W  2  % L O^+wp io  4  31  840  260  120  7.00  11  76.36  4.20  200  180  4.94  32  78Q  300  160  4.88  10  78.00  4.33  180  160  5.10  33  102Q:  260  220  4.64  12 : 85.00  5.10  200  180  4.94  34  950  320  230  2.97  12 :  79.17  4.32  220  180  6.40  35  700  260  140  5.00  10  70.00.  4.38  160  120  3.64  36  1010  280  18Q  5.61  13  77.69  4.81  210  180  5.46  37  1160  400  250  4.64  12  96.67  4.14  280  250  10.60  38  380  220  100.  3.80  7  54.29  3.45  110  80  2.09  39  690  2 60  160 .4.31  10  69.00  3.83  180  170  4.55  40  1500  440  320  15  100.00  4.41  340  320  14.52  4.69  135 Parameter Heasurements f o r Sample F54 scimen 1  .  • l L  #  y  #  Specimen  L  l  //  600  9  66.67  201  580  9  64.44  1Q00  15  66.67  21  740,  10.  74.00  3  720  9  80.00  22  780  10  78.00  4  1080  11  98.18  23  700  10  70.00  5  1120  11  101.82 :  24  . 740  9  82.22  6  106Q  11  96.36  25  900  11  81.82  7  10Q0  12-  83.33  26  1000  13  76.92  8  1220  13  93.85  27  540  8  67.50  9  1460  13  112.31  28  1220  14  87.14  2 :  10  9.0  Q  10  90. 00  29  3 60  6  60.00  11  840  11  76.36  30  900  13  69.23  12  960  11  87.27  31  1300  13  100.00  13  800.  10.  80. 00  32  1360  12  112.50  14  56Q  11  50.91  33  560  12  4 6.67  15  840  11  76.36  34  1020  11  92.73  16  720  9  80.00  35  1200  13  92.31  17  960  10  9.6.00  36  1280  13  98.46  18  1040  11  94.55  37  800  10  80.00  19  640  9  71.11  

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