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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 University, 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' ... -i n THE FACULTY OF GRADUATE STUDIES The Department of Geological Sciences We accept t h i s t hesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA October 1981 Charles Murray Henderson, 1981 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y a v a i l a b l e for reference and study. I further agree that permission for extensive copying of t h i s thesis for s c h o l a r l y purposes may be granted by the head of my department or by his or her representatives. It i s understood that copying or p u b l i c a t i o n of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. D e p a r t m e n t of Geological Sciences The University of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date October 2, 1981 i i ABSTRACT A. succession of l a t e Early through Medial Permian conodont faunas i s documented for the f i r s t time from the calcareous, f i n e grained, quartzose sandstones of the Assistance and Trold Fiord Formations on northern E l l e s -mere Island, Northwest T e r r i t o r i e s . Of the taxa i d e n t i f i e d and described, one species and three subspecies are proposed as new. The taxa include, i n chronological order: Neogondolella idahoensis subsp. indet., Neostrep-tognathodus p r a y i , Neogondolella idahoensis n.subsp. A, Anchignathodus  minutus, Neogondolella s e r r a t a ( ? ) , N. n.sp. B, N. postserrata(?), N. b i t t e r i n.subsp. C, and N. rosenkrantzi n.subsp. D. Numerous ramiform elements are also associated with Neogondolella idahoensis n.subsp. A. These elements may comprise part of a multielement Neogondolella apparatus or they may represent separate form species. In observation of t h e i r questionable status, a somewhat uns a t i s f a c t o r y dual taxonomy i s proposed f o r these elements, and includes the following taxa: N. idahoensis n.subsp. A - Xanio-gnathus t o r t i l i s , N. idahoensis n.subsp. A - E l l i s o n i a excavata, N. idahoen-s i s n.subsp. A - E l l i s o n i a t r i b u l o s a , and N. idahoensis n.subsp. A - P r i -o n i o d e l l a decrescens. S t a t i s t i c a l work on the abundant platform elements of N. idahoensis n.subsp. A provides evidence f o r minor 'evolutionary trends of increasing siz e and increasing number of d e n t i c l e s upsection. Comparison of these con-odonts with N. serrata.and N. postserrata from the Great Basin of SW USA suggests that the phylogenetic development of Permian Neogondolella f o l -lowed an,, evolutionary path.more appropriate to punctuated e q u i l i b r i a than to p h y l e t i c gradualism. The conodont taxa i n d i c a t e that the Assistance Formation i s Upper Leonardian to Uppermost Roadian i n age whereas the Trold Fiord Formation includes most of the Wordian.stage. These two formations have been separ-ated into f i v e subdivisions on the basis of both lithology. and the presence or absence of various biota 1 A sixth, subdivision i s described f o r the Sa-bine Bay Formation which underlies the Assistance and where conodonts are apparently absent. L i t h o l o g i c and b i o t i c evidence (including trace f o s s i l s and mega- and microbiota) point to shallow, offshore marine conditions well within the photic zone and characterized by low energy and slow depositional rates, for most of the conodont bearing s t r a t a . A much thicker c o r r e l a t i v e sec-t i o n to the south represents, i n large part, a de l t a front sequence. The Sabine Bay Formation, on the other hand, i s composed of shoreface sandstones, possibly i n a b a r r i e r i s l a n d s e t t i n g . The r e s u l t s of t h i s research ind i c a t e that conodonts may be very promising f o r c o r r e l a t i o n of Permian s t r a t a i n the Canadian A r c t i c A r c h i -pelago and f o r worldwide comparison. More work within the Sverdrup Basin, including both marginal and basinal sections , i s necessary to provide a good biozonation of these marine Permian s t r a t a . The use of the abundant brachiopods i n combination with the conodonts i s probably the best way to resolve t h i s zonation. The taxonomic.descriptions and subdivisions pro-posed herein should provide a foundation f o r future work. i v TABLE OF CONTENTS T i t l e page i Abstract i i Table of Contents i v L i s t of Tables v i i L i s t of Figures v i i i L i s t of Plates i x Acknowledgements x Introduction 1 Location and Scope of the Study 1 F i e l d Work 3 Previous Work 4 Laboratory and A n a l y t i c a l Methods. 6 Stratigraphy and Paleoenvironment - General Statement 10 Sabine Bay Formation 10 Hamilton Peninsula area 10 McKinley Bay area 14 Tanquary Fiord area 14 Assistance Formation 15 Hamilton Peninsula area 15 McKinley Bay area 16 Sawtooth Range area ". 17 Trold Fiord Formation 18 Hamilton Peninsula area 18 McKinley. Bay area 21 Sawtooth Range area 22 V Age and Co r r e l a t i o n . . . 23 Relative value of various f o s s i l b i o t a . . . . . . . . ":."25 Previous 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 29 Assistance Formation - 30 Trold Fiord Formation.... 30 Age assignments r e s u l t i n g from t h i s work 30 Subdivision A.... 31 Subdivision B 32 Subdivision C 34 Subdivision D. 36 Subdivision E 37 Subdivision F 38 Summary 40 Quantitative Analysis of Measurable characters for Neogondolella 40 Introduction 40 Results and Discussion 45 Overall length of platform (Ll) 46 Number of d e n t i c l e s per element (#) 54 Ratio of length to number of d e n t i c l e s (Ll/#) 54 Length from.'.tip:'."of cusp to fourth d e n t i c l e anterior (L2) . ... 57 Maximum width (Wl) 57 Width?.at posterior end (W2) 57 Height from t i p of cusp to base of the flange. (HI) 57 Ll/Wl r a t i o . '"59 v i ,L1/HI r a t i o 60 Posterior area 60 Discussion of Evolutionary trends and concepts 61 Systematic Paleontology 69 Introduction 69 Anchignathodus .minutus. 73 Neostreptognathodus p r a y i 73 Neogondolella idahoensis subsp. iridet 74 Neogondolella idahoensis n.subsp. A 75 Neogondolella idahoensis n.subsp. A v a r , g r a c i l i s 84 Neogondolella idahoensis n.subsp. A var. robustus 85 Neogondolella idahoensis n.subsp. A var. intermediatus 86 Neogondolella idahoensis n.subsp. A v a r , constrictus 86 Neogondolella idahoensis n. subsp. A var. lobatus 87 Neogondolella serrata( ?.) . . . 90 Neogondolella n.sp. B_ 92 Neogondolella postserrata(?) 94 Neogondolella b i t t e r i n.subsp. Cj 95 Neogondolella rosenkrantzi n.subsp. I). 98 Neogondolella idahoensis n.subsp. A - Xaniognathus t o r t i l i s 103 Neogondolella idahoensis n.subsp. A - E l l i s o n i a excavata 104 Neogondolella idahoensis n.subsp. A - E l l i s o n i a . t r i b u l o s a 104 Neogondolella idahoensis n.subsp. A - P r i o n i o d e l l a decrescens.... 105 References....... 106 Plates 1 - 8 113 Appendix,!-. ••• 129-135 v i i LIST OF TABLES Table 1. S t a t i s t i c s from data given i n Appendix I.... 47 Table 2. Values derived from z-tests f o r L l , # a i*d Ll/# 49 Table 3. Counts and percentages of platforms and ramiforms.of Neogondolella i n F48 - F54. 51 Table 4. S t a t i s t i c s from sample subsets (elements with 10 and 11 denticles) 52 Table 5. Values derived from t - t e s t s on sample subsets 52 v i i i LIST OF FIGURES Figure 1. Map showing locati o n s of sections studied 2 Figure 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 various shallow marine environments 11 Figure 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 c o r r e l a t i o n of formations and sections. 12 Figure 4. Correlation chart f o r s e r i e s , stages and zones of the Permian 24 Figure 5. Frequency d i s t r i b u t i o n of number of d e n t i c l e s and platform length f o r Neogondolella idahoensis n.subsp. A 43 Figure 6. Graphical representation of part of Table 1 48 Figure 7. Graphs showing r e l a t i o n s h i p of the platform length to (a,b,c) number of d e n t i c l e s per element f or F49, F54 and F96 53 Figure 8. Graph showing r e l a t i o n s h i p of the platform length to num-ber of d e n t i c l e s per element of N. idahoensis n.subsp. A, !N. serrata, and _N. postserrata ( i n chronological order)... 56 Figure 9. Graphical representation of part of(Table 1 58 Figure 10. Graph showing r e l a t i o n s h i p of Length/Height to Length of platform f o r F49 7 9 Figure 11. Graph showing r e l a t i o n s h i p of .Length/Width to Length of platform f o r F49 80 Figure 12. Graph showing r e l a t i o n s h i p of Posterior Area to Length of platform for.F53 81 Figure 13. Graph showing r e l a t i o n s h i p of Posterior Area to Length of platform f o r F49 82 Figure 14. Graph showing r e l a t i o n s h i p of Posterior Area to Length/ number of d e n t i c l e s f o r F49 83 LIST OF PLATES A l l f igures on plates are Scanning Electron Micrographs. Plate 1. Neostreptognathodus p r a y i , Anchignathodus minutus, Neo-gondolella idahoensis subsp.indet., Neogondo1ella idahoen- s i s n.subsp. A - P r i o n i o d e l l a decrescens. and Neogondolella  idahoensis n. subsp. A - E l l i s o n i a excavata. 113 Plate 2. Neogondolella idahoensis n.subsp. A - E l l i s o n i a t r i b u l o s a , Neogondolella idahoensis n.subsp. A - Xaniognathus t o r t i l i s , and Neogondolella idahoensis n.subsp. A 115 Plate 3. Neogondolella. idahoensis n.subsp.. A 117 Plate 4. Neogondolella idahoensis n.subsp.. A 119 Plate 5. Neogondolella.. idahoensis n.subsp. A 121 Plate 6. Neogondolella n.sp. IS, Neogondolella postserrataQ?), and Neo-gondolella serrata( ?) 123 Plate 7. Neogondolella b i t t e r i n.subsp. Cj and Neogondolella rosenkrantzi n.subsp. I) 125 Plate 8. Neogondolella rosenkrantzi n.subsp. I) 127 X ACKNOWLEDGEMENTS There are a number of people to whom I became indebted during the course of t h i s research. F i r s t of a l l , I would l i k e to thank my advisor, Dr. R.V.. Best, for his encouragement and patient 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 t h e s i s . His contribution has improved immensely the q u a l i t y of the f i n a l manuscript. I would also l i k e to express my gratitude to the other members of my com-mittee: Drs. G.E. Rouse and P.L. Smith of UBC and Dr. M. Orchard, of the Geological Survey of Canada, Vancouver. In p a r t i c u l a r , I would l i k e to acknowledge Mike Orchard, a fellow conodont worker, for the f r u i t f u l d i s -cussions we had with regards to conodonts and t h e i r taxonomic problems. I g r a t e f u l l y thank the Geological Survey of Canada i n Calgary for the f i n a n c i a l and l o g i s t i c a l support during the f i e l d season of 1979. In par-t i c u l a r , I would l i k e to acknowledge W.W. Nassichuk, A. Embry, and U. Mayr for i n i t i a l l y suggesting the project and l a t e r for providing support and u s e f u l discussions. Foremost, I wish to express my deepest appreciation to my wife, Betty, who not only typed part of the manuscript but also served as an endless source of encouragement and endured my many l a t e evenings spent completing t h i s t h e s i s . F i n a l l y , I would l i k e to dedicate t h i s thesis to the l a t e Dr. David G. Perry. Dave was the o r i g i n a l advisor to my t h e s i s and had joined me on Ellesmere Island for part of the c o l l e c t i n g i n early July, 1979. His pro-mising l i f e was cut short by a helicopter crash before my f i e l d season had even ended i n August, 1979. His death:.was a t r a g i c and b i t t e r blow but h i s memory provided me with the i n s p i r a t i o n and motivation to continue and com-plete t h i s research. 1 INTRODUCTION This t h e s i s records the r e s u l t s of a study of the conodont b i o s t r a t -igraphy of the Permian Sabine Bay, Assistance and Trold Fiord Formations of northern Ellesmere Island, N.W.T. To a les s e r extent brachiopods and l i t h o f a c i e s were u t i l i z e d to e s t a b l i s h c o r r e l a t i o n s . Location and Scope of the Study Ellesmere Island, the most northerly island of the Canadian A r c t i c Archipelago, i s located between 76oahd 83°North l a t i t u d e . The study i s based p r i m a r i l y on seven sections from four d i f f e r e n t areas including two from Hamilton Peninsula (80°10' N, 081°45' W), two from McKinley Bay (81°10' N, 079°10'W), two from the headof Tanquary Fiord (81°25' N, 076°30' W) and one from the Sawtooth Range (79°30' N, 083°20' W) (sections A, B, C, and D re s p e c t i v e l y on F i g . 1). Other sections have been studied i n minor d e t a i l ( F ig. 1) and"are only referred to where they proved valuable as support f o r any in t e r p r e t a t i o n s . The l i t h o l o g y studied i n the above mentioned sections represent the marginal f a c i e s for the Permian part of the Sverdrup Basin; a basin of deposition from early Carboniferous to T e r t i a r y . Despite::the d e s c r i p t i v e and reconnaissance studies by previous workers (Thorsteinsson, 1974; C h r i s t i e , 1964; Nassichuk and C h r i s t i e , 1969; and Mayr, 1976) the Permian part of the basin remains the poorest understood of the Phanerozoic systems. F o s s i l c o l l e c t i o n s and age determinations have been previously reported from the Sabine Bay, Assistance and Trold Fiord Formations by Nassichuk et a l . (1965), Harker and Thorsteinsson (1960), Nassichuk (1970), Nassichuk and Spinosa (1970) and by J.B. Waterhouse and R.E. Grant i n Thorsteinsson (1974). The material reported on herein constitutes the f i r s t systematic c o l l e c t i o n s through the 2 Figure 1. Map of northern Ellesmere Island, showing location of the sections studied. 3 complete sections: the previous reports were l a r g e l y of i s o l a t e d occurrences. These previous studies emphasized ammonoids and brachiopods to f a c i l i t a t e c o r r e l a t i o n . Both of these groups have t h e i r own peculiar problems asso-ciated with them (see p. 25) that hinder c o r r e l a t i o n schemes. This report emphasizes the use of conodonts, a group whose once many problems hindering c o r r e l a t i o n have been l a r g e l y ironed out by intensive research over the past f i v e years (Clark and Behnken, 1979; Clark et a l . , 1979 and Wardlaw and Collinson, 19 79b). It i s because of the 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 short distances, and the presence of disconformities and transgressive u n i t s that the c o r r e l a t i o n and environ-mental r e l a t i o n s h i p s of the formations pertaining to t h i s report are poorly understood. The o r i g i n a l i n t e n t i o n f or the research was to better define the age and c o r r e l a t i o n of these formations. This seems to have been accomplished through the use of conodonts although the r e s u l t s should only be regarded as a beginning, but a s t a r t that at 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 area i s by Twin Otter or DC-3 a i r c r a f t from Res-olute Bay, Cornwallis Island, to Eureka or Tanquary Fiord a i r s t r i p s . From these bases access to the section l o c a l i t i e s was accomplished through the use of Jet Ranger he l i c o p t e r s . The f i e l d work was completed during three weeks between June 16 and August 11, 1979. The section d e s c r i p t i o n was completed with the aid of an assistant from f l y camps c o n s i s t i n g of a logan and a pyramid tent at each of the l o c a l i t i e s . Radio contact was maintained at regular times with the main base at Eureka or Tanquary Fiord to report weather, and to ind i c a t e move dates and supplies required. The weather through the period indicated above was a, mixture of sun and cloud and included only two weather rel a t e d down-days. This f i f t y -seven day period saw three days with snow f l u r r i e s and nine days of showers or r a i n . During t h i s e n t i r e period the sections studied were free of i c e and snow with the exception of the minor f l u r r i e s . Otherwise sunshine was the order of the day with temperatures reaching as high as 19oC (July 30) but more t y p i c a l l y averaged 3 to 10°C. Daytime temperature f l u c t u a t i o n s were minor as at t h i s l a t i t u d e the sun remains above the horizon from A p r i l 15 to August 29 (Thorsteinsson, 1974). Part of the camp remained at Tanquary Fiord a f t e r the IT1"*1 of August but snow began to f a l l on the 1 2 t h and camp was folded for the season by the 16 when no break was i n sight. Previous Work The summary of previous work i n the area as presented herein, and e s p e c i a l l y of the early h i s t o r y , i s l a r g e l y taken from R.L. C h r i s t i e ' s Geological Survey of Canada (GSC) Memoir 331 (1964) and to whom the c r e d i t i s due. The h i s t o r y of exploration and geological i n v e s t i g a t i o n of northern Ellesmere Island i s a very auspicious and c o l o u r f u l one. The f i r s t geolo-g i c a l studies were by a B r i t i s h explorer, Captain S i r George Nares, on a 1875-76 Royal Navy expedition to Lady Franklin Bay. Captain H.W. Feilden, a n a t u r a l i s t on Nares' expedition, and others made extensive c o l l e c t i o n s of rocks and f o s s i l s i n the region between Discovery Harbour and Feilden Peninsula. Lieutenant Adolphus W. Greely of the .U.S. Army established Fort Conger i n Discovery Harbour i n 1881. Expeditions went to Lake Hazen and Greely Fiord during 1882-83 where geological and archaeological specimens were c o l l e c t e d and copious notes made. This success was tainted by the t r a -gi c end of the expedition where a l l but seven men died of starvation because 5 a planned rendesvous with a return v e s s e l was l a t e . The area was explored by a number of others over the next seventy years including Commander R.E. Peary (1898-1909), a geologist W. Elmer Ekblaw (1913-17) who c o l l e c t e d Permian f u s u l i n i d s from near the mouth of Tanquary Fiord, and a geologist Dr. J.C. Troelsen (1939-40). The f i r s t appearance of the GSC was i n 1948 by V.K. Prest along the northeast shore of Ellesmere. G. Hattersley-Smith (Defence Research Board) and R.G. Blackadar (GSC) l a t e r conducted geological reconnaissance i n the Lake Hazen area. R.L. C h r i s t i e of the GSC conducted f i e l d work i n 1954, 1957, and 1958 i n northeast Ellesmere producing a map i n h i s GSC Memoir 331. In 1956 and 1957 R. Thorsteinsson and E.T. Tozer investigated western Ellesmere Island. This work and much of the previous work was conducted by means of dog teams and canoe over extended f i e l d seasons. In 1961 and 1962 Operation Eureka, under the d i r e c t i o n of R. Thorsteinsson of the GSC, i n -cluded J . Wm. Kerr, E.T. Tozer, and H.P. T r e t t i n . During t h i s period transportation included Piper Super Cub a i r c r a f t and a G2A h e l i c o p t e r . In 1963 R. Thorsteinsson and P. Harker conducted further s t r a t i g r a p h i c studies and mapping of Ellesmere. These f i v e f i e l d seasons are the founda-r. ' . t i o n f o r Thorsteinsson's.GSC B u l l e t i n 224 (1974) which remains today as the major work on Carboniferous and Permian stratigraphy i n the area. Previous f o s s i l work was l a r g e l y on brachiopods and ammonoids as indicated i n the f i r s t part of t h i s chapter. However, l a t e Lower Permian through Middle Permian conodonts from the area have only been reported once previously (Kozur and Nassichuk, 1977) and t h i s was of j u s t two c o l -l e c t i o n s (see p. 28 for d e t a i l s ) . The designation of the three formations of t h i s report date between 1960 and 1974. The Sabine Bay Formation was named by Tozer and Thorsteins-6 son (1964) for a.section on Sabine Peninsula, M e l v i l l e Island. The A s s i s -tance Formation was named and defined by Harker and Thorsteinsson (1960) for a succession on G r i n n e l l Peninsula, Devon Island. F i n a l l y , the Trold Fiord Formation was defined by Thorsteinsson (1974) and includes a type section on a small, unnamed t r i b u t a r y of the East Cape River that issues into the northeast side of Canon Fiord on the west coast of Ellesmere (very near the Hamilton Peninsula . sections of t h i s r e p o r t ) . The most precise way to summarize the previous work i n the area on Carboniferous and Permian rocks i s to say that the reconnaissance has been completed but that d e t a i l e d studies are merely beginning. Laboratory and A n a l y t i c a l Methods Laboratory work was conducted from the f a l l of 1979 to the spring of 1981. A standard technique of a c e t i c acid d i s s o l u t i o n , wet sie v i n g , and heavy l i -quid separation (tetrabromoethane) was used to concentrate the conodonts from t h e i r host rocks. The bulk samples that were processed were of two types. The f i r s t con-s i s t e d of large s i n g l e blocks or a number of moderate sized slabs that weighed up to 25 kg (55 lbs.) but more t y p i c a l l y averaged 10 kg (22 l b s . ) . These blocks were c o l l e c t e d f or t h e i r f i n e l y s i l i c i f i e d brachiopod content which were to have been the major emphasis of the research (the emphasis switched to conodonts about half-way through the processing). The second type consisted of 2 to 3 cm diameter chips c o l l e c t e d from s i n g l e horizons s p e c i f i c a l l y for conodonts and weighing between 3.2 and 4.2 kg (7 to 9 lbs.) i n t o t a l . The large blocks were broken into two f r a c t i o n s . Small fragments were broken o f f the blocks and retained f o r conodonts while the remainder (1/2 to 3/4 of the t o t a l ) was placed i n hydrochloric acid baths ( d i l u t e d , but 7 not to s p e c i f i c percent as the only c r i t e r i o n to.be met was that bubbling was not to be so strong as to cause further breakage of the s i l i c i f i e d f o s -s i l s ) to separate the s i l i c i f i e d brachiopods. The conodont samples were placed i n p l a s t i c buckets which were subsequently f i l l e d 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 at a d i l u t i o n of 1 part acid to 6 to 9 parts water (to keep the acid at or below 10% - stronger a c e t i c acid tends to etch the conodonts while any strength of hydrochloric w i l l d i s s o l v e the con-odonts) . The samples were l e f t i n a fume hood for up to two months but more t y p i c a l l y f o r two to three weeks with the acid being changed weekly. The longer than normal d i s s o l u t i o n period, for such work was required be-cause the rocks, being calcareous quartzose sandstones, were slow to d i s -solve and contained a high percentage of insolubles (as opposed to pure carbonates which are more commonly sampled f or conodonts). Even a f t e r these long periods, the samples were r a r e l y e n t i r e l y dissolved and d i s s o l u t i o n was us u a l l y discontinued 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 residue had been separated. As a r e s u l t , i t i s impossible to report the.actual percentage of in s o l u b l e residue. It was necessary to use a modified procedure f or i s o -l a t i n g the conodonts due to the large i n s o l u b l e f r a c t i o n s . After the samples were dissolved, they were wet sieved and washed through a four sieve stack c o n s i s t i n g of 20 (.841 mm), 35 (.500 mm), 100 (.150 mm) and 200 mesh (.075;mm) standard 21 cm diameter sieves. On top of t h i s stack was a 1.2 mm nylon screen to r e t a i n the coarsest p a r t i c l e s and undissolved chunks. The two coarsest sieves were used to separate any coarse sand or small undissolved fragments from.the f i n e sand and, hopefully, conodonts which would be trapped i n the f i n e s t two sieves. This stack was necessary because of the high percentage of insol u b l e material. Normally, a s i n g l e 8 150 or 200 mesh sieve with a nylon screen on top i s s u f f i c i e n t f o r the i n -solubles of r e l a t i v e l y pure carbonates. A l l of the inso l u b l e residue was retained and l e f t to a i r dry i n porcelain c r u c i b l e s . The s i l t and clay 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 sieve was. also, c o l l e c t e d i n p l a s t i c buckets and allowed to s e t t l e . After the sediment had s e t t l e d most of the excess water was poured o f f and the wet sediment stored i n covered p l a s t i c containers. Some of these samples were l a t e r analyzed for t h e i r palynomorph content i n conjunction with a graduate course with G.E. Rouse at UBC. After drying, the 100 and 200 mesh insoluble f r a c t i o n s were placed i n separatory funnels f i l l e d with tetrabromoethane ( s p e c i f i c gravity = 2.89). The remaining coarser insolubles were placed i n a container and stored i n cabinets. The insolubles i n the gegaratory funnels divided into two f r a c t i o n s : a l i g h t 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 quartz, chert, glauconite and s i l i c i f i e d 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 sinking to the bottom and c o n s i s t i n g of opaques, ir o n coated grains, f i s h debris (teeth, plates...) and conodonts ( s p e c i f i c gravity = 2.84 to 3.10: E l l i s o n , 1944). These heavy f r a c t i o n s were then allowed to run out of the funnel onto a f i l t e r 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 f i l t e r e d onto a separate paper. The tetrabromoethane was constantly reused owing to the high cost of the mat e r i a l . These f r a c t i o n s were then thoroughly washed with acetone and l e f t to dry. The acetone with i t s dissolved tetrabromoethane i n s o l u -t i o n was placed i n an. open beaker and allowed to evaporate i n a fume hood u n t i l the tetrabromoethane was concentrated (acetone evaporates more r a p i d l y ) . This procedure allowed only minimal l o s s of heavy l i q u i d with each separation. After drying, the conodonts were picked from the heavy f r a c t i o n s with the 9 aid of a binocular microscope and a wet, very f i n e paint brush. However, i f the heavy f r a c t i o n was large and contained abundant i r o n minerals, the sample was passed through a magnetic separator where the conodonts are further concentrated i n the non-magnetic heavy f r a c t i o n . This procedure saves unnecessary time spent picking non-productive residues. The conodonts having been concentrated from t h e i r rock i n abundances, when present, ranging from one- to as many as 150 per kg (F49), were thus a v a i l a b l e f o r d e t a i l e d study. The analysis of these, faunas f i r s t consisted of simple observation under the binocular microscope and d e s c r i p t i o n . Secondly, the samples were measured for various parameters with a micro-meter mounted on a.binocular microscope. These measurements were used to enhance descriptions and subjected to various s t a t i s t i c a l procedures as out'^ l i n e d i n a l a t e r chapter. T h i r d l y , the samples were putaonl. Scanning Electron Microscope (SEM) stubs, coated with gold-palladium, and photographed with the SEM. These photos besides providing the i l l u s t r a t i o n s f o r the plates f a c i l i t a t e d even more d e t a i l e d d e s c r i p t i o n . Indeed, adequate d e s c r i p t i o n would be impossible without the SEM. The i d e n t i f i c a t i o n and comparison, to other s i m i l a r conodonts to determine 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 studied sections concluded the analysis of the conodonts. The sections were also analyzed i n terms of t h e i r l i t h o l o g y and other bi o t a . Descriptions 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 notes and a close inspection of hand specimens. Although a few t h i n sections were prepared i t was decided that time was i n s u f f i c i e n t to do an adequate study, nor did i t seem necessary i n a paleontological t h e s i s . The remaining biota were i d e n t i f i e d at high taxonomic l e v e l s and used as.a rough guide to chang-ing b i o f a c i e s . Some brachiopod genera were i d e n t i f i e d as they aided, to a l e s s e r degree, the age determinations of the s t r a t a herein described. 10 STRATIGRAPHY AND PALEOENVIRONMENT - GENERAL STATEMENT A discussion of the stratigraphy f or the marginal f a c i e s of the youngest Permian.on Ellesmere Island follows. This chapter i s based on the megascopic 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-biota and trace -f o s s i l s present. The paleoenvironmental in t e r p r e t a t i o n s : recorded herein are not meant to be d e f i n i t i v e as they are founded on " rough data. The expected c h a r a c t e r i s t i c s f o r various environments as described i n Brenner and Davies (1974), Davies et a l . (1971), Dickinson et a l . (1972), Goldring and Bridges (1973), Harms et a l . (1975), Howard (1972) and M i a l l (1978) are summarized i n Figure 2. Sabine Bay Formation A. Hamilton Peninsula area Here the Sabine Bay Formation, which o v e r l i e s the Belcher Channel Formation and o v e r l a i n by the Assistance,is characterized by c y c l i c s e d i -mentary environments ;.(Fig. 3 shows features mentioned i n t h i s chapter). The formation consists of 180 metres of medium, clean, well sorted, f r i a b l e quartzose sandstones with some f i n e and coarse sand and sparse granules and pebbles. The sandstones are porous (10 to 15% estimated) and usually uncemented although l o c a l c a l c i t e cement i s present. Fresh surfaces are generally white to l i g h t beige i n colour while weathered surfaces are dominantly yellowish brown to brownish orange but may also be :' medium brown and pale red or creamy pink. The sandstones are generally t h i c k bedded to massive but exhibit f a i n t i n t e r n a l laminae upon closer examination. Crossbedding i s not common but l o c a l l y conspicuous. The sequence i s cut by a couple of dykes (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-portant i n terms of hydrocarbon preservation as Thorsteinsson (1974) reports 11 shallow offshore t r a n s i t i o n lower shoreface upper shoreface t r a n s i t i o n foreshore -increasing depth -mcreasinl shore g distance from t h i n beds few s e d i -mentary structures s i l t s t o n e extensive bioturba-t i o n •low energy) organic r i c h conodonts abundant rieogondol-j e l l i d s occur i n deeper lower en^ ergy, lessfc nutrient r i c h environments thah idiognathodids and gn|a \-f ine, and clean sand stone tf i n e sand beds t h i c k en upwards |-siltstone i n t e r -bedded r i p p l e laminae -cross bedding -mega r i p p l e s - f i n e to mediuml d i r t y sand-stone -30 to 45 cm thick beds - p a r a l l e l laminae -lens l i k e crossbedding -abundant large d i s t i n c t trace f o s s i l s -fine to medium, clean| sandstone -well sorted -trough crossbedding -minor amount of burrowing -beds trunca-ted - r i p p l e laminae thodids. -f i n e to medium, clean sandstone -well sort-ed -trough cross-bedding -minor amount of bur-rowing -very clean, well sorted, f i n e to med ium sandstone| -low dipping, p a r a l l e l to sub-p a r a l l e l bedding -some convolute| laminae -no trace f o s s i l s 1 deposit and sediment feeders burrows more horizon-| t a l and develop branches higher d i v e r s i t y -low d i v e r s i t y of burrows, suspension feeders -unbranched, v e r t i c a l to steeply i n c l i n e d burrows Figure 2. Generalized l i s t of c h a r a c t e r i s t i c s for various shallow marine environments. Figure 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 c o r r e l a t i o n of the formations and sections pertinent to t h i s report. 13 bituminous, residues i n an outcrop on Hamilton Peninsula. A coquinoid u n i t of rugosochonetid brachiopods outcrops 30 metres from the top of the formation". Coarse ribbed Spirophyton i s present i n t h i s unit and i n the overlying 30 m, but absent below. The type of ribbing or laminae present on Spirophyton seems to be very useful f or paleoenvironmental i n t e r p r e t a t i o n f o r the Permian rocks of Ellesmere. Marintsch and Finks (1978) i n a study of Devonian Zoophycus (a trace f o s s i l . s i m i l a r to Spirophyton) demonstrated environmental s i g n i -ficance for the mean and maximum diameter.of the trace and for the menis-cus height (related to ri b b i n g diameter of the trace and maximum body diameter of the organism creating the burrow). They found that the animal i s l a r g e s t near the centre of i t s environmental range (quiet, r e l a t i v e l y deep offshore marine), smallest near the margins (shallower, higher energy) and absent i n the shallowest water beds within t h e i r sequence. Observa-tions for the Permian of Ellesmere suggest that the coarse ribbed, smaller diameter (10 to 20 cm) Spirophyton are found i n shallow shoreface environ-ments while the f i n e ribbed, larger diameter (20 to 35 cm) are found i n ." deeper, quieter offshore marine conditions. Any environments interpreted as foreshore or t r a n s i t i o n a l between foreshore and shoreface do not contain any Spirophyton. Apparently, Spirophyton i s also absent from the Van Hauen and Degerbols Formations which are the basinal equivalents of the A s s i s -tance and Trold Fiord Formations. Brachiopods were never found i n abun-dance i n beds containing Spirophyton although a few may be present near by. Two other features are noteworthy with regards to the Sabine Bay For-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 to 12 cm diameter) found on bedding surfaces with large scale r i p p l e s (wavelength = 0.9 to' 1.15 m, Amplitude = 20 to 30 cm) and probably representing an upper 14 shoreface environment. These burrows are f a i r l y , evenly spaced (0.3 m apart) suggesting high competition f o r resources. These burrows have been found at Hamilton Peninsula and McKinley Bay within the Sabine Bay Formation and at Henrietta-Nesmith (Fig. 1) i n possible shallow water equivalents of the Trold Fiord Formation. Secondly, no conodonts or any other microbiota other than palynomorphs are present i n t h i s formation. B. McKinley Bay area Here 33 metres of Sabine Bay Formation unconformably o v e r l i e the Nan-sen and are o v e r l a i n in. turn by a t h i n section of the Assistance Formation. The Sabine Bay begins with d i r t y , f i n e to very f i n e quartzose sandstone with coarse ribbed Spirophyton and carbonaceous material to clean, very f i n e a r e n i t e with a coquina of rugosochonetid.brachiopods a l l of which i s interpreted as a lower shoreface environment. This unit apparently progrades into an upper shoreface environment ( f i n e to medium grained, clean quartzose a r e n i t e s ) , which i n turn transgresses into a lower shoreface environment (brachiopod and bivalve coquinoid quartzose sand-stone), and f i n a l l y progrades into a foreshore environment ( f i n e grained, very clean quartzose a r e n i t e ) . -C. Tanquary Fiord area The formation ranges i n thickness from 36 to 70 metres, thinning towards the north and onlapping the Tanquary s t r u c t u r a l high. The Sabine Bay Formation unconformably o v e r l i e s the Canyon Fiord Formation and i s i n turn o v e r l a i n unconformably by the T r i a s s i c . Bjorne Formation as no A s s i s -tance or Trold Fiord.:equivalents are present. The Sabine Bay can be d i -vided into three u n i t s Including upper and'"lower,'uM'ts-of clean, fi h e - t o -medium grained quartzose arenites representing shallow shoreface to fo r e -15 shore environments, and a middle u n i t of poorly sorted, very f i n e sand-stone to s i l t s t o n e and minor shale with a d i s t i n c t root zone (with c o a l -i f i e d roots or other plant fragments) suggesting a backbarrier lagoon or marsh environment at le a s t p a r t l y emergent. Except for roots no other mega-f o s s i l s or traces were observed. Assistance Formation A. Hamilton Peninsula area The Assistance Formation at Hamilton Peninsula includes between 162 and 178 metres of section depending on the p o s i t i o n of the talus-covered boun-dary with the overlying Trold Fiord Formation. The lowest part of the Assistance consists of a f i n e grained, poorly sorted, quartzose sandstone with carbonaceous material, trace f o s s i l s (coarse ribbed Spirophyton and Skolithos), minor rounded pebbles and fragmented biota at the top of one bed, c a l c i t e cement, and glauconite. The Assistance i s s i m i l a r to parts of the Sabine Bay, except for the glauconite. Although the appearance upsection of glauconite i s abrupt, the remaining l i t h o l o g y suggests that the boundary between the Assistance and the underlying Sabine Bay may be gradational'and continuous. Thorsteinsson (1974) i n t e r p r e t s t h i s boundary as a disconform-i t y which, i f present, must be of short duration. The pebbly sandstone unit i s followed by s t r a t a that t y p i c a l l y weather yellowish grey to greyish orange with f r e s h surfaces being various shades df grey, and composed of very f i n e quartzose sandstone to s i l t s t o n e with v a r i a b l e amounts of calcareous cement. These rocks contain abundant carbon-aceous material, trace f o s s i l s (Asterosoma, f i n e ribbed Spirophyton, Plano-l i t e s (3 to 5 mm; diameter) and other u n i d e n t i f i e d types), and abundant mega-and microbiota. Glauconite: i s present i n a l l of these rocks but never as abundant as i n the overlying Trold Fiord Formation. A large part of the 16 section i s soft and f r i a b l e and was generally measured as cover or talus while harder more calcareous sandstones stand out prominently. Bedding i s generally t h i n to medium but poorly defined. Few other sedimentary structures were observed although carbonaceous material i s often arranged as i r r e g u l a r laminations. A l l factors point to the prevalence of shallow offshore marine conditions away from shoreface environments although a few f i n e grained, cleaner quartzose sandstone beds may represent t r a n s i -t i o n a l beds between offshore and lower shoreface. The extensive bioturba-t i o n and preseace of .- tfre-autnigenic~mirieral- : glauconite suggest that rates of .deposition were .considerably l e s s than for the Sabine Bay. Conodonts are abundant i n the lower half of the formation and include Neogondolella idahoensis n.subsp. A and Anchignathodus minutus. The Neogon-d o l e l l a fauna i s very abundant and include a s i g n i f i c a n t proportion of com-plete specimens. A large number of ramiform elements occur i n a s s o c i a t i o n with Neogondolella. No other assemblage i n the Assistance or Trold Fiord Formations has as many ramiforms compared to platforms; i n f a c t , most had none. The colours of these conodonts are brown to dark brown and have an a l t e r a t i o n index of 2.0 according to Epstein et a l . (1977). This indicates metamorphic temperatures of 60 to 140°C and a f i x e d carbon range of 55 to 70%, well within the l i m i t s for petroleum preservation. Conodonts i n the upper half of the formation are fragmented and rare and include N. serrata(?) and a couple of ramiform fragments. B. McKinley Bay area The Assistance Formation, which was not previously recognized at Mc Kin-l e y Bay, i s defined here as a t h i n (3 to 4 m) unit of. greyish yellow weather-ing, f i n e to medium grained quartzose sandstone followed by a unit of matrix supported, dark grey chert-pebble conglomerate. Large brachiopods and bryozoan fragments occur within t h i s unit which -is v a r i a b l y cemented by c a l -c i t e . R e c r y s t a l l i z e d conodonts (possibly as a r e s u l t of intense heating by a nearby dyke) were found i n one sample (F100) and included Neogondolella  idahoensis and Neostreptognathodus p r a y i . The occurrence of two Neostrep-tognathodus fragments i s unique to t h i s sample. Clark (1974) indicated that gnathodids throve i n very shallow nutrient r i c h water of moderate energy and normal s a l i n i t y , whereas gon d o l e l l i d s arid anchignathodids preferred ' . deeper water, perhaps at the l i m i t of the photic zone. There are no i n d i -cations from the l i t h o l o g y or associated biota at F100 (dominantly moder-ate sized productids and small s p i r i f e r i d s ) , to suggest that the represented environments are shallower than those of other conodont occurrences i n the Assistance at.Hamilton Peninsula. 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 within the basin to-the extent that the McKinley Bay section i s closer to the basin.margin than the Hamilton Peninsula sections. Despite the obvious c y c l i c i t y of environments at Hamilton Peninsula no Neostrepto-gnathodus specimens were found. Those samples with Neogondolella were nearly always associated with 3 to 5 mm diameter P l a n o l i t e s ; untransported megafauna intensely bored, by an e n d o l i t h i c chlorophyte alga indicate a low energy, shallow marine environment well within the photic zone. These obser-vations and i n t e r p r e t a t i o n s suggest that either the d i f f e r e n t environmental conditions 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 are subtle and as yet unrecognized for t h i s area or that Neostreptognathodus was very rare i n the Permian of the Sverdriip Basin. D. Sawtooth Range area This section of Assistance rocks i s very thick (515 to 545 m a f t e r r e -moving the 88 m s i l l ) and can be divided into two u n i t s . The lower 450 m 18 t h i c k unit i s characterized by yellow-grey weathering, f i n e grained quartz-ose sandstone, with v a r i a b l e amounts of carbonaceous material, trace f o s -s i l s (Skolithos and coarse ribbed Spirophyton), calcareous and/or s i l i c e o u s cements and p r a c t i c a l l y no megafossils. Pendants within the s i l l and a few beds above i t contain, brachiopods (Jakutoproductus(?) - see p.32 for s i g n i -ficance) and small pelecypods. As glauconite i s not present i n any of these rocks r e l a t i v e l y rapid deposition i s implied. The proposed paleoenviron-mental i n t e r p r e t a t i o n f o r t h i s unit i s of some sort of a de l t a complex. The deposition i s interpreted as occurring at the de l t a front i n lower shoreface and t r a n s i t i o n a l environments. The thickness of these deposits compared to sections to the northeast, the r a p i d i t y of deposition, dominance of dep-o s i t feeders, and the abundance of carbonaceous material a l l confirm a d e l t a i c environment (Weimer, 1970). The upper 95 m th i c k unit i s characterized by f i n e to very f i n e grained, f o s s i l i f e r o u s (almost coquinoid), quartzose sandstone with varying amounts of carbonaceous material and glauconite. These sediments are i n -tensely burrowed ( f i n e ribbed Spirophyton, and Asterosoma), have i r r e g u l a r platy bedding and are very s i m i l a r to the l i t h o l o g y ..of the Assistance at Hamilton Peninsula. The presence of abundant brachiopods and bryozoans as well as glauconite suggests lower energy conditions and slower deposition. Apparently t h i s unit i s transgressive over the lower u n i t . Transgressions are often i n i t i a t e d when a l l or part of a d e l t a system i s abandoned so that subsidence increases r e l a t i v e to the deposition. Trold Fiord Formation A. Hamilton Peninsula area The Trold Fiord Formation i s characterized by glauconite r i c h , f i n e quartzose sandstones but.also consists of minor biogenic arenaceous lime-stone, chert-pebble conglomerate, and chert (major components are sponge s p i c u l e s ) . On the basis of regional overstepping by the Trold Fiord on older formations from NW to SE, Thorsteinsson (1974) indicated a disconform-i t y at the boundary between the Assistance and Trold Fiord Formations. However, t h i s author saw no evidence f o r such an i n t e r p r e t a t i o n at Hamilton Peninsula. Unless the seas t o t a l l y vacated the basin one would expect some sections to show a nearly continuous record while others may show a major hiatus. The lowest parts of the Trold Fiord (Subdivision D) are characterized by s i l i c i f i e d coquinas of brachiopods i n a g l a u c o n i t i c , f i n e quartzose sandstone which contains only minor amounts of carbonaceous material ( i n d i s t i n c t i o n to the more carbonaceous Assistance) presumably due to decreased introduction of terrigenous plant material further offshore. The coquinas, which are t y p i c a l l y of shallow subtidal o r i g i n , are composed of dominantly small, unfragmented, and sometimes a r t i c u l a t e d productids,.and l e s s e r amounts of fragmented large productids and s p i r i f e r i d s . Other biota form only a small f r a c t i o n of the t o t a l biomass while conodonts are apparently absent. This unit weathers dusky yellow with minor red while f r e s h surfaces are greyish yellow-green and only occasionally red. The majority of the glauconite formed i n the small pores of echinoderm fragments and progressively replaced the structure u n t i l a s o l i d bleb of glauconite r e s u l t e d . Glauconite also formed i n the chambers of small forams and i n the zooecia of bryozoa. I t i s generally regarded that glauconite forms by the a l t e r a t i o n of k a o l i n i t e clays i n l o c a l l y reducing conditions (provided by the small pores of b i o t i c elements, presumably owing to con-20 centrations of decaying organic matter) but i n a generally o x i d i z i n g en-vironment (provided by act i o n of shallow marine waves). As well as being most common on the outer edge of the shelf and on topographic highs, glau-conite i s i n v a r i a b l y associated with low sedimentation rates because of the:, low concentration of soluble i r o n i n the ocean and because the g l a u c o n i t i -zation process stops a f t e r b u r i a l owing to the lo s s of the proper chemical environment (Burst, 1958). A 5% metre unit of chert granule and pebble conglomerate i s a char-a c t e r i s t i c part of Subdivision E.in the middle of the Trold Fiord Formation. The lower f i v e metres are concentrated i n lenses or channels and grade l a t e r a l l y into burrowed (8 to 10 mm diameter P l a n o l i t e s ) , very f i n e to f i n e grained g l a u c o n i t i c quartzose sandstone. The upper 0.5 metres i s represented by a s o l i d bed of red and yellow chert pebble conglomerate which may repre-sent a transgressive lag deposit. Large unbroken productids (Thamnosia) are present i n t h i s i n t e r v a l . Despite t h e i r t h i c k s h e l l s , the energy of the depo s i t i o n a l environment must have been low for the s h e l l s to remain unbro-ken, which seems somewhat anomalous i n l i g h t of the grain s i z e . The brach-iopods are also intensely bored by polychaetes and. barnacles suggesting that they were exposed to the marine environment f o r a s i g n i f i c a n t time before t h e i r incorporation into the conglomerate. The conodonts, which are rare and fragmented i n t h i s u n i t , include Neogondolella n.sp. 13 and N. postserrata(?). The upper parts of'.the Trold Fiord (Subdivision! F) begin with coquinas of s i l i c i f i e d brachiopods s i m i l a r to those lower i n the section. Some beds are dominated by bryozoans where these encrusting sheet-like and c y l i n -d r i c a l trepostomes comprise up to 50% of the rock volume. A minor amount of blue-grey chert also occurs, i n t h i s i n t e r v a l . Carbonaceous blebs and fi l m s are sporadic and u s u a l l y associated with burrowing. The sandstones through-out t h i s unit are. very f i n e to f i n e grained and moderately to well sorted. Weathering colours include greyish orange, greyish yellow and greyish yellow -green whereas f r e s h surfaces are greyish o l i v e to dusky yellow-green. Abundance of conodonts. i n t h i s unit is.comparable to that of Neogon- d o l e l l a n.subsp. A assemblages. Between these two assemblages conodont populations are sparse. The conodonts i n t h i s Trold Fiord u n i t , some of which are complete, include Neogondolella b i t t e r i n.subsp. C and Neogondol-e l l a rosenkrantzi n.subsp. B. Only one small fragmented and u n i d e n t i f i e d ramiform element was found i n a s s o c i a t i o n . Colours of these conodonts are almost o r i g i n a l amber and have a colour a l t e r a t i o n index of 1.5 (very pale brown) according to Epstein et a l . (1977). This indicates metamorphic temperatures of 50 to 90°C and a f i x e d carbon range of 55 to 70%, well within the l i m i t s for petroleum generation and preservation. The highest beds of the Trold Fiord are burrowed (medium ribbed Spirophyton), f i n e grained, quartzose sandstone with small s p i r i f e r i d s and phosphatic nodules containing moderately large i n a r t i c u l a t e brachiopods (Lingula) and f i s h debris. The environments represented by t h i s unit are l a r g e l y shallow subtidal marine to possibly t r a n s i t i o n a l shoreface at 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 Bjorne sandstone. B. McKinley Bay area Here the Trold Fiord Formation ranges i n thickness from 24 to 61 metres over a distance of 1.6 km (1 m i l e ) . The base of the formation consists of one metre of red weathering chert granule and pebble conglomerate with large s p i r i f e r i d s and moderate sized productids (Thamnosia?). Above the conglomerate, s i l i c i f i e d coquinas of brachiopods (mostly of small size) 22 occur i n a very fine,grained sandstone to arenaceous carbonate. At l e a s t part of t h i s unit demonstrates features suggesting hardground development; brachiopod coquinas (most are i n l i f e position) t o t a l l y surrounded by en-cr u s t i n g trepostome bryozoans with the zooecia l a t e r f i l l e d with glauconite. The brachiopods are intensely bored by e n d o l i t h i c algae and acrothoracican barnacles, the density of which may be an indica t o r of r e l a t i v e exposure time. The remainder of the section consists of f i n e grained, g l a u c o n i t i c , quartzose sandstones with minor burrowing (Spirophyton) and rare brachiopods and gastropods. I n a r t i c u l a t e brachiopods characterize the highest beds. Only a couple of u n i d e n t i f i a b l e Neogondolella fragments were found at t h i s l o c a l i t y . D. Sawtooth Range area This sequence of Trold Fiord s t r a t a i s thicker and contains less, biota then the Hamilton Peninsula section. The lower parts of the formation are' characterized by f i n e to very f i n e g l a u c o n i t i c , quartzose sandstones with shaly layers, minor burrowing, and rare brachiopods. The few brachiopod horizons that do occur have been leached of a l l o r i g i n a l s h e l l . The lowest 169 metres are 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 debris. Most of t h i s i n t e r v a l weathers dark green and has more glauconite than quartz grains, however, two samples (L38 and L46) weathered blue-grey i n colour and had f a r l e s s glauconite. The cements i n these sandstones are"dominated by s i l i c a but minor c a l c i t e i s also present. Once again the middle part of the Trold Fiord i s characterized by chert granule and pebble conglomerates. The conglomerate i n t e r v a l i s 20 metres th i c k but discontinuous as the granules and pebbles are concentrated i n lenses or channels. The majority of t h i s unit i s red weathering and has 23/.; abundant abraded and bored b i o c l a s t s (polychaetes, sponge(?), and barnacle borings) including a few Thamnosia and many large s p i r i f e r i d s . Large P l a n o l i t e s (8 to 10 mm diameter), i d e n t i c a l to those at Hamilton Peninsula, are also present. Above the conglomeratic u n i t are about 40 metres of sand-stone, s i m i l a r to the lowest 169 metres. The next 56 metres of section consist, of f i n e grained, v a r i a b l y g l a u c o n i t i c quartzose sandstones that . weather green with occasional p u r p l i s h l a y e r s , lacking i n carbonaceous material, and contain a few small P l a n o l i t e s burrows. There appears to be evidence f o r channeling i n these sandstones as coquinas of brachiopods are l e n s - l i k e i n d i s t r i b u t i o n . The next 30 metres consist of f i n e grained sandstone and minor dark greenish grey chert that would be best described as.a s p i c u l i t e . The section ends with monotonous l i g h t green weathering, f i n e grained quartzose sandstone which, except for one l o c a l i t y with i n a r -t i c u l a t e brachiopods, i s generally devoid of megabiota. It i s d i f f i c u l t to d i s t i n g u i s h p a r t i c u l a r environments but the majority of the section i s undoubtedly of shallow subtidal shelf o r i g i n . I t i s perplexing why brachio-pod l o c a l i t i e s are so few despite the slow depositional rates denoted by the presence of abundant glauconite. No samples from t h i s section were processed for conodonts, l a r g e l y because no s i l i c i f i e d faunas were present. AGE AND CORRELATION A-.correlation chart for Permian stages and zones i s provided i n Figure 4 ; i t includes schemes from Grant and Cooper (1973),.Furnish (1973), Ward-law and Collinson (1979a) y Water house (197.6), and the scheme adopted for t h i s study which i s a combination of the others. The research conducted for 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 Dorashamian D j u l f i a n P u n j a b i a n K a z a n i a u Kungurian ( U f i m i a n ) B a i g e n d z i n i a n Sakmarian A s s e l i a n BB Changhsinglan Chhidruan A r a k s i a n Amarassian C a p i t a n i a n Wordian Road l a n L e o n a r d i a n A k t a s t i n i a n S t e r l i t a m a k i a n T a s t u b i a n A s s e l i a n C C l T a r t a r i a n K a z a n i a n Kungurian A r t i n s k i a n Sakmarian CC2 Ochoan Guadalupian Changhsingian Chhidruan A r a k s i a n Amarassian C a p i t a n i a n Wordian Leonard i a n Wolfcampian Roadian L e o n a r d i a n A k t a s t i n i a n S t e r l i t a m a k i a n T a s t u b i a n A s s e l i a n PD l a) • H TH >u u tH a) 3 cn X N C ttj •rH p. 3 i H to nj (V n) u 5 01 tn a n) •H A i tn tn •H H tu < to Zone 6 Zone 5 EE Zone A M Wordian b a r r e n i n t e r v a l Zone 3 'Zone 2 X Zone 1 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 Dorashamian D j u l f i a n Amarassian C a p i t a n i a n Roadian Leonard i a n Aktastinian Sakmarian T r o l d F i o r d F o r m a t i o n A s s i s t a n c e F o r m a t i o n S a b i n e Bay Form a t i o n S u b d i v i s i o n F S u b d i v i s i o n E S u b d i v i s i o n D S u b d i v i s i o n C S u b d i v i s i o n B S u b d i v i s i o n A F i g u r e 4. Corr AA EE 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 zones o f t h e Permian. Waterhouse ( 1 9 7 5 ) , BB = F u r n i s h (1973), CC = Moore e t a l . T h i s s tudy. L = Lower Permian,M (1965) 1 = Europe, 2 M i d d l e Permian,U « N o r t h A m e r i c a , DD Upper Permian = Wardlaw and C o l l i n s o n (1979a,b). 2 5 r e l a t i o n s h i p s of the Assistance and Trold Fiord Formations through the use of conodonts as the b i o s t r a t i g r a p h i c index. Before discussing the r e s u l t s i t seems appropriate to r e l a t e the state of the art p r i o r to t h i s research. Relative Value of Various F o s s i l Biota Five groups have led the way over a l l others f or the determination of b i o s t r a t i g r a p h i c subdivisions of the Permian. .These include brachiopods, ammonoids, f u s u l i n i d s , palynomorphs and conodonts. Brachiopods were the predominant marine megafauna of the Permian. It i s because of t h i s dominance that they can be used to c o r r e l a t e more rocks and on a wider basis than any other group. It i s the opinion of many authors that as brachiopods are l a t i t u d i n a l l y ( c l i m a t i c a l l y ) c o n t r o l l e d and r e l a t i v e l y long ranging, 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 decreased. However, Waterhouse (1976) states that t h i s i s a widely r e i t e r a t e d misap-prehension and that brachiopod species and genera, during the Permian, were l e s s l a t i t u d i n a l l y or 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 or ammonoids, and j u s t as s h o r t - l i v e d . Even i f Waterhouse i s proven correct by t h i s ' statement, the 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 at the s p e c i f i c l e v e l , very d i f f i c u l t indeed. Waterhouse (1976) also indicates that 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 solved u n t i l paleontologists i n the USSR reexamine brachiopod faun-ules i n the Permian type or standard sections of the Urals. In the forma-tions pertinent to t h i s study brachiopods (productids and s p i r i f e r i d s ) are by f a r the dominant biota but need to be studied at the s p e c i f i c l e v e l since many genera range throughout the e n t i r e section. Because of the problems at the s p e c i f i c l e v e l , age determinations have been of minor value or c o n f l i c t i n g to o f f i c e r s of the GSC concerned with.the Carboniferous and 26 Permian of A r c t i c Canada. Ammonoids are very s h o r t - l i v e d , often even at the generic l e v e l , and thus prove of i n f i n i t e value f o r c o r r e l a t i o n s (Furnish, 1973). However, t h e i r value i s quickly diminished when one considers the r a r i t y of t h e i r : occurrence. According to Waterhouse (1976; f i d e . R.E. Grant, pers. comm.) det a i l e d studies i n West Texas produced only 5,000 ammonoids from 97 l o c a l i t i e s compared to some 3,000,000 brachiopods from about 800 l o c a l i t i e s . Waterhouse (1976) also indicated that i n over 1500 l o c a l i t i e s from the Yukon T e r r i t o r y only f i v e yielded ammonoids. Although ammonoids have been found previously i n the Sabine Bay, Assistance and Trold Fiord Formations (however, rarely) t h i s author found none i n his d e t a i l e d sections. Fusulinids are s h o r t - l i v e d and often, when present, as abundant as brachiopods. In opposition to these p o s i t i v e aspects i s the strong l a t i -t u d inal 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 . Fusulinids are generally r e s t r i c t e d to warm waters and although present i n the Carboni-ferous and E a r l i e s t Permian of Ellesmere Island they are absent from the l a t e Lower and Middle Permian sections covered by t h i s report. Carboniferous and T e r t i a r y systems have long dominated pa l y n o l o g i c a l studies although the Permian i s becoming increasingly important (Hart, 1965; Jansonius, 1962). Palynomorphs could prove very valuable for c o r r e l a t i o n of those parts of sections where other biota are rare or absent. Two samples from the Assistance Formation at Hamilton -•Peninsula yielded palynomorph assemblages dominated by V i t t a t i n a which, according to Hart (1965), i s Kungurian i n age. This form i s s i m i l a r to V. simplex described by Jansonius (1962) from the Permian Belloy Formation of the Peace River area, Canada. Two 27 samples from the Trold Fiord Formation at Hamilton Peninsula yielded ' V . V i t t a t i n a c f . V. l a t a which was f i r s t described from the Guadalupian Flowerpot Formation of Oklahoma, USA (Wilson, 1962). These f l o r a l d i f f e r -ences suggest that these palynomorphs may be us e f u l f o r disc r i m i n a t i n g Assistance and Trold. Fiord equivalents. These palynomorphs were found i n the same samples, as those containing conodonts. A c o r r e l a t i o n scheme combining the d i s t r i b u t i o n s of both of these groups would be us e f u l f o r resol 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 at Henrietta-Nesmith (Fig. 1) which lack conodonts e n t i r e l y . Furthermore, palynomorphs (as well as conodonts) can be used to e s t i -mate the temperatures that t h e i r host s t r a t a were subjected to. The two Assistance samples had an average Thermal A l t e r a t i o n Index (TAI) of be-tween 2.8 and 3.0 (S t a p l i n 1969, 1974) whereas the Trold Fiord samples aver-aged 2.8. According to Epstein et a l . (1977) conodont colour a l t e r a t i o n does not begin u n t i l l a t e stages of palynomorph diagenesis; explaining the very minimal differences f o r palynomorph.TAI's upsection. According to S t a p l i n (1969, 1974). TAI 1s on the order of 2.8 indicate hydrocarbon p o t e n t i a l f o r o i l and wet gas and a mature organic metamorphic f a c i e s . Temperatures of 100°C are 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 within the range of 50 to 140°C suggested by the conodonts. Conodont colour a l t e r -a t i o n indexes of 1.5 to 2.0 and palynomorph TAI's of 2.8 to 3.0 are e n t i r e l y consistent with comparisons presented'in Epstein et a l . (1977). Conodonts, compared to these other groups, are s t i l l i n t h e i r infancy i n terms of t h e i r use as a b i o s t r a t i g r a p h i c index f o r the Permian. As many problems arose from the early intensive study of Permian conodonts Waterhouse (1976) was.;led to express h i s doubt that they w i l l ever be able 28 to provide worldwide c o r r e l a t i o n for :the marine Permian. In f a c t , no conodonts have been found from the cold water Permian of east A u s t r a l i a despite intensive search (N i c o l , 1975). 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 during the l a s t f i v e years, l a r g e l y through the work of Clark,,Behnken, Wardlaw and Co l l i n s o n . This fa c t i n combination with the abundance and excellent preservation of conodonts found i n the cold water faunas of the Assistance and Trold Fiord Formations ( t h i s study) which are c l o s e l y a l l i e d to warm water faunas of West Texas, Wyoming, Montana, Utah, and Idaho ( t h i s cannot be said for the brachiopods from the same areas) j u s t i f y optimism that conodonts w i l l enjoy a very bright future f o r worldwide.correlation of the Permian. Previous Conodont Work The only previous Permian conodont work on Ellesmere Island was based on a s i n g l e sample from the Assistance Formation on Hamilton Peninsula (near the p o s i t i o n of F54) and a sin g l e sample from the base of the Deger-bols Formation (basinal equivalent of the Trold Fiord Formation) from near Otto Fiord as summarized by Kozur and Nassichuk (1977). The Assistance sample yielded s i x conodont specimens, three of which were assigned to N. idahoensis and the remainder to an. intermediate p o s i t i o n between N. idahoensis and N. serrata or N. nankingensis. These samples seem very much l i k e the population samples defined i n t h i s report as N. idahoensis n.subsp. A which are intermediate between N. serrata (not N. nankingensis since serrations are not present on the posterior parts) and N. idahoensis. The authors ( i b i d . ) placed t h e i r fauna i n the Upper Roadian ( t h i s report places i t i n the Lower Roadian). The Degerbols sample yielded several fragmentary conodont specimens 29 a l l belonging to single.species N. c f . N. g r a c i l i s and possibly intermedi-ate between _N.. idahoensis and N. g r a c i l i s to which the authors assigned an> Upper Roadian age (younger than type Roadian but older than Wordian sensu s t r i c t o ) . As indicated i n t h i s report, forms similar, to N. g r a c i l i s occur i n populations of N. idahoensis n.subsp. A which are assignable to the Lower Roadian. However, as these forms are few i n number they are therefore un-l i k e l y to be the only forms present i n a small sample. As indicated e a r l i e r i t seemed reasonable to suggest that the v a r i e t i e s within N. idahoensis n.subsp. A, i f separated as peripheral i s o l a t e s could, following speciation, lead to populations of _N. g r a c i l i s . If t h i s i s indeed the case then the Upper Roadian age i s e n t i r e l y consistent. However, there may be some e c o l -o g i c a l requirements that increase the number of one v a r i e t y or another i n d i f f e r e n t environments. In other words, i t cannot be discounted that the g r a c i l i s v a r i e t y of _N. idahoensis n.subsp. A becomes the dominant member i n the more basinal environment of the Degerbols Formation (as opposed to a subordinate member on the margin^) leading to an age assignment for Kozur and Nassichuk's N. c f . N. g r a c i l i s of Lower Roadian. More work i s indeed necessary to c l a r i f y t h i s problem. 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 area as i t i s understood at present. E a r l i e r Age Assignments for A r c t i c Permian Formations Sabine Bay Formation: (Thorsteinsson, 1974) In 1974 no f o s s i l s had been observed i n the Sabine Bay on Ellesmere Island but i t s age was given early 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 Belcher Channel Formation and below the Assistance. A r t i n s k i a n ammonoids i d e n t i f i e d as Sverdrupites-were reported from basal beds of the Sabine Bay Formation on M e l v i l l e Island. 30 Assistance Formation (Thorsteinsson, 1974) On the Bjorne Peninsula the Assistance Formation i s dated as early A r t i n s k i a n or Aktastinian i n age on the basis of ammonoids (Nassichuk et a l . , 1965) and brachiopods ( i d e n t i f i e d " by J.B. Waterhouse as brachiopod fauna "E" of the N. Yukon and probably to the Jakutoproductus zone). However, t h i s age assignment i s considerably older than the type A s s i s -tance ( G r i n n e l l Peninsula,. Devon Island) and the Assistance i n the v i c i n i t y of Hamilton Peninsula. Harker and Thorsteinsson (1960) suggested an age equivalent to the Baigendzhinian subseries (upper Artinskian) on the basis of brachiopods. Brachiopods from Hamilton Peninsula ( i d e n t i f i e d by J.B. Waterhouse) in d i c a t e an age of Ufimian or Kungurian. Ammonoids (Nassichuk, 1970; Nassichuk et a l . , 1965) ind i c a t e both l a t e s t Early Permian and l a t e s t A r t i n s k i a n (Baigendzhinian) age. Trold Fiord Formation (Thorsteinsson, 1974) Brachiopods ( i d e n t i f i e d by J.B. Waterhouse) from the Trold Fiord Formation indi c a t e a Kazanian age (Wordian substage). A sin g l e ammonoid (Nassichuk et al..., 1965), Neogeoceras macnari, indicates a Guadalupian age. Age Assignments Resulting From This 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 are based on conodonts - as. opposed to the previous work i n the area with abun-dant brachiopods and rare ammonoids. The r e s u l t s r e f i n e , but do not dras-t i c a l l y a l t e r , the ages;.assigned by previous workers. Six subdivisions are proposed for the l a t e Lower and Middle Permian of Ellesmere Island represented by the Sabine Bay, Assistance and Trold Fiord Formations. The t r a c e a b i l i t y of these subdivisions for the e n t i r e A r c t i c Archipelago i s impossible to assess at t h i s time because of the rather infant stage of Permian conodont work i n the area and because of the environmental and r e s u l t i n g l i t h o l o g i c changes into the b a s i n a l equivalents of the described sections. The s i x u n i t s are referred to as subdivisions-, rather than zones because they are i n part based on l i t h o l o g y and environ-ment and i n part on paleontology. The writer i s o p t i m i s t i c that future work w i l l eventually lead to further refinement and synthesis of >;aj' good traceable biozonatiqn based p r i m a r i l y on conodonts but also supplemented by brachiopods. Subdivision A This subdivision i s assigned to the Sabine Bay Formation and regarded 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 . Shallow marine tongues do occur near the top of the formation at both the McKinley Bay and Hamilton Penin^ sula sections. Brachiopods and a few pelecypods were c o l l e c t e d at both of these l o c a l i t i e s - apparently the f i r s t reported f o s s i l s i n Ellesmere I s -land exposures of the formation. Marine tongues with ammonoid f o s s i l s were reported from the Sabine Bay Formation on M e l v i l l e Island. The brachiopods are dominated, almost ex c l u s i v e l y , by rugosochonetids (Neochonetes or Svalbardia). Although the range i s much greater, these brachiopods may be rel a t e d to Waterhouse's (Bamber and Waterhouse, 1971) brachiopod ;fauna of /Leonardian to Roadian age from the N. Yukon. Because of t h e i r p o s i t i o n below the Assistance Formation an'early Leonardian or Baigendzinian age i s assigned. Previous age assignments to the Sabine Bay Formation from M e l v i l l e Island indicated an Aktastinian age at the base of the formation. Rocks assigned to the basal part of.the Assistance Formation on Bjorne Peninsula, Ellesmere Island indicated a c o r r e l a t i o n with Waterhouse's ( i b i d ) 32 Jakutoproductus zone of the.N. Yukon and an'Aktastinian age. F o s s i l c o l l e c -tions from near the base of the Sawtooth Range section ( F i g . 3) contain productids u n l i k e any seen at Hamilton Peninsula and t e n t a t i v e l y i d e n t i f i e d as Jakutoproductus. These c o l l e c t i o n s are followed by a few hundred metres of l a r g e l y u n f o s s i l i f e r o u s section before abundant brachiopod f o s s i l s are once again encountered - these faunas being very s i m i l a r to those assigned a Late Leonardian to Early Roadian age at Hamilton Peninsula. The age for the basal parts of:the Bjorne Peninsula and Sawtooth Range sections suggest greater c o r r e l a t i o n . t o the Sabine Bay Formation than to the Assistance For-mation. L i t h o l o g i c differences have resulted in.these s t r a t a being assigned to the Assistance Formation but they might be better described as a new formation. Furthermore, i t would appear that the Sabine Bay Formation on Hamilton Peninsula and at McKinley Bay i s younger than that at Bjorne Peninsula or at the Sawtooth Range. Inosummary, although the unit 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 that the unit ranges i n age from Aktastinian to Early and possibly Medial Leonardian of Baigendzinian. Subdivision B This subdivision i s defined as a l i t h o s t r a t i g r a p h i c unit at the base and a b i o s t r a t i g r a p h i c range zone at the top".. In other words, i t includes a l l that section above the top of the Sabine Bay Formation Assignable to the Assistance formation up to the top of the range for N. idahoensis n.subsp. A. In the Hamilton Peninsula section t h i s includes the s t r a t a from the base of the Assistance Formation to the top of F54 (Fig. 3). At McKinley Bay t h i s subdivision comprises a l l the s t r a t a assigned to the Assistance Formation. Neogondolella idahoensis has been reported extensively i n sections from 33 the western United States.. Similar assemblages to the conodont fauna of subdivision B i n the Assistance Formation have been reported from the Meade Peak Phosphatic Shale Member of the Phosphoria Formation of Idaho, Wyoming, and Utah (Youngquist et a l . , 1951; Clark and Ethington, 1962; Clark and Behnken, 1971; Wardlaw and C o l l i n s o n ( f i g . 3), 19;79b)and assigned a Roadian age. Other s i m i l a r faunas occur i n the Bone Spring Limestone, V i c t o r i o Peak Formation, Guadalupe Mountains, Texas (Leonardian i n age according to Behnken (1975) and Upper Leonardian to Lower Roadian by Wardlaw and Collinson (1978)). According to Wardlaw and Collinson (1979a) N-. idahoensis c e r t a i n l y ranges through the l a t e s t part of the Leonardian and doubtfully into the early part of the Roadian. N. serrata on the other hand has been reported from the Cutoff, Brushy Canyon and Getaway Member of the Cherry Canyon Formation of West Texas and from the Meade Peak Phos-phatic shale member of the Phosphoria Formation and assigned a Roadian to Early Wordian age (Clark and Behnken, 1979; Behnken, 1975; and Clark and Ethington, 1962). Neostreptognathodus p r a y i has been reported from the K a i -bab of Nevada and Utah and the Bone Spring Limestone and the V i c t o r i o Peak Formation of West Texas where i t i s apparently r e s t r i c t e d to the Late Leonardian. It appears that subdivision B can be correlated with the Late Leonard-ian to Early Roadian of the western United States. The assemblage for F100 (Fig. 3) at McKinley Bay.which has N. idahoensis subsp. indet. i n associa 1? t i o n with Neostreptognathodus prayt suggests a Late Leonardian age. The faunas q u a n t i t a t i v e l y studied i n d e t a i l at Hamilton Peninsula (F48 to F54) appear to be morphologically intermediate between Neogondolella idahoensis and N. serrata suggesting that an early Roadian age i s more appropriate. 34 No serrations were noted on samples from F100 and because of the associa-. t i o n with Neostreptognathodus p r a y i these representatives of Neogondolella  idahoensis are considered older than those from F48 to F54. However, since the d i s t r i b u t i o n s of Neostreptognathodus p r a y i and Neogondolella  idahoensis are f a c i e s c o n t r o l l e d to the extent that they r a r e l y occur i n t e r -bedded i n a s i n g l e section, i t i s d i f f i c u l t to assess the s i g n i f i c a n c e of Neostreptognathodus p r a y i without i t s presence at Hamilton Peninsula. As well i t i s impossible to assess whether Lower Roadian s t r a t a are present and condensed, were o r i g i n a l l y present and eroded, or unrecognized, or were never deposited at McKinley Bay. This subdivision can be correlated i n part with the Kapp St a r o s t i n Formation of Spitsbergen (Szaniawski and Malkowski, 1979) where the authors report N. idahoensis, N. c f . N. g r a c i l i s , and Neostreptognathodus s v a l - bardensis,. Sweetocristatus a r c t i c u s and several ramiform elements. This s u b d i v i s i o n i s also correlated with Wardlaw and Collinson's (1979a)well defined biozonation f o r the Great Basin-Rocky Mountain Region USA as including t h e i r Zone 1 (Peniculauris i v e s i - Neostreptognathodus  p r a y i zone) and the lower part of Zone 2 (Peniculauris bassi - Neostrepto- gnathodus s u l c o p l i c a t u s zone). As a r e s u l t of these age assignments the base of the Assistance Forma-t i o n at Hamilton Peninsula can be regarded as no older than Late Leonardian. Subdivision C This subdivision i s very loosely defined and i s represented by a couple of sparse conodont c o l l e c t i o n s at F63 and F73 from the Hamilton Peninsula section. The base of the unit, i s defined by the top of subdivision B while the upper l i m i t i s defined as the highest part of the Assistance Formation 35 (although covered by t a l u s ) . A large part of subdivision C i s covered by talus r e s u l t i n g i n sparse c o l l e c t i o n s . The low number of c o l l e c t i o n s i s also owing to the decreased abundance and possible lower- d i v e r s i t y of brachiopods apparent i n the rocks. Conodonts show an equal~. i f not more dramatic, decrease i n numbers when they are present at a l l . Those conodonts reported are questionably assigned to N. serrata(?) because of t h e i r smaller s i z e at apparently similar, growth stages to N. idahoensis n.subsp. A, sharper more compressed posterior d e n t i c l e s , and! the lack of four d i s t i n c t i v e node-like d e n t i c l e s j u s t anterior of the cusp. As reported i n the discussion for subdivision B, N. serrata has a range d i s t r i b u t i o n of Medial Roadian to Early Wordian. Because of the age assignment to the overlying subdivisions, t h i s subdivision C can be assigned a Medial to Late Roadian age. Subdivision .C can be correlated with Wardlaw and Collinson's (1979a) biozonation as including the upper part of Zone 2, Zone 3 (Peniculauris  b a s s i - Neostreptognathodus sp. C zone), and t h e i r barren i n t e r v a l at the top of the Roadian. Perhaps t h i s paucity of conodonts seen for the Roadian at Hamilton Peninsula i s r e a l and not r e l a t e d to preservational factors since Wardlaw and Collinson (1979a) also report a barren i n t e r v a l i n the Upper Roadian to lowermost Wordian. The conodonts may have undergone a c r i s i s l i k e that ofpre-Wolfcampian conodonts (Clark, 1972) which would i n part explain the subsequent increased abundance and d i v e r s i t y i n the middle and upper parts of the Trold Fiord Formation (as i s often the case a f t e r a near-extinction). The top!of the Assistance can therefore be regarded as no younger than Upper Roadian and the f u l l range for the Assistance Formation at Hamilton Peninsula i s from the Upper Leonardian to Upper Roadian. Subdivision. D This subdivision i s also lacking i n conodonts but dominated by the brachiopod Cancrinelloides. Above unit D the presence of N. postserrata(?), N. b i t t e r i n,. subsp.::C and N. rosenkrantzi n.subsp. J) i n d i c a t e a Late Wordian or Kazahian age, so that D i t s e l f i s regarded as Early Wordian or Late Kungurian i n age. Waterhouse, i n Bamber and Waterhouse (1971), reports a Cancrinelloides zone i n the N. Yukon which he assigns a Kazanian age. He does indi c a t e , however, that Cancrinelloides can occur i n s l i g h t l y lower beds included/ i n h i s Thamnosia zone; Thamnosia i s the dominant genus i n t h i s report's subdivision E. Waterhouse assigns a Late Ufimian or Kungurian age to the Thamnosia zone. For t h i s reason i t seems reasonable to assign an Early Wordian age to unit D. Wardlaw and Collinson (1979a)also define a Thamnosia depressa zone which they assign an Early Wordian age. I f the overlying subdivision E at Hamilton Peninsula i s dominated, by Thamnosia and various Neogondolella species then subdivision D cannot be as young as Kazanian. Furthermore, since the base of unit D i s defined by the base of the Trold Fiord Forma-t i o n then at l e a s t part of the Trold Fiord i s older than Kazanian; u n l i k e previous reports which r e s t r i c t e d i t to the Kazanian. The four d i s t i n c t i v e features of t h i s unit are the apparent lack of conodonts, the lack of. large t h i c k - s h e l l e d productids l i k e Thamnosia, the presence of Cancrinelloides and occurrence i n the base of the green sand-stones of the Trold Fiord Formation. The environmental 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 these brachiopods i s not well understood (Waterhouse, 1973). U n t i l more sections i n the area are studied, encompassing a wide v a r i e t y 37 of environments, both marginal and 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 -cance of t h i s unit D cannot be established. Subdivision E Unit E i s the c l o s e s t to a r t r u e range or acme biozone as i t i s based on the presence and dominance of the brachiopod Thamnosia and the occur-rence of reasonably abundant conodonts. Also c h a r a c t e r i s t i c of t h i s subdivision i s a conglomeratic unit that, except for varying thickness, i s i d e n t i c a l i n the McKinley Bay, Hamilton Peninsula and Sawtooth Range sec-t i o n s . If t h i s conglomerate can be shown to be synchronous then i t could prove a very u s e f u l marker horizon(Fig. 3) - possibly r e l a t e d to a s i n g l e tectonic pulse or other short-duration p h y s i c a l phenomenon. At Hamilton Peninsula subdivision E includes f o s s i l c o l l e c t i o n s F83 to F90 and F35 to F44. F36 and F83 contain specimens of a new species; Neogondolella n. sp. J3. Both of these c o l l e c t i o n s occur below the chert-peb-ble conglomeratic u n i t . F87 includes conodont fragments questionably r e -fe r a b l e to N. postserrata(?) . This.-collection occurs within the conglomera-t i c u n i t . N. postserrata has previously been reported from the Southwells Member of the Cherry Canyon Formation to the Lower McCombs Member of the B e l l Canyon Formation of Idaho and Texas and assigned a Wordian and Capi-tanian age (Behnken, 1975; Clark and Behnken, 1979) . Wardlaw and Collinson (1979a)describe a Thamnosia depressa zone from the upper part of the Plympton Formation, and Rex Chert Member of the Phosphoria Formation i n the Great Basin and Rocky Mountain region of the western USA to which they assign an Early to Medial Wordian.age. They indi c a t e that t h i s zone i s more or l e s s equivalent to the Neospathodus ar c u c r i s t a t u s assemblage (Clark and Behnken, 1971; Clark et a l . , 197.9) which Clark et a l . assign a Wordian age. 38 Clark et a l . (1979) indi c a t e the equivalence of t h i s zone to the Neogon- d o l e l l a d e n t i c u l a t a fauna of West Texas despite giving i t a Capitanian to Amarassian age i n Clark and Behnken (1979). Wardlaw and Collinson (1979a) and Clark et a l . (1979) both describe overlying faunas o f N. b i t t e r i and _N. rosenkrantzi assigned to a Late Wordian and Capitanian age. For these reasons N. postserrata (which i s older than N. denticulata) cannot be any younger than Wordian. Assuming ages assigned to the type N. postserrata and f o r the Thamnosia depressa zone the c o l l e c t i o n s below and within the * ;-conglomerate of subdivision E can be regarded as Medial Wordian or Upper-most Kungurian (Ufimian) i n age. Supporting t h i s assignment i s another Thamnosia zone described by Waterhouse (Bamber and Waterhouse, 1971) from the N. Yukon. Here Waterhouse assigns a Late Ufimian age st a t i n g that the Thamnosia of h i s zone-Ft are more evolved than those found i n the Assistance Formation. Thamnosia i s abundant throughout the conglomeratic unit and s l i g h t l y above i t and thus defines the top of unit E, Late Kungurian or Medial Wordian i n age. Subdivision F Subdivision F occurs above the zone with dominant Thamnosia and i n -cludes the f o s s i l c o l l e c t i o n s F91 to F97 and F45 to F47 . The conodonts i d e n t i f i e d from t h i s zone include Neogondolella b i t t e r i n.subsp. C and N. rosenkrantzi n.subsp. I). Brachiopods.include various s p i r i f e r i d s and productids i n varying degrees of abundance; e s p e c i a l l y common are species of Yakovlevia and Kuvelousia. However, both of these genera occur i n many underlying zones and require s p e c i f i c i d e n t i f i c a t i o n before they could be <used :for range-zone..determination. The fauna j u s t described compares very well with that present i n Wardlaw and Collinson'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 b i t t e r i zone) from the Great Basin -Rocky Mountain region of western USA and assigned a l a t e Wordian age. The conodonts at hand are very s i m i l a r to faunas figured i n Wardlaw and Collinson (1979b) from the Retort Phosphatic Shale Member of the Phosphoria Formation from Montana ( i n the case of N. rosenkrantzi) and Wyoming (for _N. b i t t e r i ) . Representatives f o r both.species from the Gerster Formation appear more advanced. N_. rosenkrantzi has also been described from East Greenland (where i t was named by Bender and Stoppel,l965)though the age r e l a t i o n s h i p s were not c l e a r at the time. Clark and Behnken (1979) and Clark et a l . (1979) assign a Capitanian and Amarassian age to faunas described from the Radar, McCombs, and Lamar Members of the B e l l Canyon.Formation of Texas and the Gerster Formation of Nevada and Wyoming. The main reason for contradicting age assignments are the differences of opinion for the recognition of N. b i t t e r i and N. rosenkrantzi by Wardlaw and Collinson and Clark et a l . Clark et a l . (1979) i d e n t i f i e d specimens of N. rosenkrantzi (according to Wardlaw and Collinson, 1979b)as N. b i t t e r i . Therefore, N. rosenkrantzi can occur down into the Late Wordian,(see Clark et a l . , 1979 for discussion and d e s c r i p -tion) . Having resolved these contrasting age assignments and stated that the conodonts at hand are l e s s advanced than Capitanian specimens from the Gerster limestone, I assign subdivision F to a Late Wordian or Kazanian age. One other feature that 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 subdivision at the Sawtooth Range and Hamilton Peninsula i s the dominance of i n a r t i c u l a t e brachiopods near the top. Although such a feature i s probably environ-mentally controlled,the horizon could prove to be a useful marker ( F i g . 3). 40 The absence of i n a r t i c u l a t e s at McKinley Bay suggests that a l l or part of t h i s subdivision, i s missing here: perhaps as a r e s u l t of regression i n the l a s t part of the Trold Fiord Formation. The top "of u n i t F i s coincident with the top of the Trold Fiord For-mation, the l a s t Permian s t r a t a i n the! A r c t i c . The B l i n d Fiord or Bjorne Formations of Lower T r i a s s i c age r e s t unconformably on the Trold Fiord. Indications are that the Trold Fiord Formation ranges i n age from the Lower Wordian or Upper Kungurian to the Uppermost Wordian or Upper Kazanian. Although there i s no d i r e c t evidence for i t , a Capitanian age represented i n the upper parts of the Trold Fiord Formation cannot be e n t i r e l y ruled out since N. b i t t e r i and N./rosenkrantzi can range t h i s high. I t i s also impossible to say whether younger rocks may have been.present and eroded, or never deposited at a l l . I t i s f a i r to say that the time represented by the Trold Fiord/Bjorne unconformity i s considerable: l n the order of 10 m i l l i o n years. Summary The Sabine Bay, Assistance and Trold Fiord Formations have been separ-ated into s i x subdivisions which can be c o l l e c t i v e l y r eferred to as Aktas-t i n i a n to Latest Wordian or Kazanian age. Although the three formations are separated by unconformities (Thorsteinsson, 1974) t h i s author.considers them, " i f present at a l l , to be Q f short"duration. Perhaps more basinal sections could i n d i c a t e continuous sedimentation with the transgressions and regressions only a f f e c t i n g the margins of the basin. QUANTITATIVE ANALYSIS OF MEASURABLE CHARACTERS. FOR NEOGONDOLELLA  Introduction Q u a l i t a t i v e observation of the conodont populations i n samples F48, F49, F52, F53 and'F54 indicated that there are no d i s t i n g u i s h a b l e d i f f e r -ences with respect to o v e r a l l platform shape and d e n t i c l e configuration. A d e s c r i p t i o n based on these observations was s u f f i c i e n t to erect a new subspecies of Neogondolella idahoensis but did nothing to elucidate any evolution i n the populations. It also seemed expedient to have more than j u s t my subjective opinion on which to base the new subspecies. As a. r e s u l t , a quantitative analysis was.^undertaken to determine i f there was any demonstrable evolution i n the populations and to provide unbiased c r i t e r i a on which to base the taxonomyr(see Appendix I for data) v The measurable parameters analyzed include the o v e r a l l platform length ( L l ) , the length from the posterior cusp to the fourth d e n t i c l e a n terior of the cusp (L2), the height from the t i p of the cusp to the base of the flange (Hi), the maximum width (Wl), the width at the posterior end of the platform (W2) , and the number of d e n t i c l e s (//) on the platform. Some of these parameters have been measured on s i m i l a r conodont populations by other workers ( L l and # by Behnken, 1975; L l , HI and // by Dzlk and Trammer, 1980). In a d d i t i o n a number of r a t i o s were determined including L l / H l , Ll/Wl and Ll/#. The f i n a l parameter analyzed i s a function of area i n the posterior end of the platform determined by the equation L2(Wl + W2)% These various parameters were analyzed by c a l c u l a t i n g the mean, stan-dard error for the mean, the standard deviation of the mean and, f i n a l l y , a comparison using z- and t - t e s t s to determine the s t a t i s t i c a l s i g n i f i c a n c e of any differences between populations. Before l i s t i n g and discussing the r e s u l t s of these c a l c u l a t i o n s the background, assumptions and implications of such s t a t i s t i c a l t e s t i n g should be outlined. Above a l l one must remember that i n each, case we are dealing with a sample of the population and not the population i t s e l f . Any two samples 42-from the same population w i l l vary; however, i n samples of s u f f i c i e n t s i z e t h i s d i f f e r e n c e isminimal. Since the population i s the unit of evo l -ution,-, ::the: samples studied herein can only be regarded as approximating the evolutionary trends. A requirement for a sample to be representative of the population i s that i t be selected at random. The conodonts selected for t h i s study were a l l . those (large and small) i n each f o s s i l c o l l e c t i o n that were s u f f i c i e n t l y preserved to allow for the measurement of the various characters. The only processes i n the s e l e c t i o n of the specimens were those as a r e s u l t of the environment of deposition, the diagenetic h i s t o r y and the sample processing. Although these processes can be non-random ( p r e f e r e n t i a l breakage or etching of more f r a g i l e specimens), the r e s u l t i n g frequency curves ( F i g . 5) reasonably approximate a normal d i s t r i b u t i o n , suggesting that the processes were too small to be s e l e c t i v e . Having a r e -presentative approximation of the population, the sample i s now a v a i l a b l e for s t a t i s t i c a l a n a l y s i s . Calculation of the mean of any character involves EX the simple solving of the equation'M'= — (M = mean, ZX = sum of the values for a character, and N = the number of specimens within the sample). However, Burma (1948). points out that the mean of a sample c o n s i s t i n g of a growth seri e s i s merely the mean size of half grown specimens, the min-imum s i z e i s that of the smallest of the youngest specimens and the maxi-mum s i z e i s the largest of the oldest specimens. Burma (1948) furthermore states that such a procedure i s meaningless and lacking i n b i o l o g i c a l s i g -n i f icance: .and that i f one dictum i s established i n quantitative paleontology i t should be that comparisons of one character, to be v a l i d , must be made at comparable growth stages only. His point i s well, 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 the conodonts. Such a procedure would E l e m e n t R e p r e s e n t a t i o n fo r N u m b e r of D e n t i c l e s F 4 8 F 4 9 F 5 2 F53 F54 F49+F52 F53 + F54 r 5 E : 8 13 Ju A i i i 15 8 13 7 16 6 15 7 15 6 16 o v e r a l l 16 E l e m e n t R e p r e s e n t a t i o n f o r P l a t f o r m L e n g t h (pm) F 4 8 F 4 9 F52 F53 F54 F4 9+F52 F53+F54 I I I 1 • I in o LO 8 id JUL .lk. O O S Q o o •0 io £ u> io io g o o o o o o in in io to 05 io m * v eo co a co * o v e r a l l Figure 5. 4 4 be easy with a group l i k e the ammonites where features are present that allow one to d i s t i n g u i s h an adult conch. However, no such features e x i s t for the conodont platform (Dzik and Trammer, 1980). Are we defeated be-fore even beginning the analysis?. Other workers have shown that conodonto-phores l i k e Neogondolella Have a complex ontogeny. M e r r i l l and Powell ' (1980) demonstrated an ontogeny of Pennsylvanian Gondolella where the ap-paratus began as ramiform elements only and subsequently developed into platform ("juvenile") and ramiform elements and f i n a l l y into a platform only ("mature") apparatus. Other workers have also suggested that more than one pair of platforms, each of which are at::different developmental stages, comprise the apparatus. In other words, the developmental stage of the platform may bear no r e l a t i o n s h i p to the actual age of the conodont animal. Unless the samples display some unusual mortality rate, i t may be v a l i d to compare the e n t i r e range of platform sizes as the majority of them probably reached a c e r t a i n stage i n development before death. With these considerations i n mind I proceeded with the s t a t i s t i c a l comparisons of the ent i r e sample but also separated, the data into two subsets (those elements with 10 d e n t i c l e s and those with 11) of possible p a r t i c u l a r growth stages. After c a l c u l a t i n g the mean of each sample the standard error of the mean was calculated at the 95% confidence l e v e l (a = —S- where a-== standard m . N deviation or measure of c e n t r a l tendency of 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 set at the 75% and 95% l e v e l s (M - 1.15a arid M -2.0a r e s p e c t i v e l y ) . Proper use and f u l l value of. these c a l c u l a t i o n s as-sumes a near normal d i s t r i b u t i o n of the characters. These values were then graphed for v i s u a l impact and compared i n the cases of length and the r a t i o length/number of d e n t i c l e s using z- and t - t e s t s to 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 . The t - t e s t assumes a normal d i s t r i b u t i o n and equal standard deviations for the two samples being compared (Hodges et a l . , 1975) and i s valuable for samples of low number. The z-test does not de-pend on the same assumptions but i s only useful f o r samples of approxi-mately 20 or more specimens. It i s generally thought that the quantitative approach i n science i s the only t r u l y objective approach (Raup and Stanley, 1971; p. 42). Despite t h i s a s s e r t i o n the technique i s often met with objection by paleontolo-g i s t s . Many workers state that the method i s good for a large number of specimens but not for a few. However, s t a t i s t i c a l methods are a v a i l a b l e fo r study of samples with a very few specimens or with 1000 or more (Burma, 1948) . Secondly, anything which a person attempts to do with a small sam-ple, which he could not do by s t a t i s t i c a l analysis w i l l probably be founded on error ( i b i d . ) . Another objection i s that misuse of s t a t i s t i c s by those who apply them to paleontology i s often either owing to lack of knowledge or lack of appreciation, of the philosophy behind them ( i b i d . ) . So long as one recognizes the l i m i t s of i n t e r p r e t a t i o n for the derived information t h i s l a s t objection should be minimized. The value of such an analysis to define minute changes i n the specimens through time, and to:.indicate the f u l l range of v a r i a t i o n of any species or i n f r a s p e c i f i c unit f a r out-weighs these objections. A l l species that are created with the concept of species being 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 with suspicion ( i b i d . ) . Results and Discussion The measurements for the specimens studied are included i n Appendix A. A l l measurements were completed with a micrometer set i n a binocular micro-scope at 75 power.. This, method allowed an approximate accuracy of + 5 um for the length .of a 1000 um platform; i n other words an accuracy of approx-imately 1.0% and allowing f o r at l e a s t three s i g n i f i c a n t f i g u r e s . Except for F54, the samples were measured i n terms of a l l the parameters studied. Only length and the number of d e n t i c l e s were determined f o r F54 as these specimens were placed on SEM stubs and photographed before i t seemed appro-p r i a t e to proceed with the more d e t a i l e d a n a l y s i s . Overall length of platform (Ll) Perhaps the most obvious r e f l e c t i o n of increasing maturity of the platform i s an increase i n length. More s i g n i f i c a n t i s the progressive increase upsection f o r the mean value of length from 816 um to 905 um (Table 1). Although the v a r i a t i o n i n the population samples overlaps for the most part, the increase i n the mean value i s e n t i r e l y consistent upsection ( F i g . 6) for N. idahoensis n.subsp. A. However, N. rosenkrantzi n.subsp. D_ represented i n F96 has a :mean length considerably l e s s than that for F48 to F54 i n d i c a t i n g a r e v e r s a l i n the trend. The large v a r i a b i l i t y i n length 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 , at best, of 16% (Table 2) when subjected to the z-test. This i s of even les s s i g n i f i c a n c e than that represented f o r the change of length from F49 to F54 (11.5%). Despite t h i s lack of s i g n i f i c a n c e f o r the dif f e r e n c e , the trend indicated i s consistent with the evolution of N. idahoensis to N. rosenkrantzi according to the r e s u l t s of other workers.(Behnken, 1975; Clark and Behnken, 1979). Although the trend from F48 to F54 may possibly be explained by evolutionary processes, i t could also be the r e s u l t of the ontogenetic stages preserved. M e r r i l l and Powell (1980) indicated that the ontogeny of Pennsylvanian Gondolella proceeded from ramiform only to platform only "ST" • Sample Parameter N. Data — F48 N - (9-10) F49 N = (58-60) F52 N - (24-25) F53 N =(38-40) F54 N = (37) ^lpm M ± 2a m a 75% range* 95% range** 816 ± 134 212 572 - 1060 402 - 1230 847 ± 48 184 635 - 1059 479 - 1215 847 ± 80 199 618 - 1076 449 - 1245 888 ± 82 259 590 - 1186 370 - 1406 905 ± 84 255 612 - 1198 395 - 1415 L2)im M + 2a in a 75% range* 95% range** 297 + 26 40 251 - 343 217 - 377 279 ± 10 42 231 - 327 195 - 363 286 ± 16 38 242 - 330 210 - 3 62 303 ± 16 53 24 2 - 364 197 - 4 09 lvim M ± 2o m a 75% range* 95% range** 181 ± 28 44 130 - 232 93 - 269 183 ± 1.6 61 113 - 253 61 - 305 200 ± 18 44 149 - 251 112 - 288 209 + 18 55 146 - 272 99 - 319 W2um M ± 2o m a 75% r a n g e * 95% range** 152 ± 24 38 108 - 196 76 - 228 163 ± 8 31 127 - 199 101 - 225 174 ± 18 47 120 - 228. 80 - 268 180 ± 16 53 119 - 241 74 - 286 1pm M ± 2a m a 75% range* 95% range** 149 + 2 0 29 116 - 182 91 - 207 164 ± 10 35 124 - 204 94 - 234 166 ± 18 42 118 - 214 82 - 250 172 + 14 44 121 - 223 84 - 260 9 M ± 2o ra a 75% r a n g e * 95% range** 10.00 + 1.02 1.61 8.15 - 11.85 6.78 - 13.22 10.78 ± .42 1.62 8.92 - 12.64 7.54 - 14.02 10.80 ± .60 1.50 9.07 - 12.53 7.80 - 13.80 11.02 ± .66 2.12 8.58 - 13.46 6.78 - 15.26 10.92 ± .60 1.80 8.85 - 12.99 7.32 - 14.52 M i 20 m a 75% r a n g e * 95% range** 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 - 88.5 58.6 - 96.6 79.3 + 3.6 11.1 66.5 - 92.1 57.1 - 101.5 82.0 + 5.0 15.3 64.4 - 99.6 51.4 - 112.6 V W1 M ± 2o m 0 75% r a n g e * 95% r a n g e * * 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 L l / H l M i 2a m a 75% r a n g e * 95% r a n g e * * 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 L 2(W 1+W 2)5i I O 4 2 nm M ± 2 0 m a 75% r a n g e * 95% r a n o e * * 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 Table 1. S t a t i s t i c s from data given i n Appendix I. See page 44 f o r d i s c u s s i o n of s t a t i s t i c a l values used. - 1 1 1 1 1 1 1 I 7 J j ^ 9 10 EC 11 1 1— 12 i i 13 L , / x10" 2 3 3 D SE • B 8 9 I A 10 11 12 13 +- 1——I 1 1 i 1——f 1- h 1 1—-I 1 1 1 1 -I——I 1 1 1 1 H 650 850 j-.'iM.'i ••'•.••••V.>: 1050 1250 1450 1650 l'r-:-'.' A«BMlSlim^ l.-!>W.!'.-. •i H 6 650 • B =3 A 850 1050 •+- 1 —I 1 1 1 h r-1250 H h 1 r-1450 1650 H H 1 1 1 h 3D H 1-17 8 10 11 12 13 14 15 16 F9& denticles HE • B 6 » i II 8 9 10 11 12 . 9 I 13 14 • • < 15 F54 F53 F52 F49 F48 16 Figure 6. Graphical representation of part of" Table 1. Lines i n f u l l bla_ck show range of M±-2'a^. Stippled pattern = 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 z-Test f o r # 'z-Test for' L /#' Samples Compared T~ z-score t t P-value E n t i r • t z-score e Sample I n 1 + + P-value - t z-score P-value^ F49 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 1 4.35 0.000 Table 2. Values derived from z-tests f o r L l , // and'Ll/7/. the z-score of the di f f e r e n c e between two independent samples (1, 2) M = mean a = standard d e v i a t i o n N = number of specimens ft= the area under a normal curve ( t o t a l area = 1) to the l e f t or r i g h t of M ± za which indicates the p r o b a b i l i t y that the d i f f e r e n c e between samples 1 and 2 could occur by chance alone (a two sided t e s t i s equal to the t o t a l area to the l e f t and r i g h t arid i n d i c a t e s the combined chance of getting a deviation i n either d i r e c t i o n t = -Mi - M2 Nl N 2 50 apparatuses-. If t h i s i s true for the Neogondolella.. apparatus as w e l l , then the increasing degree of platform overrepresentation or, rather, ramiform underrepresentation, could be r e l a t e d to increasing maturity and larger ;,:' s i z e of the preserved sample. Table 3 l i s t s the data:for element repre-sentation and demonstrates a f l u c t u a t i o n i n the platform to ramiform r a t i o . Except for the change from F49 to F52 the data show a c l e a r increase i n the ramiform underrepresentation comparing well, with the increase i n length f o r the same i n t e r v a l . The mean value f o r length i n F52 i s equal to that for F49 which, although i t should be a decrease, approximates the trend i n element representation. Although i t seems c l e a r that t h i s concept may be important for .explaining some of the change i n length nevertheless i t i s not completely s a t i s f a c t o r y . Heeding Burma's (1948) plea that only s i m i l a r growth stages should be s t a t i s t i c a l l y compared the data for each sample were divided into two sub-sets (Table 4): the f i r s t with a l l those elements with 10 d e n t i c l e s and the second with a l l the. elements with 11 d e n t i c l e s . The number of d e n t i c l e s cannot be considered a perfect i n d i c a t o r of r e l a t i v e age, however, i t i s true that t h e i r number increases during the i n f e r r e d ontogeny of the e l e -ment. Furthermore, 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 -ing r e l a t i v e age. The values derived from t - t e s t s of the r e s u l t i n g sub-sets provide some i n t e r e s t i n g r e s u l t s . The diff e r e n c e between the samples with 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 with 11 d e n t i c l e s i n F49 and F54 i s significant., even at the 1% l e v e l . The graph i n Figure 7 i l l u s t r a t e s t h i s observation well i n that as d e n t i c l e number increases the data points for the two samples pl o t progressively further apart. Early stages cannot be distinguished at the subspecific or Sample Number of Platforms Number of Complete Platforms Percentage of Complete Platforms Number of Ramiforms Number of Platforms: Number of Ramiforms F48 28 1Q 36% 12 2.33 : 1 F49 70.5 60 9% 175- 4.03 : 1 F52 95 25 26% 48 1.98 : 1 F53 174 40 23% 32 5.44 : 1 F54 190 37 19% 25 7.60 : 1 Tota l 1192 172 : 14% 292 4.08 : 1 Table 3. Counts and percentages df platforms and ramiforms of Neogondolella i n F48 - F54. 52 S t a t i s t i c a l Data D e r i v e d from Sample Subsets — S a m p l e F48 F49. F52 F53 F54 P a r a m e t e r s . Data \ s . N = 1, 2 N = 12, 12 N = 6, 5 N = 11, 7 N = 7, 9 h M 630 772 733 752 811 10 d e n t i c l e s a a 1 = 78.1 — 75 44 75 88 h M 1Q6Q 863 816 923 984 11 d e n t i c l e s a a 1 = 105.3 — 1Q8 46 105 102 10 d e n t i c l e s M 1 a a = 7.81 63.0 77.2 7 - 5 73.3 4.4 75.2 7.5 81.1 8.8 M 96.4 78.5 74.2 83.9 89.5 11 d e n t i c l e s a o 1 = 9.5? - 9.6 4.2 9.6 9.4 Table 4. S t a t i s t i c s from sample subsets (elements with 10 and 11 d e n t i c l e s ) . t - T e s t f o r and 1^/i Samples Compared 10 D e n t i c l e s 11 D e n t i c l e s t - s c o r e * P - v a l u e * * t - s c o r e * P - v a l u e * * 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 CF49+F52) -»• (F53+F54)! 0.58 0.28 2.75 0.005 Table 5. Values derived from t - t e s t s on sample subsets A Mo s i / 1 / % +.'1/N2 s = / - CJJ+N2 02 Nj_+ N 2- 2 0= standard deviation Efx - M N Efx = sum of the squares:of values for a character **= the p r o b a b i l i t y that the difference between two sample means could occur by chance ( i . e . i f Z-score = 1.2, then P-value = .115, thus there i s a 11.5% chance that the differe n c e occurred by chance alone). E • 1 5 . 1 4 r 1 3 - 1 2 a i n 0 ' 1 0 53 Figure 7a,b,c. Graphs showing the r e l a t i o n s h i p of the platform length to the num-ber of de n t i c l e s per element for F49, F54 and F96. o 9 0»(o/e . N . r o s e n k r a n t z i n . s u b s p . D ( F 0 6 ) ~'*N. i d a h o e n s i s n . s u b s p . A ( F 6 4 ) ( T o r 2 + s p e c l m e n s ) 7 a. F54 a n d F96 3 0 0 5 0 0 7 0 0 6 0 0 1 I O C 1 3 0 0 1 6 0 0 l e n g t h o f p l a t f o r m (^jm) e 1 «! o E ' e 1 5 . 1 4 7b. Regions defined by the c e n t r a l 3/4 of c l o s e l y spaced spec-imens for F49 and F54, CI a i 3 a o - 1 2 a i i i r o * 1 0 h E 1 6 • 1 6 • • 1 3 o Z 1 2 c a> •o 1 1 o i o • 0 E 7 6 a ° 8 E 3 c 7 6 ( c e n t r a l N . I d a h o s n a i s n . s u b s p . A F S 4 F 4 9 of s p e c i m e n s ) 3 0 0 5 0 0 7 0 0 0 0 0 1 1 0 0 1 3 0 0 1 6 0 0 , ' 9 l e n g t h o l p i s i f o r m ( ^ m ) > 09 9 + . am 7c. F49 N. i d a h o e n s i s n . s u b s p . A ( F 4 9 ) (1 o r 2 s p e c i m e n s ) 3 0 0 5 0 0 7 0 0 0 0 0 1 1 0 0 1 3 0 0 1 6 0 0 l e n g t h o f p l a t f o r m (^im) 54 s p e c i f i c l e v e l . A second i n t e r e s t i n g observation i s the increase i n mean length from 10 to 11 d e n t i c l e s f o r each sample. This increase i s approx-imately 85 um for F49 and F52 and about 170 um for F53 and F54 (Table 4). The importance of t h i s observation w i l l be discussed i n the section f o r length/number of d e n t i c l e s . Number of Denticles per Element (#) The mean number of d e n t i c l e s increases from F48 to F53 but decreases marginally for F54. If the number of d e n t i c l e s can be used as a "rough" guide to r e l a t i v e stages of development and i f platform overrepresentation i s d i r e c t l y r e l a t e d to ontogenetic stages, then the trends f o r these two parameters should roughly coincide. This i s not the case, however (Table 3, F i g . 6). This indicates that either length 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 or that platform overrepresentation does not explain the observations suggesting that the changes are rel a t e d to ev o l -ution..:.: The diff e r e n c e s recorded i n the number of de n t i c l e s upsection are i n s i g n i f i c a n t ( s i g n i f i c a n c e i s considered at 5%) according to z-scores (Table 2). Ratio of length to number of d e n t i c l e s (Ll/#) The comparison of two parameters together tends to produce some i n t e r -esting r e s u l t s with regards to possible evolutionary implications that could not be deciphered through analysis of the two parameters alone. This i s not sur p r i s i n g as animals or. t h e i r s k e l e t a l remains are not d i f f e r e n t i a t e d from one another by t h i s or that character but rather by the sum of many charac-ters (Burma, 1948). The r e s u l t s i n d i c a t e a decrease i n the r a t i o of length to number of d e n t i c l e s for F48 through F52 followed by an increase i n sam-ples F53 and F54 ( F i g . 6). Z-tests f o r the e n t i r e sample indi c a t e a f a i l -55 ure of s i g n i f i c a n c e even at the 10% l e v e l (Table 2). As a r e s u l t , the pos-s i b i l i t y that t h i s v a r i a t i o n could.be r e l a t e d to a process of random sam-p l i n g cannot be discounted. Further evidence suggests, however, that the observed trend may be s i g n i f i c a n t . The t - t e s t for Ll/# indicates that the data are s i g n i f i c a n t at the 5% l e v e l f o r those elements with 11 den t i c l e s from F49 to F54. Further upsection to F96,. a very sharp decrease i n the Ll/# r a t i o occurs which has a very high s t a t i s t i c a l s i g n i f i c a n c e (Table 5). Obviously, to proceed from,F54 to F96, at le a s t one more point of i n f l e c t i o n i s required to obtain, this,>decrease. What s t a r t s to become apparent i s a f l u c t u a t i n g mode of evolution for t h i s p a r t i c u l a r r a t i o . Combining data from Behnken (1975).with my data f o r length versus number of de n t i c l e s , these trends may be made gr a p h i c a l l y v i s i b l e ( F i g . 8). The evolution of Neogondolella. idahoensis (F54) to _N. serrata and f i n a l l y to N. postserrata-N. rosenkrantzi (F96) produces a n l i n i t i a l l y large decrease i n the length for an element of given d e n t i c l e number followed by a s l i g h t increase. These major i n f l e c t i o n points are thus s i g n i f i c a n t at the s p e c i f i c l e v e l . Perhaps s i m i l a r l y shaped but smaller points of i n f l e c t i o n l i k e that seen for F48 to F54 are s i g n i f i c a n t at the subspecific l e v e l . Another f a c t o r that becomes apparent from an analysis of the data i s that the more mature the element i s ( i e . the larger and more d e n t i c l e s i t has) the greater one's a b i l i t y to d i s t i n g u i s h between the population samples. In support of t h i s statement are t - t e s t s for Ll/# for 11 de n t i c l e s (Table 5). As was pointed out e a r l i e r , F49 to F52 increases i n length by 85 ym between 10 and 11 de n t i c l e s whereas F53 and F54 increase by 170 ym. Grouping these v a r i a t i o n s together and te s t i n g for the s i g n i f i c a n c e between F49 plus F52 and F53 and F54 provided the cl 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 for z-tests 6-5 * * 1 * * • 3 0 0 5 0 0 7 0 0 9 0 0 1 1 0 0 1 3 0 0 1 5 0 0 l ength of p l a t f o r m (jum) Figure 8. Graph showing the r e l a t i o n s h i p of the platform length to the number of d e n t i c l e s per element of N. idahoensis n.subsp. A, _N. serrata,. and N. postserrata ( i n chronological order). A f l u c t u a t i n g mode of evolution i s indicated by the decrease i n length for given d e n t i c l e number from _N. idahoensis n.subsp. A to N. serrata, followed by an increase i n length from N. serrata to N. postserrata. N_. idahoensis n.subsp. A region defined by F49 to F54 from Ellesmere Island whereas N_. serrata and N_. postserrata are based on data presented i n Behnken, 1975. 57 for t h i s i n t e r v a l (however, the only s i g n i f i c a n t t e s t s at 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 Figure 7 that the increase i n mean Ll/ / / 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 that for 10 to 11 between F49 and F54 as the f i e l d s represented by the data points become even more divergent. Length from t i p of cusp to fourth d e n t i c l e anterior of the cusp (L2) The f i r s t four d e n t i c l e s a n t e r i o r of the cusp tend to be more c l o s e l y spaced and more c i r c u l a r i n cross section than the remaining l a t e r a l l y com-pressed d e n t i c l e s . The maximum width of the element often occurs at about t h i s same point on the platform. Consequently, i t seemed appropriate to analyze t h i s parameter as i t could be valuable even for fragmental specimens. Except f o r F48 (based on the smallest sample) the trend i s one of increas-ing length upsection s i m i l a r to the increase f o r L l . ( F i g . 9). This s i m i l -a r i t y of trends.points to the value of t h i s parameter for samples where only fragmental specimens are a v a i l a b l e . Maximum width (Wl) The data here ind i c a t e a consistent increase i n mean width upsection ( F i g . 9). More discussion w i l l follow i n the section for Ll/Wl. Width at posterior end (W2) This parameter shows a consistent increase upsection s i m i l a r to that for Wl ( F i g . 9). The main reason for taking the two width measurements was for the determination, of an area function i n the posterior 1/3 to 1/2 of the d e n t i c l e . Height from t i p of cusp to base of flange (Hi) Dzik and Trammer (1980) found that t h i s parameter was very u s e f u l f o r d i s c r i m i n a t i n g T r i a s s i c Gondolella species. The height increases upsection 2 -r-3 W 1 x10" *\ 1 A * ± 1 1 I I I 50 250 B Hi 50 250 H 1 1 1 1 1 h i i i i 1 2 I V! W2*10 r2 B A 1 2 3 H 1- 1 1 I 1 1 1 h i u « 4 5 6 3D l IC *il IB w. JMhtSJkJ—I A 4 6 33E l>:-.v~-».v.'-'.' • B 4 5 6 7 H 1 1 1 1 1 1 1 1 1 r-3 4 5 6 I I —I 1 1 1 h 200 400 D A 4 8 200 400 H 1 I 1 1 1 +-9 10 11 L2(W,+W2)H 3 B 1 ° 4 „ m 2 jim 1 • 3 4 -j i • • 5 -i i_ 6 A 8 9 10 -j i _ _ 11 • Figure 9. Graphical representation of part of Table 1. Lines i n f u l l black show range of M + 2a M. Stippled pattern = 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 ) . 59 very c o n s i s t e n t l y . So f a r i t has been established that the two lengths, the two widths and the height measurements a l l increase, with some minor f l u c t u a t i o n s , from F48 to F54. What i s i n t e r e s t i n g , however, i s the trend i n the r a t i o s of these values as they a l l decrease upsection. Ll/Wl Ratio Despite the increases i n both length and width the Ll/Wl r a t i o dee-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 greater increase i n width r e l a t i v e to the increase i n length. This would be r e f l e c t e d i n a platform becoming more "square" i n shape rather than elongated and narrow. Z-test scores for t h i s trend i n d i c a t e that the changes from F48 to F53 and from F49 to F53 are s i g n i f i c a n t at the 5% l e v e l (even for a two-sided t e s t ) . I t i s impossible to t e l l whether t h i s trend fluctuates l i k e that for Ll/# or whether t h i s decrease i s translated through the e n t i r e section of Permian rocks discussed i n t h i s study. Two fragments of N . serrata(?) suggest a Ll/Wl ratio: of about 4 while the r a t i o f or N . rosenkrantzi varied between 4.0 and 4.7, averaging 4.3. A look at i l l u s -t r a t i o n s of specimens from the Great Basin of the USA suggests a s i m i l a r pattern of Ll/Wl r a t i o s for N . idahoensis (3.9), Ni.. serrata (3.3), N . post- serrata (3.6), N . b i t t e r i (3.6) and N . rosenkrantzi (4.0). (,Cl?ark'-and Behn-ken', 1979; Clark et a l . , 1979; Wardlaw and Collinson, 1979b). Admittedly, such comparisons, stand on shaky ground as the measurements are based on a few specimens preselected by the above authors and which do not take into account the v a r i a t i o n of Ll/Wl during ontogeny (although I did t r y to se-l e c t s i m i l a r intermediate to mature specimens from the figured specimens). Furthermore, samples from Texas (Behnken, 1975) suggested a continuous decrease from 4.3 to 3.05 f o r N . idahoensis to N . rosenkrantzi. These 60 _N. rosenkrantzi are more elongate than those from Ellesmere Island or the Great Basin. Probably the trends vary with environment and/or geography. Nevertheless there seems to be some merit i n suggesting a f l u c t u a t i n g trend at the s p e c i f i c l e v e l f or Ll/Wl s i m i l a r to that for Ll/# i n the Ellesmere Island samples of Neogondolella.. L l / H l Ratio Here again although both L l and HI increase fro'm F48 to F53 the r a t i o L l / H l decreases for the same i n t e r v a l . However, z-test scores i n d i c a t e that the d i f f e r e n c e between F48 to F53 i s i n s i g n i f i c a n t and could be the r e s u l t of random sampling. Despite t h i s , the trend i s s t r i k i n g l y consistent. Once again i t i s impossible to t e l l i f t h i s trend continues for the remainder of the section, nor are there data, to make inferences from. I n t u i t i v e l y , how-ever, the shorter length of N. serrata(?) and the high, robust cusps of N. n.sp. .B would indi c a t e decreased L l / H l r a t i o s while the short cusps of N. b i t t e r i n.subsp. C and N. rosenkrantzi n.subsp. D would indi c a t e a subse-quent increase i n the L l / H l value. L2(W1 + W2)% „ :—7 Posterior area 1 Q4 : Data for t h i s parameter indicates an increase i n t h i s area value for F48 to F53. Z-scores.::indicate that the d i f f e r e n c e between F48 and F53 i s s i g n i f i c a n t only at the 7%level whereas the d i f f e r e n c e between F49 and F53 i s s i g n i f i c a n t at the 2% l e v e l , even for a two sided t e s t . Measurements suggest that t h i s area value would be much l e s s f o r N. serrata(?) but great-er again for N. n.sp. B to N. rosenkrantzi n.subsp. D i n the Trold Fiord Formation. Once again a f l u c t u a t i n g mode i s suggested for t h i s parameter as w e l l . 61 Discussion of Evolutionary Trends and Concepts Evidence for the Ll/# r a t i o from population samples of N. idahoensis n.subsp. A from Ellesmere Island compared to data from population samples of N. serrata and N. postserrata (Behnken, 1975) from western USA in d i c a t e a f l u c t u a t i n g mode i n the evolution of these platform elements with respect to t h i s parameter. Furthermore, the r e s u l t s for N. idahoensis n.subsp. A suggest that t h i s f l u c t u a t i n g mode may be important at the subspecific l e v e l . Although not backed up by s t a t i s t i c a l data from large samples, conodonts upsection to N. idahoensis which are c l o s e l y a l l i e d to N. serrata, N. post-serrata and N. rosenkrantzi in d i c a t e a s i m i l a r trend to that seen for the comparison with Behnken's samples (1975) from western USA. In addit i o n s i m i l a r f l u c t u a t i n g trends seem apparent f o r several other parameters mea-sured from the Ellesmere Island samples. I t therefore seems reasonable, with a l l t h i s supporting documentation to suggest that t h i s f l u c t u a t i n g tendency i s the r u l e rather than the exception f or the evolution of Permian neogondolellids. Having proposed a mode for the evolution of these conodonts I have opened myself to the argument of i n t e r p r e t a t i o n of t h i s trend and; i n par-t i c u l a r , am^obliged to face the question of p h y l e t i c gradualism (of which many authors are proponents) versus punctuated e q u i l i b r i a (Eldredge and Gould, 1972; Gould and.Eldredge, 1977). As I pondered over t h i s problem I t r i e d to take heed of. Eldredge and Gould's warning (1972) that a l l obser-vation i s coloured by theory and expectation. My o r i g i n a l i n t e n t i o n for the s t a t i s t i c a l a n alysis was to see i f there was any quantitative d i f f e r e n c e between f i v e population samples that I could not, d i f f e r e n t i a t e q u a l i t a t i v e l y ( e s p e c i a l l y because of the high degree of v a r i a b i l i t y ) and then determine 62 i f any differences were s i g n i f i c a n t f o r the determination of subspecies. It was not u n t i l well into the analysis that I saw the p o s s i b i l i t y of purporting an evolutionary scheme. I did not then, and I do not now wish to get into a deep phi l o s o p h i c a l discussion of the merits of one or the other scheme. Rather I would l i k e to indicate my data and suggest an in t e r p r e t a t i o n and leave t h i s i n t e r p r e t a t i o n open to c r i t i c a l a n a lysis by other workers as part of the continuing process of evolutionary model syn-thesis f o r t h i s group of biota. I c e r t a i n l y cannot claim that my data would ever solve the problem of ...phyletic gradualism versus punctuated mode of organic evolution. The concept of p h y l e t i c gradualism states that new species evolve by the slow and continuous transformation of ent i r e populations r e s u l t i n g i n an unbroken gradation of f o s s i l forms (Eldredge and Gould, 1972). These unbroken gradational s e r i e s are r a r e l y ( i f ever) found because of the "sup-posed" imperfections i n the geologic record. The theory of "punctuated e q u i l i b r i a " states that new species evolve r a p i d l y from small, p e r i p h e r a l l y i s o l a t e d l o c a l populations ( a l l o p a t r i c speciation) r e s u l t i n g i n many breaks i n the f o s s i l record since the new species evolve i n an area remote from i t s ancestors (Eldredge and Gould, 1972). The hi s t o r y of evolution i s , therefore, not one of s t a t e l y unfolding, but a story of homeostatic e q u i l -i b r i a , disturbed only r a r e l y by rapid and episodic events of speciation ( i b i d . ) . Since 1972 when Eldredge and Gould f i r s t published t h e i r "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 evolution a few workers have come out to support i t while many have come out i n opposition, and contend that t h e i r research data indicates p h y l e t i c gradualism. This prompted Gould and E l -63 dredge to write a second paper i n 1977 to refute these claims and indic a t e the success of t h e i r model. In order to refute t h e i r punctuated e q u i l i -b r i a model a researcher must show gradational species l e v e l lineages well preserved over the f u l l span of an extensive geographic and temporal range. As Gould and Eldredge (1977) adequately point out most of the claims f o r phy l e t i c gradualism are based on l o c a l sections(not t h e i r f u l l geographic range) and short duration (too small a scale) and are therefore i n v a l i d as the data are i n s u f f i c i e n t f o r a test'. In defence of t h e i r openmindedness Gould and Eldredge (1977) do accept one case of gradualism as being very impressive. This case i l l u s t r a t e d the increase i n prolocular diameter of a verbeekinoid foraminifer i n 34 r e l a t i v e l y large samples spanning the Middle to Upper Permian, from southeast. Asia, southern China and Japan. As Gould and Eldredge (1977) state, "We.:are delighted with these r e s u l t s as we expect some countercases, e s p e c i a l l y among predominantly asexual forms". Their discussion of another g r a d u a l i s t i c case, that of Gingerich (1976) f o r Early Eocene mammals i n northern Wyoming, i s of p a r t i c u l a r i n t e r e s t to t h i s work. Gingerich (1976) claimed that species of the Eocene mammal Hyopsodus evolved i n a manner conforming to Cope's r u l e (increasing s i z e through'time)" based on the increase upsection of the logarithm of length times width of the f i r s t lower molar. However, t h i s o v e r a l l increase was achieved only a f t e r a number of f l u c t u a t i o n s (of nine descendant species, f i v e evolve toward smaller s i z e and only four to l a r g e r ) . Gould and Eldredge (1977) counter that Gingerich's s p l i t t i n g of lineages f i t s t h e i r model of punctuated e q u i l -i b r i a better. They found long segments of apparent s t a s i s within h i s sup-posedly g r a d u a l i s t i c sequences. Furthermore, they state the f l u c t u a t i n g pattern towards increase i n tooth s i z e confirms the most important i m p l i -cation of punctuated e q u i l i b r i a , that speciation i s e s s e n t i a l l y random with 64 respect to the d i r e c t i o n of a macroevolutionary trend (Wright's r u l e , Wright, 1967). Stanley (1975) wrote that macroevolutionary trends are not a r e s u l t of g r a d u a l i s t i c orthoselection, but a r i s e from a "higher l e v e l s e l e c t i o n " of c e r t a i n morphologies from a random pool, of speciation events produced by punctuated e q u i l i b r i a . According to.Gould and Eldredge (1977) the phylogeny of Hyopsodus af firms :;Wright' s r u l e where s i z e increase i n the e n t i r e clade arose from the d i f f e r e n t i a l , success of larger species i n a random subset of c l a d i s t i c events. I t i s my opinion that the phylogeny of Neogondolella idahoensis n.subsp. A to _N.. rosenkrantzi n.subsp. JJ equally supports the model of punctuated e q u i l i b r i a and affirms Wright's r u l e . The data displayed on Figure 8 i s based on N. serrata and _N. postserrata from West Texas (from Behnken, 1975) and on _N.. idahoensis n.subsp. A from north-ern Ellesmere Island. The material thus covers a large geographic area and a s i g n i f i c a n t portion of the temporal range of the Neogondolella serrata complex, that of the l a t e Lower Permian through Middle Permian (about 15 m i l l i o n years), both necessary pr e r e q u i s i t e s f o r an adequate t e s t . The data indicates an o v e r a l l increase i n number of d e n t i c l e s per unit length upsection but only a f t e r a f l u c t u a t i n g path where N. serrata has more den t i c l e s per.:unit length than the r e s u l t i n g descendent N. postserrata. N. rosenkrantzi n.subsp. D from Ellesmere Island f a l l s within the same f i e l d represented by data points for N. postserrata from West Texas. How-ever, a l l of these members of the clade have a larger value for d e n t i c l e number per u n i t length than that f o r the ancestral.form, N. idahoensis n. subsp. A. Having accepted a punctuated e q u i l i b r i a mode of evolution to ex-p l a i n the f l u c t u a t i n g pattern of d e n t i c l e number per. u n i t length I must also accept some of the other features of the model; namely, that between these spe c i a t i o n events the forms did not change, that i s they underwent a 65 period of s t a s i s . However, at f i r s t glance I cannot do t h i s , as some con-s i s t e n t trends were apparent i n the samples F48 to F54 studied quantita-t i v e l y i n d e t a i l (Table 1). Gould and Eldredge (1977) i n d i c a t e that the norm for a species during the heyday of i t s existence as a large population i s morphological s t a s i s , minor non-directional f l u c t u a t i o n i n form, or minor d i r e c t i o n a l change bearing no r e l a t i o n s h i p to pathways of a l t e r a t i o n i n subsequent daughter species. The nature of degree of t h i s minor change can be best understood by r e a l i z i n g that the r a p i d i t y of speciation i n such a model does not r e -quire the intermediate stage of a recognizable subspecies (Stebbins, 1977). My naming the studied populations represented by F48 to F54 as a new sub-species of Neogondolella idahoensis indicates that I f e e l recognizable changes do occur between the rapid speciation events and that these are s i g -n i f i c a n t at the subspecific l e v e l . The time represented from F48 to F54 (15 m of a 200 m section) may represent as much as one m i l l i o n years but i s probably more on the order of 500,000 years (assuming continuous sedimenta-t i o n and time scales for the Permian). Clark and Behnken (1979) indi c a t e that the average species duration for Permian neogondolellids i s 3.3 m i l l i o n years but that t h i s may vary from 2 to 10 m i l l i o n years (the l a t t e r f i g u r e for N. idahoensis - considered high by t h i s author). Even i f we assume the average duration df2 3 .3 m i l l i o n years for N. idahoensis, the time represent-ed between F48 and F54 i s . o n l y a small f r a c t i o n of t h i s temporal range. In other words even i f the change from F48 to F54 i s i n s i g n i f i c a n t , i t could, i f extended to the presumed f u l l range of the species, become s i g n i f i c a n t . Z-tests for the parameters L l , #, and Ll / / / indicated that differences between samples were i n s i g n i f i c a n t . However, when the samples were separ-ated into j u s t those elements with 11 d e n t i c l e s a t - t e s t indicated that 66 dif f e r e n c e s between F49 and F54 were very s i g n i f i c a n t . This s i g n i f i c a n c e alone could not be used, to d i f f e r e n t i a t e between subspecies but may i n d i -cate that d i f f e r e n c e s between e n t i r e samples, i f extended through greater temporal range, could become s i g n i f i c a n t . Although Gould and Eldredge (1977) do allow f o r some minor changes i t i s with the amount that many authors seem to be at odds with them. I would prefer to believe that between these rapid speciation events some gradualism does occur. However, Newell (1956) indicated that spurious "phyletic change" may a r i s e i n l o c a l sections by successive immigration of normal geographic variants responding to changing l o c a l environments. If the environment was changing i n a progressive man-ner (eg. shallowing during regression) then samples upsection would change i n one d i r e c t i o n with respect to a parameter. Given the genetic and physio-l o g i c a l complexity of any. population of organisms, many d i f f e r e n t ways of adjusting to a new factor of the environment are possible (Stebbins, 1977), suggesting that any change i n a l o c a l section with respect to environment need not be accompanied by a s i m i l a r consistent gradual change i n b i o t i c response. If the immigrants (normal geographic variants) respond i n many ways to environmental change then the r e s u l t s are u n l i k e l y to be consistent upsection 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 non-direc-t i o n a l ( s t a b i l i z i n g ) or d i r e c t i o n a l i s that of d i v e r s i f y i n g s e l e c t i o n where, i f environmental heterogeneity i s increasing over time, the response of the population 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 population breaking up into several d i f f e r e n t l y adapted subunits (Stebbins; 197.7). The v a r i a b i l i t y of those populations from F48 to F54 seems to be increasing as evidenced by the a l -most c o n s i s t e n t l y increasing standard deviation of the samples upsection. How these 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 ; i n t e r a c t 67 and whether the r e s u l t i n g gradation upsection can s t i l l be regarded as spurious rather than r e a l , i s d i f f i c u l t to say unequivocally. There are many complicating.factors, obviously. Perhaps easier to demonstrate would be whether or not the minor change i n form bears a r e l a t i o n s h i p to the evolution of subsequent daughter species. If one considers the i n t e r v a l represented between N. idahoensis and N. serrata the expected trends would be towards o v e r a l l decreasing s i z e and increasing d e n t i c l e number. The r e s u l t s for F48 to F54, which repre-sents part of t h i s j u s t mentioned i n t e r v a l , depict increasing length, width, height and posterior area upsection (exactly opposite the expectation) and o v e r a l l increase but f l u c t u a t i n g d e n t i c l e number (approximates the expected trends). These between-speciation trends could be regarded as random or chance events i n the developmental pathway because only one of the two trends bears any . r e l a t i o n s h i p to the speciation trends. It seems appropriate at t h i s time to compare the r e s u l t s of a study by Dzik and Trammer (1980) which i n many respects i s s i m i l a r to t h i s one. Their analysis i s the r e s u l t of study of 25 samples over about 23 metres of section from the Holy Cross Mountains of Poland which contain T r i a s s i c neogondolellids (gondolellids to them). Their r e s u l t s indicate a general decrease i n d e n t i c l e number and length but along a f l u c t u a t i n g path. They int e r p r e t t h e i r r e s u l t s as the r e s u l t of p h y l e t i c gradualism and not of punctuated e q u i l i b r i a . However, these r e s u l t s f a i l i n providing an ade-1 -. quate test since they represent a l o c a l section of short duration. In such a section one would not expect to see trends that are the r e s u l t of punc-tuated e q u i l i b r i a . Furthermore, the f l u c t u a t i n g path ( i f i t were over a larger time frame) could be better interpreted i n a punctuated e q u i l i b r i a 68 model. These r e s u l t s could be interpreted as "spurious" d i r e c t i o n a l phyle-t i c change because they are from a l o c a l s ection. Furthermore, a compli--cating f a c t o r previously discussed, that of increasing v a r i a b i l i t y , does not seem to be the case here. They in d i c a t e , however, that platform cono-donts i n the uppermost Muschelkalk of Germany ( s l i g h t l y younger than the Holy Cross Mountain specimens) represent i n morphology a progress along the trend d i r e c t i o n recognized i n the conodonts from the Holy Cross Mount-ains suggesting that t h e i r trends were not "spurious" and that the popular, tions of Gondolella inhabiting the Central European basin were evolving simultaneously and regardless of l o c a l f a c i e s changes. Despite the fa c t that the time frame represents a large f r a c t i o n of the Middle T r i a s s i c , the demonstrated evolution i s of three temporal subspecies. Two of these subspecies, are represented i n the l o c a l Holy Cross Mountain section - i n the uppermost and lowermost samples. As indicated e a r l i e r the trends at t h i s taxonomic l e v e l should be more gradual. These r e s u l t s give me some reason to accept my bias that the changes from F48 to F54 are r e a l at the subspecific rank and that gradual changes should be expected at t h i s taxo-nomic l e v e l . In summary, the r e s u l t s f or the Neogondolella species described from the Assistance and Trold Fiord Formations of northern Ellesmere Island, A r c t i c Canada compared to species from the Great Basin of the western USA, are best interpreted as the r e s u l t of evolution consistent with a punctuated e q u i l i b r i a model. The r e s u l t s for a small f r a c t i o n of t h i s i n t e r v a l seem to ind i c a t e that 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 i n gradual changes at the subspecific l e v e l . This should not. be considered i n opposi-t i o n to the punctuated e q u i l i b r i a model but regarded as a feature that en-69 hances the resultant changes during the rapid but ..punctuated speciation events. Just as Gould and Eldredge (1977) regard s t a s i s as r e a l so should gradual change between punctuated speciation events be regarded as a r e a l i t y at the subspecific l e v e l . This does not seem to. be an unreasonable s t a t e -ment when one considers the many varying evolutionary 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 are a number of problems with the designation of the genus Neogondolella that make i t s concept unclear. It was o r i g i n a l l y erected for forms that developed from the genus Spathognathodus (now Neospathodus) i n the Lower T r i a s s i c . Subsequently, Upper Carboniferous, Permian and Lower T r i a s s i c species have.been assigned to Neogondolella by many authors including a l l American authors. Kozur (1968) re t a i n s the genus Gondolella for a l l of these forms. The present controversy over the designation i s one of the opposing views between the European and North American "schools". The revised diagnosis (Sweet, 1970; f i d e . Z i e g l e r , 1973) includes conodont species i n which the s k e l e t a l apparatus comprises elements of a singl e morphologic type. " These elements, which are elongate, paired, and i n d i v i d u a l l y asymmetrical, have a terminal or subterminal p o s t e r i o r cusp; a median nodose or denticulate carina; and f i n e l y to coarsely p i t t e d , l a r g e l y unornamented, platformlike l a t e r a l extensions, which are joined p o s t e r i o r l y i n most species by a more or l e s s well developed brim that encloses 1 the posterior end of the carina. Underside of elements marked by a l o n g i t u d i n a l l y grooved keel that widens p o s t e r i o r l y to enclose .a p i t beneath the cusp (Zie g l e r , 1973, p. 127-128)." Kovacs and Kozur (1980) 70 suggest that t h i s diagnosis of Neogondolella Bender and Stoppel, 1965 i s i n s u f f i c i e n t to separate t h i s genus from Gondolella Stauffer and Plummer, 1932. I t appears that many of these features (posterior brim, ornamenta-tion) are v a r i a b l e within the phylogeny of t h i s group and ontogeny of i t s members (eg. smooth and p i t t e d platform of N. idahoensis versus serrated and p i t t e d platform of N. serrata and lack of s e r r a t i o n i n early ontogeny of N. postserrata). Kovacs and Kozur (1980) indi c a t e that the main argument for separation of the two genera i s the assumption that Neogondolella has a s i n g l e element apparatus (platform only) whereas Gondolella has a m u l t i -element apparatus (platform plus ramiforms). Because of the c o n f l i c t i n g opinion of a number of senior workers i t i s not c l e a r which argument has greater merit. Von B i t t e r and M e r r i l l ' s ' (19/7) suggestion that a Pennsyl-vanian Neogondolella had a multielement apparatus was rejected by Clark and Behnken (1979). One of the reasons for t h i s c o n f l i c t i n g opinion i s the phenomenon of platform overrepresentation; although the frequency v a r i e s i n any l o c a l section, i t generally increases from Carboniferous to T r i a s s i c . M e r r i l l and Powell (1980) have shown that t h i s platform overrepresentation i n Missourian Gondolella i s probably the r e s u l t . o f an ontogenetic s e r i e s of only ramiform apparatuses, to.ramiform plus platform, to apparatuses comprising platform only. In other words, they suggest a mechanism for t h i s low index of mutual occurrence which indicates-that a pure s t a t i s t i c a l a n a lysis could lead to serious mistakes i n the combination of conodont apparatuses. They agree with Kovacs and Koziir's (1980) opinion of a m u l t i -element apparatus for both of these genera but at the same time r e t a i n the designation of Neogondolella. Von B i t t e r (1976) suggests that the r e t i c u -l a t e d microstructure i s d i s t i n c t i v e ; covering much more of the o r a l surface 71 in. Neogondolella. Sweet (1970) indicated that i f Neogondolella contained elements of a si n g l e type then Neogondolella was fundamentally d i f f e r e n t from Gondolella. Evidence seems to indic a t e that the two genera are not fundamentally d i f f e r e n t but that minor differences do exist ( r e t i c u l a t i o n , v a r i a b l e but much l e s s ribbed or serrated ornament) to the extent that many authors ( i f not most: Kozur the notable exception) r e t a i n the genus Neogon-d o l e l l a . Furthermore, the o r i g i n and phylogeny of the genus remains clouded. The o r i g i n a l Neogondolella was erected as a form derived from Neospathodiis but Mosher (1968) ..indicates that N. mombergensis (the type species) i s not rel a t e d to Neospathodus making the o r i g i n a l diagnosis impractical. Szaniaw-s k i and Malkowski (1979) in d i c a t e that the evolutionary development of the Permian neogondolellids shows c l e a r l y that, they represent one phylogenetic branch and that the recognized ontogeny bears no r e l a t i o n s h i p to Neospatho-dus. Clark and Behnken (1979) in d i c a t e that N. b i s s e l l i , the ancestor of a l l Permian and younger species, occurs several hundred feet above Gondol-e l l a b e l l a with which i t bears l i t t l e morphologic s i m i l a r i t y . Movshovich et a l . (1979, f i d e . Kovacs and Kozur, 1980) in d i c a t e that "G". p r a e b i s s e l l i i s intermediate between G. b e l l a and "G". b i s s e l l i . There appears at pre-sent no cl e a r s o l u t i o n for a student attempting to decipher these reports i n terms of an adequate phylogeny. I t does seem c l e a r that more work i s necessary from an unbiased point of view to determine the s i g n i f i c a n c e of differences between Neogondolella and Gondolella. Only at t h i s point could the evolution of these forms be deciphered. U n t i l a revised diagnosis i s presented that i s widely acknowledged and refutes the genus Neogondolella, t h i s author r e f e r s h i s gondolelliform e l e -72 merits to species of Neogondolella as diagnosed i n Ziegl e r (1973). Using t h i s diagnosis indicates that descriptions for the species at hand assume a s i n g l e element-type apparatus. For t h i s reason the platform elements of Neogondolella w i l l be discussed separately from the ramiforms. Recognizing that the ramiform elements could belong to multielement "Neogondolella" apparatus a dual taxonomic nomenclature i s adopted f o r these elements as follows: Multielement species - Form species. This method of de s c r i p t i o n i s somewhat unsatisfactory but i t does r e f l e c t the present controversy and the desir e by the author that the question be resolved i n the near future. Should the genus be found to have a multielement apparatus then the form' species name should be placed i n synonymy while i f the genus i s found to have.a s i n g l e element apparatus then the multielement species name should be dropped. Description of these ramiform elements w i l l follow that of the platforms of Neogondolella. As the platform elements evolved r a p i d l y and are consequently s t r a t i -g r a p h i c a l l y very important they are described i n d e t a i l , whereas the rami-1 form elements evolved slowly and are, as a result, of l i t t l e value s t r a t i -g r a p h i c a l l y and thus are not described i n d e t a i l . The platform elements of Neogondolella species are described i n chron-o l o g i c a l order. Two other genera, including Anchignathodus minutus and Neostreptognathodus p r a y i are discussed f i r s t because of t h e i r a s s o c i a t i o n with the oldest Neogondolella platform species (N. idahoensis). These two genera are only b r i e f l y described as they are unimportant -in the area owing to t h e i r slow evolutionary rates and very rare occurrence, r e s p e c t i v e l y . In summary, the order of appearance w i l l be Anchignathodus minutus, Neostreptognathodus' p r a y i , platform elements of various Neogondolella spe^ c i e s , and f i n a l l y the ramiform elements . 73 Systematics Genus .ANGHIGNATHODUS. Sweet, 1971 Type species Anchignathodus minutus ( E l l i s o n ) , 1941 ASGHIGNATHODUS MINUTUS ( E l l i s o n ) , 1941 PI. 1, f i g s . 3-6. Spathodus minutus E l l i s o n , 1941 Occurrence: Lower Assistance Formation, Hamilton Peninsula section (F48, F49, F52, F53, F54, F63 arid F75). Description: This element possesses a short, t h i n , l a t e r a l l y s t r a i g h t to s l i g h t l y curved blade about three times as long as wide and with s i x to nine 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 posterior t o . the cusp and zero to three short d e n t i c l e s a n t e r i o r to the cusp. The den-t i c l e s are o f f s e t abruptly to the cusp which i s large and tr i a n g u l a r i n o u t l i n e . The basal c a v i t y of the blade i s broadly f l a r e d i n the mid region, e s p e c i a l l y under the d e n t i c l e s posterior to the cusp. The cav i t y reduces to a narrow groove at both the anterior and posterior ends of the aboral surface. The deepest point of t h i s basal c a v i t y i s below the f i r s t or sec-ond d e n t i c l e posterior to the main cusp. Discussion: Representatives of t h i s species range from Chesterian (Late Mississippian) to Roadian (early Medial Permian) i n age (Behnken, 1975; Zieg l e r , 1973). They are d i f f e r e n t i a t e d from the younger A. t y p i c a l i s by the abrupt o f f s e t i n l a t e r a l p r o f i l e posterior to the cusp as opposed to a gradual diminution of the l a t e r a l p r o f i l e . Genus NEOSTREPTOGNATHODUS'- Clark, 1972 Type species Streptognathodus s u l c o p l i c a t u s (Youngquist, Hawley and M i l l e r , 1951) 74 NEOSTREPTOGNATHODUS PRAYI Behnken, 1975 P l . 1, f i g s . 1,2. Occurrence: Assistance Formation, McKinley Bay section (F100). Description: This element consists of a subsymmetrical, p o s t e r i o r l y pointed platform with c l o s e l y spaced, s u b p a r a l l e l , transverse ridges on the o r a l surface which extend almost to completely across the medial groove. Discussion: No anterior free blades were observed. The two fragments appear to represent forms intermediate i n ontogeny, following descriptions by Behnken (1975). According to Wardlaw and Collinson (1979a) and Clark et a l . (1979) the youngest occurrence of t h i s species i s Latest Leonardian. Genus NEOGONDOLELLA" Bender and Stoppel, 1965 Type species Gondolella mombergensis (Tatge) NEOGONDOLELLA IDAHOENSIS (Youngquist,-Hawley and M i l l e r , 1951) subsp. indet. " - . P l . 1,. f i g s . 7-13. Gondolella idahoensis Youngquist, Hawley and M i l l e r , 1951 Gondolella phosphoriensis Youngquist, Hawley and M i l l e r , 1951 Occurrence: Assistance Formation, McKinley Bay section (F100). Discussion: These specimens are very s i m i l a r to N. idahoensis n.subsp. A except that no serrations or d i s t i n c t v a r i e t i e s were recognized. This i s p a r t l y owing to the low number of specimens and poor preservation (recry-s t a l l i z e d ) . . For these reasons and because of i t s occurrence with Neostrep-tognathodus p r a y i (suggesting a s l i g h t l y older age from Neogondolella idaho-ensis n.subsp. A) a subspecific determination was not made. Their descrip-t i o n i s s i m i l a r to that of N. idahoensis n.subsp. A (except for the varied", t i e s ) which follows. NEOGONDOLELLA IDAHOENSIS. n.subsp. A PI. 2, f i g s . 9-19; P i . 3; Pi.' 4; P i . 5. Occurrence: Lower Assistance Formation, Hamilton Peninsula section (F48, F49, F52, F53, F54).;-Diagnosis: This subsymmetrical unit has a carina composed of 6 to 16 de n t i c l e s (average 10 to 11) which are d i s c r e t e i n early growth stages, becoming p a r t i a l l y fused and f i n a l l y completely fused i n gerontic forms. The platform elements are very v a r i a b l e , ranging from long and slender forms to robust and serrated.. The maximum width t y p i c a l l y occurs at some point i n the posterior h a l f , a f t e r which the element tapers gradually and f i n a l l y r a p i d l y i n the anterior h a l f . Except for the anterior t i p the platform 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 generally absent from the well developed furrows and carina, i s present on these i n some mature robust forms. During ontogeny the keel on the aboral side changes from narrow and high, terminating i n an oval loop to wide and low with a terminal t r i a n g u l a r loop. The cusp i s large, erect and i n c l i n e d p o s t e r i o r l y while the four den-t i c l e s a n t e r i o r to i t are low, node-like, and c l o s e l y spaced. The remain-der of the de n t i c l e s increase i n height and are progressively compressed a n t e r i o r l y . Description: A. Juvenile - The element i s small (length = 350 to 800 um; maximum width = 100 to 180 um), subsymmetrical, and s l i g h t l y arched. The t h i n platform has i t s l a t e r a l margins upturned more i n the c e n t r a l portions of the element than at either end. The p o s i t i o n of the maximum width,;:thoug v a r i a b l e , i s generally s l i g h t l y a n t e r i o r of the.posterior t i p . The l a t e r a l margins are su b p a r a l l e l to only s l i g h t l y tapering a n t e r i o r l y f or much of t h e i r length corresponding to the p o s i t i o n of the r e t i c u l a t e d micro-orna-ment. The remaining 1/3 of the length bears l i t t l e or no r e t i c u l a t e pat-tern and tapers much more r a p i d l y than the posterior 2/3. The r e t i c u l a t e ornament i s r e s t r i c t e d to the o r a l surface on the edges of the platform and i s absent on both the carina and the furrows l a t e r a l to i t . The carina generally consists of 6 to 9 l a t e r a l l y compressed tri a n g u l a r and nodose d e n t i c l e s . The posterior cusp i s higher than a l l other 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 oval i n crossnsection, i n c l i n e d p o s t e r i o r l y but with a slight, a n t e r i o r l y d i r e c t e d curvature, and situated v a r i a b l y be-hind the posterior margin of the platform. The cusp thus extends as a posterior f ree blade i n the e a r l i e s t recognized stages but t h i s feature i s quickly l o s t as the platform extends to the posterior edge of the cusp and f i n a l l y forms a brim posterior to the cusp i n l a t e r ontogenetic stages. The next three or four d e n t i c l e s anterior to the cusp are small and s l i g h t -l y compressed but are more generally nodose compared to the r e s t of the ca r i n a . This feature remains i n a l l ontogenetic, stages and would seem to be of major genetic s i g n i f i c a n c e and important to the diagnosis. The r e -maining d e n t i c l e s increase i n si z e r a p i d l y and become compressed, i n c l i n e d 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 anterior t i p i s approached. In the e a r l i e s t stages the platform does not reach the anterior portion where the carina extends as a free blade, but i n l a t e r stages i t lengthens to encompass t h i s free blade. The aboral surface bears a narrow and very high keel terminating p o s t e r i o r l y as an elevated oval loop. The basal groove i s very narrow, extends the e n t i r e length of. the keel and terminates p o s t e r i o r l y as an elongated, narrow and.curved p i t . The crimp i s very wide, about 3/4 of the platform width, and smooth In contrast to the ornamented o r a l surface. B. Intermediate - As the element increases i n length the platform becomes thicker and wider. As i s the r u l e rather than the exception for these cono-donts s the values and r a t i o s f o r the. measured parameters are extremely v a r i a b l e . This v a r i a b i l i t y i s e s p e c i a l l y large f o r L l / H l and Ll/Wl r a t i o s . In general, however, the length of the intermediate element ranges between 800 and 1100 um and the width between 180 and 220 um. The p o s i t i o n of max-imum width i s generally 1/3 to 1/4 the length from the cusp to the a n t e r i o r . The platform extends from the posterior edge of the cusp to. the anterior t i p : no free blades exist at t h i s stage. I t i s at t h i s stage that v a r i e -t i e s s t a r t 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 discussed, i n d e t a i l . Generally, taper of the platform i s gentle towards the anterior for much of the length but increases f o r the anterior 1/3 to 1/4. The r e t i c u l a t e pattern extends to the anterior t i p but is. s t i l l r e s t r i c t e d to the thickened margins of the platform. The carina consists of 10 to 11. l a t e r a l l y compressed, t r i a n -gular, nodose d e n t i c l e s . The;posterior,.cusp.(is s i m i l a r to that described for the juvenile.stage except that i t i s l a r g e r . The next 3 to 5 d e n t i c l e s anterior to the cusp ( i n most specimens i t i s 4 denticles) are low, roughly c i r c u l a r i n cross section and node-like. The remaining d e n t i c l e s increase i n height and are progressively compressed a n t e r i o r l y . A l l d e n t i c l e s are e n t i r e l y d i s c r e t e at t h i s stage. . The aboral surface bears a wider and low-er keel (as compared, to j u v e n i l e stages) which terminates p o s t e r i o r l y as an .elevated oval to s l i g h t l y square shaped loop. The basal groove i s s i m i -l a r to that i n j u v e n i l e stages, but a l i t t l e less, narrow. The crimp i s narrower, 2/3 to.3/4 of .the platform width, and smooth. C. Mature - The v a r i a b i l i t y of form for these conodonts i s very high, as was mentioned above, and becomes accentuated i n the mature stages. It i s at 78 these stages that d i f f e r e n t v a r i e t i e s can. be distinguished. Some of the v a r i e t i e s are so d i s t i n c t i v e . t h a t , i f found alone, they might well be des-cribed as d i f f e r e n t speciesj however, i t seems that the v a r i a b i l i t y i s ac-t u a l l y that within a s i n g l e subspecies. The graphical evidence (Figs. 10-14) f a i l s i n a l l cases to i s o l a t e these, v a r i e t i e s from each other. Those graphical plots that to a c e r t a i n degree separate the two extreme v a r i e t i e s , namely the posterior area versus length/denticle number r a t i o and the post-e r i o r area versus length (Figs. 12-14), do so with some overlap between themselves and a large number of intermediate forms. It seems apparent that such a feature 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 si n g l e gene pool. An anisometr.ic type of growth, indicated by the graph of area versus length ( F i g . 12), demonstrates q u a n t i t a t i v e l y that shape of the element changes during ontogeny: a fa c t alluded to throughout the des-c r i p t i o n of t h i s species. The two very d i f f e r e n t v a r i e t i e s present for t h i s subspecies w i l l be described as the g r a c i l i s v a r i e t y and the robustus v a r i e t y . A number of forms that cannot q u a l i t a t i v e l y be separated into one or another of these v a r i e t i e s w i l l be referred to as the intermediatus v a r i e t y . The g r a c i l i s v a r i e t y , so-named because of i t s very long and slender platform, i s very s i m i l a r to the species Neogondolella g r a c i l i s Clark and Ethington, 1962. The v a r i e t y robustus resembles the species Neogondolella serrata Clark and Ethington, 1962.except f o r the much, larger length per d e n t i c l e number and the lack of ridges associated with the serrate margin. The intermediatus v a r i e t y most resembles the species Neogondolella idahoensis Youngquist, Hawley and M i l l e r , 1951 leading to the new subspecies being referred to N. idahoensis rather than N. g r a c i l i s or N. serrata. I t was considered that r e f e r r i n g the populations i n question to a new s p e c i f i c rank required unsub-•g • r • r • r . g N. idahoensis n.subsp. A(F49) • one specimen • two specimens i j i i i ' 400 600 800 1000 1200 1400 L e n g t h - L1 Figure 10. Graph showing the r e l a t i o n s h i p of Length/Height to Length of the platform f o r F49. r= var. robustus g= var. g r a c i l i s , the remainder are var. intermediatus +r N. idahoensis n.subsp. A(F49) • one specimen • two specimens -L 400 600 800 1000 L e n g t h - L 1 1200 1400 Figure 11. Graph showing the r e l a t i o n s h i p of Length/Width to Length of the platform for F49. r= var. robustus g= var. g r a c i l i s , the remainder are var. intermediatus. 81 14 13r 12 11 10k < LU CC < tr o E LU H CO 2* / / i v / / a 1 A * ' / / / -I L. -1 l _ 3 4 5 6 7 8 9 10 11 12 13 14 15 16 LENGTH x*f 2 Figure 12. -.Graph showing the r e l a t i o n s h i p of Posterior Area to Length of the platform for F53. r=~var. robustus g= var. g r a c i l i s , the remainder are var. intermediatus 82 / / / I 1 I I i I I I I I i -r / /• / / • / / / •/. . / / / / / ' ' ' / :9 / / 9 / / . *.-r / -g ' . . . »/.g / -g / y a /• • •• / / / -I 1 1 1 1 1 1 1 I 2 3 4 5 6 7 8 . 9 10 11 12 13 14 15 16 LENGTH x 10"2 Figure 13. Graph showing the r e l a t i o n s h i p of Posterior Area to Length of the platform for F49. r= var. robustus g= var. g r a c i l i s , the remainder are var. intermediatus. tg- a •g • N. idahoensis n.subsp. A(F49) — i i i I i ' 0 60 70 80 90 100 110 L e n g t h / n u m b e r of d e n t i c le s - L 1 / * Figure 14. Graph showing the r e l a t i o n s h i p of Posterior Area to Length/number of d e n t i c l e s for F49. r= var. ;robustus g= var. g r a c i l i s , the remainder are var. intermediatus. 84 stantiated taxonomic. s p l i t t i n g . I f quantitative methods indicated a separ-a t i o n of t h i s form from older N.. idahoensis. similar, to the separation f o r _N. serrata and N. postserrata (Behnken,.1975 - he r e f e r r e d to N. postserra-t a as N. serrata postserrata which was subsequently elevated to s p e c i f i c rank) then perhaps t h i s new subspecies could be l a t e r elevated to s p e c i f i c rank. I t i s possible that these v a r i e t i e s . i f separated as.peripheral i s o -l a t e s could, following speciation and migration, lead to populations more r e l i a b l y "described as _N. g r a c i l i s . o r _N. serrata. As well as these three named morphologies a. number of "experiments" or. unusual forms, are mentioned below. Perhaps, as these unusual forms are few i n number, they could be considered as mutations, that were not p a r t i c u -l a r l y advantageous; at l e a s t one, however, has a platform shape s i m i l a r to that of the T r i a s s i c species N. c o n s t r i c t a . N. idahoensis n.subsp. A'Var. g r a c i l i s The platforms are t y p i c a l l y long and slender ( L l =850 to 1200 ym; Wl = 160 to 220 ym), subsymmetrical, and gently arched. The platform ex-tends the f u l l length of the element and i s f l u s h with the p o s t e r i o r edge of the cusp. The l a t e r a l edges of the platform are upturned, e s p e c i a l l y i n the c e n t r a l regions. The l a t e r a l margins of the p o s t e r i o r h a l f of the platform are s u b p a r a l l e l to p a r a l l e l and taper gently i n the a n t e r i o r h a l f . The posterior margin i s gently rounded and with the r e t i c u l a t e d .portion tapering to the cusp. The r e t i c u l a t i o n ornament tapers on the l a t e r a l margins i n the same fashion as the platform i t s e l f tapers but i s absent i n the most anterior portions. The r e t i c u l a t e pattern i s absent on both the d e n t i c l e s and furrows. Some f a i n t ridges were observed on the cusp of a couple of specimens - these may represent the very e a r l y formation of r e t i -85 culae. The carina t y p i c a l l y comprises 12 or 13.discrete d e n t i c l e s , the anterior, four of which are low and nodiform, a l l of which are connected by a noticeable crease or mid-line. No a n t e r i o r serrations have been noted on t h i s v a r i e t y . N. idahoensis n.subsp. A var. robustus As the name indicates t h i s v a r i e t y i s t h i c k e r , wider and generally more stout or robust compared to var. g r a c i l i s . The length of the element i s generally between 800 and 1250 um and the width between 200 and 270 um. 2 4 This r e s u l t s i n posterior areas averaging 6.2 u n i t s (um X 10 )', consider-ably more than the 4.4 unit average for. the g r a c i l i s v a r i e t y . The platform extends the f u l l length o f the element, i n some cases forming a minor posterior brim but normally f l u s h with the rear edge of the cusp. The margins of the platform are s u b p a r a l l e l for the posterior 3/10, increase i n width over the next 2/10, and f i n a l l y taper over the anterior h a l f : slowly at f i r s t and more r a p i d l y close to the t i p . The posterior margins are quite square as opposed to the rounded g r a c i l i s v a r i e t y . The r e t i c u ^ l a t e d pattern p a r a l l e l s the platform margin, and i s missing on the anterior -most edges where the margin tapers most r a p i d l y . The r e t i c u l a t e pattern migrates onto the furrows and the d e n t i c l e t i p s i n many specimens. This feature seems to be r e l a t e d to early fusion of the d e n t i c l e s - a feature representative of gerontic i n d i v i d u a l s . The ridges that c o n s t i t u t e the 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 middle of i t s range, fl a t t e n e d but d i s t i n c t on the outer edge, and fading and elongate towards the furrows (PI. 5, f i g . 1). . Denticles (generally 11 to 13)except for some fusion of the posterior.ones,conform to e a r l i e r d e s c r i p t i o n s . A number of specimens exhibit a serrate platform margin i n the a n t e r i o r 1/3. The serrations are v a r i a b l y developed but generally weaker than those described 86 by.Clark and Behnken (197.9) for N. serrata and younger Permian conodonts and without the ridges accompanying the serrations as. i n N. serrata. The keel i s wide and .low and terminates i n an extensive t r i a n g u l a r loop. The basal groove i s wider than i n more j u v e n i l e forms and terminates i n the basal p i t . The smooth crimp t y p i c a l l y covers 6/10 to 2/3 of the aboral surface. !N. idahoensis n.subsp. A var. intermediatus The form of the carina, p o s i t i o n of the r e t i c u l a e , and development of the aboral surface are s i m i l a r to those described f o r the other v a r i e d , t i e s . The platform shape i s intermediate between that f o r the g r a c i l i s and robustus v a r i e t i e s with the maximum width generally corresponding to the p o s i t i o n of the fourth d e n t i c l e anterior to the cusp. Areas i n the poster-i o r region average about 5.0 u n i t s . The platform tapers more or l e s s gra-d u a l l y from the maximum width to the anterior t i p . Anterior serrations are present but l e s s .common ' than i n the robustus v a r i e t y . The main .differences i n platform shape are the average maximum width, and the manner, p o s i t i o n , and degree of taper. These features have been described i n d e t a i l f or the extreme v a r i e t i e s , whereas i t seems adequate to state that the representatives of var. intermediatus form the c e n t r a l part of a continuous gradation and t y p i f y the population means. Introduction to the unusual v a r i e t i e s of N. idahoensis n.subsp. A Those forms that are unusual in.terms of t h e i r platform shape are generally recognized at the mature to gerontic stages of development and do not show observable ontogeny. These forms comprise only a small f r a c -t i o n of any populatiom: sample. N. idahoensis n.subsp. A var. constrictus This v a r i e t y has a cons t r i c t e d platform margin i n the posterior of the element. The platform i s rounded i n the posterior and tapers s l i g h t l y a n t e r i o r l y for about 1/4 of i t s length where i t widens r a p i d l y to i t s widest point about 3/8 of i t length. From t h i s point the platform tapers normally for the subspecies. The i l l u s t r a t e d specimen i s a gerontic form and has fused posterior d e n t i c l e s ( P i . 4, f i g . 7). Var. c o n s t r i c t u s has also been recognized i n specimens with a l l d e n t i c l e s d i s c r e t e . This p l a t -form shape i s c h a r a c t e r i s t i c of N. 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 posterior margin. N. idahoensis n.subsp. A var. lobatus This form has a very wide and t h i c k platform o v e r a l l that i s s i m i l a r , except.;for the posterior end, to N. rosenkrantzi. The posterior margin of the platform i s t r i - l o b e d : one lobe surrounding the cusp and"separated by furrows from the two lobes forming the p o s t e r o - l a t e r a l margins of the element. The.furrows are wider-and s l i g h t l y deeper than t h o s e i n . similar- : N. rosenkrantzi n.subsp. :B and ate not-directed posfeero-laterally as welli• The d e n t i c l e s increase only s l i g h t l y i n height a n t e r i o r l y and are not d i s -t i n c t and node-like as i s generally c h a r a c t e r i s t i c f o r the subspecies. The cusp i s also more compressed then normal and not directed p o s t e r i o r l y but rather s t r a i g h t upwards. Discussion: Juvenile forms of N. idahoensis were described f o r the f i r s t time by Szaniawski and Malkowski (1979) from the Kapp S t a r o s t i n Formation of Spitsbergen. The ontogeny described by these authors i s very s i m i l a r to that for N. idahoensis n.subsp. A, except that the j u v e n i l e carina has 6 to 9 d e n t i c l e s rather than 8 to 10 for the Kapp S t a r o s t i n specimens. Szaniawski and Malkowski (1979) stated that the ontogeny f o r N. idahoensis was very s i m i l a r to that of N. b i t t e r i (Kozur) as described by Clark and Behnken (1971) and concluded that these two species bore a close r e l a t i o n -88 ship. The j u v e n i l e forms, of N . idahoensis n.subsp. A are very s i m i l a r to those of N . rosenkrantzi n.subsp. D...., Furthermore j u v e n i l e forms of N . post-serrata as i l l u s t r a t e d i n Behnken (1975) are also very s i m i l a r to those of the other species j u s t mentioned. There seems to be good reason to believe that the close s i m i l a r i t y of the j u v e n i l e forms described above i s the re.-, s u i t of close phylogenetic a f f i n i t y f o r a l l those species belonging to the serrata complex (Clark and Behnken, 1979) and including N . idahoensis n.sub-sp. A. Clea r l y , species cannot be determined on the basis of j u v e n i l e ma^ t e r i a l alone. The differences between members of N . idahoensis n.subsp. A from F49 to F54 are d i r e c t l y proportional 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 ontogenetic development and i l l u s t r a t e s the need for intermediate and mature forms be-fore d i f f e r e n t i a t i o n of subspecies and species can be attempted. Mature specimens d i f f e r l i t t l e from specimens previously referred to N . idahoensis. Most workers a t t e s t to a high v a r i a b i l i t y i n shape and i n length to width r a t i o although none have attempted to quantify t h i s . V a r i a -b i l i t y i s a c h a r a c t e r i s t i c of N . idahoensis n.subsp. A (Figs. 10, 11). Perhaps the most c l o s e l y comparable specimens that I have seen are those .'. i l l u s t r a t e d and described by Szaniawski and Malkowski (1979) from Spitsber-gen. As well as _N. idahoensis (137 fragments) these authors described _N. c f . N . g r a c i l i s (3 fragments) and IT. sp. A (1 specimen). In-the 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 these fragments would be included i n a si n g l e species. The N . c f . N . g r a c i l i s i s s i m i l a r to the Ellesmere Island var. g r a c i l i s whereas the N . sp. A i s s i m i l a r to Var. lobatus. These spec-imens d i f f e r from H. idahoensis n.subsp. A i n terms of the lack of anterior serrations, the number of d e n t i c l e s (8 to 13 d e n t i c l e s compared to 6 to 16), and i n the manner of tapering (widest point close r to posterior t i p ) . 89 Their specimens appear closer to the type specimens for the species and are probably s l i g h t l y older than N. idahoensis n.subsp. A. Clark and Mosher (1966)' regarded N. phosphoriensis Youngquist, Hawley and M i l l e r with i t s posterior r i d g e - l i k e carina, as a l a t e r growth stage of N. idahoensis where the posterior carina has become fused. In the E l l e s -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 gerontic forms only. As well, the fusion of the carina i s shown to be gradational ( P l . 4, f i g s . 5, 6, 10). These observations most c e r t a i n l y lend further support to Clark and Mosher's conclusion that N. phosphoriensis be placed i n synonymy with N. idahoensis. The basal loop shows a development from a high elongate oval to rounded shape i n j u v e n i l e and intermediate forms to a low, large, t r i a n g u l a r shape i n mature and gerontic forms. Observation of the growth lamellae i n a few specimens ( P l . 5, f i g s . 9, 10) shows that some of the l a t e r a l ' t r a c e s of the lamellae are truncated at the p o s t e r i o r of the loop. The ^ truncation i s apparently caused by resorption as described by Muller and Nogami (1972) which, according to these authors, i s a common phenomenon'for the Conodont-iformes. It resulted i n squaring-'off the posterior of the loop such that subsequent regeneration of the loop led to a t r i a n g u l a r shape. Resorption, then, seems to be an important phenomenon within the ontogeny of t h i s con-odont. Muller and Nogami (1972) conclude that the conodont element, besides having the function to support a t i s s u e , may also have served as an organ for the temporary deposition of phosphatic,substance, which might l a t e r be u t i l i z e d to form another element i n the same animal. One cannot help but speculate whether t h i s resorption phenomenon observed i n the platform e l e -ment of Neogondolella also occurred i n the ramiforms ( i f a multielement 90 apparatus indeed existed) r e s u l t i n g i n a platform-only apparatus (as sug-gested by M e r r i l l and Powell, 1980): the excess phosphate being used to form the large, thick-margined mature and ger.ontic platform elements. NEOGONDOLELLA SERRATA(?) (Clark and Ethington, 1962) PI. 6, f i g s . 7-9. Gondolella serrata Clark and Ethington Gondolella nankingensis Ching, 1960 Occurrence: Upper Assistance Formation, Hamilton Peninsula section (F63, F73) . Description: This designation i s based only on fragmental specimens. Es-timates of length were made by taking into consideration the gentle taper. Specimens that would be regarded as intermediate i n ontogeny have an e s t i -mated length of 400 to 580 ym and a maximum width of 100 ym. Although the anterior d e n t i c l e s were not observed, the t o t a l number of d e n t i c l e s f o r these intermediate forms i s on the order of 11. The four d e n t i c l e s anterior to the cusp are sharp s fused at t h e i r bases, l a t e r a l l y compressed and not d i s t i n c t from the other d e n t i c l e s as i n N. idahoensis n.subsp. A. The cusp i s high'and l a t e r a l l y compressed, e s p e c i a l l y the anterior h a l f . The p l a t -form i s gently tapering, l a t e r a l l y upturned, arched and rounded at the post-e r i o r end. The lower surface of the platform i s smooth and bears a high and r e l a t i v e l y narrow keel. The r e t i c u l a t e " p a t t e r n on the upper surface i s sharp, i r r e g u l a r i n shape and r e s t r i c t e d to the l a t e r a l margins as i t fades quickly towards the carina, reaching the l a t t e r only at the p o s i t i o n of the cusp. Discussion: These specimens d i f f e r from the older specimens of N. idahoen-s i s n.subsp. A i n t h e i r smaller si z e and they do not have the four nodiform d e n t i c l e s .distinct from the others anterior to the cusp. Reduction i n s i z e 91 at comparable growth stages has been observed.in the evolution of N. i d a -hoensis to N'. serrata i n other regions. Another c h a r a c t e r i s t i c of N. ser-rata that i s present i n these specimens i s the fusion of d e n t i c l e s i n i n t e r -mediate rather than gerontic stages of growth. Since anterior fragments of the platform were not observed i t i s impossible to assess whether the anterior serrations, diagnostic of the species and providing the d e r i v a t i o n of the name, are present. According to the figures mentioned above, the values for L l and Wl i n d i v i d u a l l y average about two standard deviations smaller than the mean for N. idahoensis n.subsp. A.- The r a t i o Ll/number of d e n t i c l e s i s more than two standard deviations l e s s than the means.for N. idahoensis n.subsp. A and almost two standard deviations l e s s than that for N. rosenkrantzi n. subsp. _D. Despite the fa c t that these figures are based on only very few specimens t h e i r s i g n i f i c a n t departure from the means for specimens lower i n the section suggest that they are indeed a d i f f e r e n t species. Furthermore, since they share some of the features of N. serrata they are assigned to ; that taxon but are more probably intermediate between N. idahoensis n.subsp. A and N. serrata. The specimens also plot i n or near the f i e l d of''data points for L l versus number of d e n t i c l e s of N. serrata as defined by Behn-ken (1975). The assignment i s l i s t e d as i n d e f i n i t e because of the poor preservation and paucity of specimens. The i r r e g u l a r r e t i c u l a t e pattern on the upper platform surface also seems worthy of further discussion. 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 microstructure of an intermediate and a mature N. postserrata. The microstructure i s very r e g u l a r l y shaped and arranged i n l i n e a r rows (ordered) i n the intermediate form whereas microstructure of 92 the mature form i s i r r e g u l a r and arranged i n a roughly sinuous manner ( d i s -ordered) . Similar ontogenetic v a r i a t i o n , although not as marked, was ob-served i n specimens of N. idahoensis n.subsp. A. Perhaps the feature i s i n d i c a t i v e of r e l a t i v e maturity. This would suggest that the small s p e c i -mens ref e r r e d here to _N. serrata(?) are approaching maturity since they exhibit disordered r e t i c u l a t i o n . A purely q u a l i t a t i v e observation of the platforms referred to N. ser- rata(?) i s t h e i r general degenerate appearance, lacking the robustness of the older faunas. There appears to be evidence that the lack of conodonts for a s i g n i f i c a n t part of the section above _N. serrata(?) i s the r e s u l t of a b i o l o g i c c r i s i s . Perhaps the degenerate appearance of these specimens of N.. serrata(?) r e f l e c t 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 the barren i n t e r v a l are equally sparse:' a feature common to any c r i s i s or near e x t i n c t i o n . N. rosenkrantzi marks the reappearance of robust and abundant specimens. . NEOGONDOLELLA n.sp. B PI. 6, fi g s v 1-4. Occurrence: Lower part of the Trold Fiord Formation, Hamilton Peninsula section (F36, F83). Diagnosis: This very symmetrical platform element i s distinquished by i t s very large cusp which i s round i n cross section and directed straight up-wards. Other diagnostic features include a well developed, f a i n t l y t r i -lobed brim posterior to the cusp i n mature elements. This brim bears a coarsely s t r i a t e to f a i n t l y r e t i c u l a t e ornament on the o r a l surface. Fur-thermore, the r e t i c u l a t i o n on the platform margin ends very abruptly at the furrow margin. 93 Description: This new species i s based only, on the posterior regions of the platform owing to fragmentation of the elements. Nevertheless the features present on the posterior h a l f are very d i s t i n c t i v e . In the intermediate form the platform margins are s u b p a r a l l e l (how they taper.in the anterior region i s unknown) and rounded on the posterior end where i t meets the cusp. The cusp i s large, rounded i n cross section, directed almost s t r a i g h t upward, and shows l i t t l e to no l a t e r a l compression. The d e n t i c l e s are a l l low, l a t e r a l l y compressed, and fused at t h e i r bases. The platform margins are r e t i c u l a t e d i n a regular fashion. The r e t i c u l a t i o n does not reach the rounded parts of the posterior of the platform margin. The furrows and d e n t i c l e s are.smooth up to/the point where the r e t i c u l a t i o n begins abruptly. In mature forms a platform margin forms posterior to the cusp. This posterior margin i s f a i n t l y t r i - l o b e d i n o u t l i n e on most specimens but i n others can be very narrow. The cusp i s generally very large, c i r c u l a r i n cross section, and directed s t r a i g h t upwards. The r e t i c u l a t e ornament i s s i m i l a r to that i n the intermediate form except that the r e t i c u l a t i o n reaches the posterior margin and the cusp. The r e t i c u l a t i o n generally gives way to a coarse s t r i a t e d ornament on the brim posterior to the cusp. The d e n t i c l e s are smooth, l a t e r a l l y compressed (some can be node-like), and fused up to half of t h e i r height. The aboral surface i s smooth and bears a low, wide keel which terminates into an equally low, rounded basal loop. The narrow basal groove terminates i n a s l i g h t l y curved, elongated.oval p i t The crimp occupies about 2/3 of the aboral width. Discussion: S u p e r f i c i a l l y , the intermediate form looks s i m i l a r to N_. i d a - hoensis n.subsp. A. However, N. n.sp. B_ has a l a r g e r , l e s s compressed cusp that i s directed upwards unl i k e that for the older species. The r e t i c u l a t e 94 ornament fades towards the furrows ending i n faint, l i n e a r ridges perpendi-cular to the length i n N_. idahoensis n.subsp. A whereas the same ornament ends abruptly and lacks any l i n e a r ridges i n N. n.sp.. B_. Other d i f f e r e n -t i a t i n g features are the p a r t i a l fusion of d e n t i c l e s .at t h e i r base and the lack of r e t i c u l a t i o n on the posterior "shoulders" of the platform. Further-more, the element of N. n.sp. B_„is much more symmetrical than i n older species owing to the p o s i t i o n of the cusp and the p a r a l l e l margin i n the posterior of the platform. These forms look very s i m i l a r to a younger species i d e n t i f i e d by Clark and Behnken (1979) as N. w i l c o x i although the cusp i n N. n.sp. B_ i s generally l a r g e r . Mature forms are distinguished from N. idahoensis n.subsp. A i n terms of the well developed p o s t e r i o r brim surrounding the cusp that i s ornamented with coarse striations". This posterior brim i s present i n younger species (N_. babcocki and N. w i l c o x i i n p a r t i c u l a r ; see Clark and Behnken, 1979, PI. 2, f i g s . 14, 21), however, the ornament i s r e t i c u l a t e d rather than of coarse s t r i a t i o n s . N. n.sp. B_ d i f f e r s from N. b i t t e r i n.subsp. C_ i n being much more symmetrical. The aboral surface i s s i m i l a r to that i n younger and older species. The measurements f o r L2 which range between 240 and 310 um are s i m i l a r to that for N. idahoensis n.subsp. A whereas the measurements for.maximum width 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 serrata postserrata Behnken, 1975 Occurrence: Middle Trold Fiord Formation, Hamilton Peninsula section (F87). Description: This i d e n t i f i c a t i o n i s based on only a very few fragments that are d i s t i n c t l y unlike any others seen through the e n t i r e sequence. 95 The gently arched to almost f l a t platform possesses . su b p a r a l l e l margins and an abruptly squared-off p o s t e r i o r . The de n t i c l e s i n the mature form are large, c i r c u l a r i n cross section and d i s t i n c t . In a small mid-platform fragment, the d e n t i c l e s are l a t e r a l l y compressed and almost e n t i r e -l y fused suggesting t h i s may be a gerontic form. The cusp i s a c t u a l l y a rectangular shaped node projecting to one side of the platform. A small ridge marks the posterior border on the side opposite to the 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 the margins of the p l a t -form reaches the posterior margin and ends abruptly where the furrows l a t -e r a l to the carina begin. Very f a i n t ridges perpendicular to the length can be seen on the otherwise smooth furrows of the mid-platform fragment but '.: there are no serrations on the platform margins. Measurements for Wl (mean = 200 um) and L2 (mean = 260 ym) are very s i m i l a r to those for N. idahoensis n.subsp. A. Discussion; These specimens are ref e r r e d to N_. postserrata(?) because of th 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 to other species i n the same section and because of t h e i r s i m i l a r i t y to some of the square-ended specimens figured by Clark and Behnken (1979, P l . 1, f i g . 17). The shape of the cusp and the squared posterior end are the main d i s -tinguishing features. However, Behnken (1975) indicates that N. postserrata can have both rounded and squared posterior margins, so that the above d i s t i n g u i s h i n g features should not be considered exclusive to the species. NEOGONDOLELLA BITTERI n.subsp. _C P l . 7, f i g s . 1-8. Occurrence: Upper Trold Fiord Formation, Hamilton Peninsula section (F96). Diagnosis: A species characterized by a th i c k platform with a low, wide cusp of c i r c u l a r cross section surrounded by a brim with 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 carina. The subspecies i s characterized by a d i s t i n c t asymmetry r e s u l t i n g from the off-centre p o s i t i o n of the larger posterior lobe. Description: This designation i s based on a number of posterior end f r a g -ments which have very d i s t i n c t i v e features. The p o s t e r o - l a t e r a l margins are p a r a l l e l to s u b - p a r a l l e l while the posterior end i s lobed and d i s t i n c t l y asymmetrical. The posterior margin extends beyond the cusp as a well, developed brim. The posterior asymmetry i s formed by a large lobe occupying one side of the other of the platform centre. The cusp i s positioned near the median but may be to one side or the. other.:as w e l l . The low, wide cusp has a c i r c u l a r cross section. The d e n t i c l e s on the carina are low and rounded, forming nodes that are par-t i a l l y fused at t h e i r bases. Longitudinal furrows adjacent to the carina are r e l a t i v e l y deep, narrow and smooth. The t h i c k and l a t e r a l l y upturned platforms are r e t i c u l a t e d on the margin. The r e t i c u l a t e ornament extends very close to the carina fading into l i n e a r ridges as the ornament approach-es the carina. The r e t i c u l a t i o n extends around the.entire posterior brim as well. Measurements for the width of the platform range between 180 and 220 ym whereas those for L2 range from 260 to 300 ym. The aboral surface has a s t r a i g h t oval shaped p i t surrounded by a roughly t r i a n g u l a r shaped loop which roughly follows the o u t l i n e of the posterior end. The p i t extends a n t e r i o r l y as a narrow groove bordered by a*low, wide keel. The crimp i s smooth and occupies 6/10 of the aboral surface width i n the posterior r e -gion and 7/10 i n the medial, to anterior parts. Discussion: According to Wardlaw and 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 platform that abruptly narrows i n the anterior t h i r d or fourth of i t s length, a low cusp of c i r c u l a r cross section, and low denti-r c l e s on the carina. They d i s t i n g u i s h t h i s species from _N. rosenkrantzi which i s characterized by a wide platform that has a blunt posterior end and that commonly gradually tapers a n t e r i o r l y and by a large cusp of elon-gate-oval cross section. Unfortunately, other authors d i f f e r i n t h e i r determinations for the same material. Clark and Behnken (1971) and Clark et a l . (1979) include forms Wardlaw and Collinson (1979b) r e f e r to N. b i t t e r i within N. rosenkrantzi and N. babcocki. Designation of any species i s a subjective and a r b i t r a r y procedure by the paleontologist. He j u s t i f i e s , t h i s procedure by separating h i s species on comparable morphologic v a r i a b i l i t y exhibited by d i f f e r e n t but r e l a t e d extant species. This i s impossible to accomplish with conodonts. Further-more, with the lack of a f u n c t i o n a l model for conodonts (Bengtson, 1980), i t i s d i f f i c u l t to i n t e r p r e t taxonomic problems by comparison to other b i o t -i c forms. This leaves the i n t e r p r e t a t i o n of importance of various morphol-ogic features, for. d i f f e r e n t i a t i n g species open to subjective and a r b i t r a r y procedure without means of r e s o l u t i o n . However, the success and v a l i d i t y of any morphological model i s determined by the ease with which another student of these conodonts can apply the model to h i s material. Wardlaw arid Collinson'S -diagnoses seem more appropriately to f i t the material from Ellesmere Island. The specimens here r e f e r r e d to N. b i t t e r i are i d e n t i f i e d as such bee.: cause of t h e i r very t h i c k platforms, t h e i r cusp of c i r c u l a r cross section, and t h e i r low. nodiform d e n t i c l e s on the carina. Wardlaw and Collinson(1979b) f i g u r e specimens of N. b i t t e r i from the Gerster Limestone i n Nevadatand. tUtah 98 and from the Retort Phosphatic shale Member of the Phosphoria Formation,. Wyoming. There seem to be minor differences 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 subspecific l e v e l . The specimens from the Retort Member appear to have s l i g h t l y rounder posterior margins and taper more gradually then those from the Gerster Limestone. The Retort Member includes asymmetric forms-owing to the .off-centre p o s i t i o n of the l a r g e r lobe (Wardlaw and C o l l i n s o n (1979b) PI. 1, f i g s . 11, 12). These specimens are "very simiiar:,--indeed 'almost-'identical,.to - those from E l l e s -mere Island. These specimens d i f f e r from N. n.sp. B_ i n terms of t h e i r d i f f e r e n t symmetry, generally smaller cusp, and r e t i c u l a t e microornament on the p l a t -form brim as opposed to s t r i a t i o n s . The measurements for L2 and Wl are s i m i l a r to those f o r N. idahoensis n.subsp. A but the other features make the separation c l e a r . The Ellesmere samples do not exhibit any p o s t e r o - l a t e r a l d e n t i c l e s that can be present i n N. b i t t e r i , but Wardlaw and Collinson (1979b) i n d i -cate that those that do*are. rare v a r i a n t s . NEOGONDOLELLA ROSENKRANTZI n.subsp. D PI. 7, f i g s . 9-12; PI. 8. Occurrence: Upper Trold Fiord Formation, Hamilton Peninsula section (F96). Diagnosis: This species i s characterized by a thick,^wide platform with almost blunt to s l i g h t l y rounded posterior which gradually tapers a n t e r i o r -l y , by a prominent, often modified, cusp of oval cross section surrounded by a well developed brim, and by a narrow but shallow furrow l a t e r a l to:the carina and d i r e c t e d p o s t e r o - l a t e r a l l y towards the. corners of the posterior margin. The subspecies i s based on the enlarged posterdr-lateral platform 99 margins, the gradual taper throughout the e n t i r e length, and by the s l i g h t l y rounded posterior,as opposed to a s t r a i g h t and blunt margin. Description: A. Juvenile - The element i s small, subsymmetrical, s l i g h t l y arched and upturned on i t s margins. The degree of upturning i s greatest i n the anterior half whereas the posterior h a l f i s barely upturned at a l l . The carina has at l e a s t 9 d e n t i c l e s including the cusp which, as.the p l a t -form extends only to the anterior t i p of the cusp forms a free blade pos-t e r i o r l y . The cusp i s high, elongate oval i n cross section, t r i a n g u l a r i n o u t l i n e , l a t e r a l l y compressed, and d i r e c t e d s l i g h t l y p o s t e r i o r l y . The den-t i c l e s a n t e r ior to the cusp are pointed, d i s t i n c t , l a t e r a l l y compressed and increase s l i g h t l y i n height a n t e r i o r l y . The platform tapers gradually towards the anterior and bears a r e t i -c u late ornament r e s t r i c t e d to the margins over the.entire length of the 1 platform. The aboral surface i s smooth and bears a high narrow keel which terminates p o s t e r i o r l y i n a high elongate oval basal loop or flange. B. Intermediate - The element i s subsymmetrical, s l i g h t l y arched, and scarcely upturned on i t s margins. This s l i g h t upturning i s greatest i n the anterior 1/3 and the posterior 1/3 whereas the middle part i s f l a t . The platform margins are s u b p a r a l l e l to gradually tapering over the poster-i o r 2/3 whereas the anterior 1/3 tapers a l i t t l e more r a p i d l y . The platform extends the e n t i r e length of the element arid i s adjacent to, or forms a .. small brim, behind, the cusp. The posterior margin i s rounded i n o u t l i n e . The 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, occurs throughout the e n t i r e length of the element. The aboral surface i s smooth and bears a r e l a t i v e l y high, wide keel which terminates i n a rounded to s l i g h t l y squared-off basal loop. TOO The cusp i s high, oval i n x r o s s section, pointed, and directed s t r a i g h t upwards i f not barely a n t e r i o r l y . The d e n t i c l e s a n t e r i o r to the cusp, of which there are 11 or 12, increase i n height, and compression a n t e r i o r l y and are fused at t h e i r bases. The f i r s t four d e n t i c l e s are barely perceptibly more c l o s e l y spaced, of equal height and s l i g h t l y d i s t i n c t from the others-on the carina. The t i p s of the d e n t i c l e s are f l a t to s l i g h t l y pointed i n the posterior half and pointed i n the anterior h a l f . C. Mature to Gerontic - I t i s at t h i s stage that the many diagnostic fea~. „. tures of the species and subspecies become apparent. Only minor differences exist even between t h i s species and N. idahoensis n.subsp. A at the i n t e r -mediate and e s p e c i a l l y at the j u v e n i l e growth stage. The platform element has a roughly elongated t r i a n g u l a r o u t l i n e and a very robust appearance owing to i t s width (Wl = 200 to 260 ym) and very t h i c k margins. Arching of .:the element i s greater than that observed for the intermediate forms whereas the degree of upturning i s imperceptible. Some specimens a c t u a l l y show a degree of downturning. The posterior margin i s st r a i g h t to s l i g h t l y rounded and sometimes lobed but not with 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 are en^ -larged 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 sub-p a r a l l e l u n t i l they begin to taper more r a p i d l y .in the anterior 1/3. Whereas.an extensive' brim i s formed about the cusp on the posterior of. the p l a t -form i t i s absent from the anterior t i p , where the l a s t 2 to 3 almost t o t a l l y fused d e n t i c l e s form a free blade. The margins of the platform bear a r e t i c u l a t e microornament which i s widest near the mid-length and extends over the e n t i r e length. The i n t e r i o r border of the r e t i c u l a t i o n does not end abruptly but rather fades into f a i n t ridges directed towards 101 the carina. On some specimens t h i s r e t i c u l a t i o n can be found on the d e n t i -c l e s and cusp but even i n these specimens at l e a s t part of the furrow i s •-. smooth. The r e t i c u l a t i o n extends around the brim i n most specimens. In 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 almost s t r i a t e d i n appearance. The smooth furrows are shallow and narrow and p a r a l l e l the carina up to the pos-i t i o n of the cusp where they diverge towards the p o s t e r o - l a t e r a l corners. The cusp i s generally low, elongate oval i n o u t l i n e and positioned at about the median of the platform and directed s t r a i g h t upwards. In some specimens the cusp i s directed either dextral or s i n i s t r a l to the midline whereas i n a few of- the specimens i t i s fused with a couple of adjacent d e n t i c l e s forming an elongate ridge directed towards a p o s t e r o - l a t e r a l corner. The majority 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 10) are flat-topped, l a t -e r a l l y compressed, fused to about 1/2 to 2/3 of t h e i r height and of equal height. From t h i s point, which roughly coincides with the more r a p i d l y tap-ering of the platform, the d e n t i c l e s f i r s t increase i n height and then de-crease again. The smallest anterior d e n t i c l e s are almost t o t a l l y fused, forming a c h a r a c t e r i s t i c blade which i s i n part free of the platform. In other specimens the d e n t i c l e s are more node-like. Rare specimens have the f i r s t four d e n t i c l e s anterior to the cusp smaller and more c l o s e l y spaced than the others. The aboral surface i s smooth and bears a low, wide keel which termin-ates i n a higher and roughly t r i a n g u l a r basal loop. The shape of t h i s ba-s a l loop i s quite v a r i a b l e . The narrow basal groove terminates p o s t e r i o r l y i n a s t r a i g h t elongate oval p i t . The keel increases,.;in height and narrows i n a b l a d e - l i k e ridge which connects with the o r a l free blade at the anter-i o r t i p . The crimp covers roughly 6/10 of the width i n the posterior parts of the platform. 102 Discussion: The early growth stages are very s i m i l a r even to N_. idahoensis n.subsp. A. Szaniawski and Malkowski (1979) pointed out the s i m i l a r i t y of early 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 early growth stages throughout the phylogeny of Neogondolella points to the close r e l a t i o n s h i p of a l l the species. Some mature forms of N. i d a - hoensis n.subsp. A a c t u a l l y mimic N_. rosenkrantzi. Mature specimens of N. rosenkrantzi n.subsp. D_differ i n terms of a more tr 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 arrangement of the d e n t i c l e s including the anterior blade (a feature not seen i n N.. idahoensis) , shallower furrows, and thicker platform margins. However, for r e l i a b l e i d e n t i f i c a t i o n t h i s mimicry points to the need, not only for mature forms, but also f o r enough representatives to include 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 these species. As pointed out i n the discussion for N. b i t t e r i n.subsp. C. the main features d i s t i n g u i s h i n g N.. rosenkrantzi. from N.. b i t t e r i are the s l i g h t l y -straight to almost blunt posterior ends, and a large cusp of elongate-oval cross section. As well the enlarged p o s t e r o - l a t e r a l margins and d i f f e r e n t symmetry d i s t i n g u i s h N. rosenkrantzi n.subsp. I) from N. b i t t e r i n.subsp. C^. Wardlaw and Collinson (1979b) figure.specimens of N. rosenkrantzi from the Retort Phosphatic Shale Member of the Phosphoria Formation and from the Gerster Limestone.. The l a t t e r have a very blunt posterior end and ap-pear more "advanced" than the Ellesmere specimens, which more c l o s e l y r e -semble the Retort Member specimens i n posterior o u t l i n e and manner and de-gree of tapering. There seem to be grounds for suggesting that these d i f -ferences are s i g n i f i c a n t at the i n f r a s p e c i f i c l e v e l (subspecies). To c a l l a t t e n t i o n to some of these d i f f e r e n c e s , therefore, I have re f e r r e d these 103 specimens to a d i s t i n c t subspecies. The specimens d i f f e r from other species i n the Trold Fiord Formation i n t h e i r d e n t i c l e shape and configuration, the shape and p o s i t i o n of the cusp, the greater thickness of the platform margins and the p o s i t i o n and configuration "of r e t i c u l a t e ornament. Ramif orm.. Elements NEOGONDOLELLA IDAHOENSIS n.subsp. A - XANIOGNATHUS TORTILIS (Tatge) PI. i, f i g s . 1-4. Ozarkodina t o r t i l i s Tatge, 1956 Occurrence: Lower Assistance Formation, Hamilton Peninsula section. Description: This blade-shaped element has a long anterior process with as many as 8 sharp, pointed, subequal and l a t e r a l l y compressed d e n t i c l e s , a l l i n c l i n e d p o s t e r i o r l y . The cusp i s high, sharp and l a t e r a l l y compressed. At l e a s t three d e n t i c l e s are present on the short posterior process, which i s twisted to one side or the other. The undersurface of both processes i s grooved and terminates i n a pronounced basal p i t d i r e c t l y below the main cusp. The de n t i c l e s a l l bear a very f i n e s t r i a t e microornament.. Discussion: This species i s distinguished from the older X. abstractus by the l e s s robust blade and the twisted (as opposed to straight) posterior process. According to Behnken (1975) the range of X. t o r t i l i s begins at about the end of that of N. serrata . This would indi c a t e a Late Roadian age but t h i s species occurs withNeogondolella platforms of Early Roadian age. These specimens may be intermediate between X. t o r t i l i s and the older X. abstractus as the base of the den t i c l e s appear more robust than specimens of X. t o r t i l i s figured by Behnken. (1975) ; nevertheless they--do Appear closer to X. t o r t i l i s in most respects. 104 NEOGONDOLELLA.IDAHOENSIS n.subsp. A - ELLISONIA EXCAVATA Behnken, 1975 PI. 1, f i g . 15. Occurrence: Lower Assistance Formation, Hamilton Peninsula section. Description: The specimens have a v a r i a b l e hindeodelliform morphology with , beneath the- cusp,- a small conical'basa'l- p i t which i s l a t e r a l l y compressed and i n c l i n e d p o s t e r i o r l y . The posterior bar i s long and bears at l e a s t 10 to 12 d i s c r e t e pointed d e n t i c l e s that are of v a r i a b l e s i z e . along the bar. 3 d e n t i c l e s are present on the downward projecting short anterior bar. Discussion: The. figured .spec imehif. probably: represents the LB.: element of a multielement species which includes U, LA, and LF elements which were not observed. Behnken (1975) indicates that t h i s species occurs with N. idaho- ensis i n West Texas., and would thus have a Leonardian age. NEOGONDOLELLA IDAHOENSIS n.subsp. A - ELLISONIA TRIBULOSA (Clark and Ething-ton, 1962) PI. 2, f i g s . 5-8 Apatognathus t r i b u l o s u s Clark and Ethington, 1962 Occurrence: Lower Assistance Formation, Hamilton Peninsula Description: Of the U, LAI, LA2 and LC elements LAI ( f i g s . 6, 7) and LC ( f i g s . 5, 8) elements are present and described. LAI - This i s a s l i g h t l y asymmetrical lohchodiniform element with a high, l a t e r a l l y compressed and p o s t e r i o r l y i n c l i n e d cusp. The posterior bar bears 3 to 4 d e n t i c l e s whereas the short anterior bar bears 2 pointed but larger d e n t i c l e s . The basal p i t below the cusp i s formed by very s l i g h t expansion of a groove which i s present over the e n t i r e length of the aboral surface. LC - This i s an enantiognathiform element with a long posterior 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 su b p a r a l l e l f o r much of t h e i r length. The main cusp i s t r i a n -gular i n cross section and.the corners are extended into sharp ridges. The short downward projecting anterior bar bears one to two de n t i c l e s and pro-j e c t s at an angle of about 60 degrees to the posterior bar. Below the cusp i s a large t r i a n g u l a r basal p i t . Discussion: This species occurs through the range of N. serrata and N. postserrata i n d i c a t i n g a Roadian to Early Wordian age. Perhaps the pre-sence of both E. excavata and E..' / t f i b u l o s a together indicates overlap of t h e i r ranges i n the Early Roadian which f i t s i n well with the Early Roadian age assigned to N. idahoensis n.subsp. A because of i t s intermediate p o s i - v ' . t i o n between N. idahoensis and N. serrata. NEOGONDOLELLA IDAHOENSIS n.subsp. A - PRIONIODELLA DEGRESCENS Tatge, 1956 P l . 1, f i g . 14. Occurrence: Lower Assistance Formation, Hamilton Peninsula section. Description: This- specimen' Is a short, s t r a i g h t and denticulate "element without a main cusp. The s i x d i s c r e t e and pointed d e n t i c l e s are subequal i n height. Discussion: Behnken (1975) includes t h i s species within the range of N. postserrata, i n other words, Early Wordian. However, s i m i l a r forms have much greater ranges. Other elements which may be assigned to P r i o n i o d e l l a species are fragmental and u n i d e n t i f i e d . Addendum: A paper, i n preparation by the author describes Neogondolella  idahoensis n.subsp. A, N_. nsp. By N. b i t t e r i n.subsp. C and N. rosenkrantzi n.subsp. D a s N. idahoensis praeserrata n.subsp., N.. p e r r y i n.sp., N. b i t t e r i a r c t i c a n.subsp. and N_. rosenkrantzi ellesmerensis n.subsp., r e s p e c t i v e l y . 106 REFERENCES Bamber, E.W. and Waterhouse, J.B. 1971: Carboniferous and Permian stratigraphy.and paleontology, northern Yukon T e r r i t o r y , Canada; B u l l , of Can. 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Waterhouse, J.B., Waddington, J . and Archbold, N. 1978: The evolution of the Middle Carboniferous to Late Permian brachiopod subfamily S p i r i f e r e l l i n a e , Waterhouse; In Western and A r c t i c Canadian Biostratigraphy, CR. Stelck and B.D.E. Chatterton (Eds.), Geol. Assoc. of Can. Spec. Paper, 18, pp. 415-443. Weimer, R.J. 1970: Rates of d e l t a i c sedimentation and in t r a b a s i n deformation, Upper Cretaceous of Rocky Mountain Region; In De l t a i c Sed-imentation: Modern and Ancient, J.Pi Morgan (Ed.), Society of Economic Paleontologists and Mineralogists Spec. Publ. #15, pp. 270-292. Wilson, L.R, 1962: Plant M i c r o f o s s i l s , Flowerpot Formation;'. Oklahoma Geol. Surv. C i r c u l a r , 49, 50p. Wright, S. 1967: Comments on.the preliminary working papers, df Eden and Wadding-ton; In Mathematical. Challenges to the Neo-Darwinian Theory of Evolution, P.S. Moorehead. and M.M'. Kaplan (Eds.), The Wistar Inst. Symposia Monograph #5, Philadelphia, pp. 117-120. Youngquist, W., Hawley, R.W. and M i l l e r , A.K. 1951: Phosphoria conodonts from S.E. Idaho; J. Paleontol., 25, pp. 356-364. Zie g l e r , W. (Ed.) 1973: Catalogue of Conodonts; -E". Schweizerbart' sche Verlagsbuchhand-lung, 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 El e c t r o n Micrographs. Fig s . 1, 2 Neostreptognathodus p r a y i Page 73 1. Oblique l a t e r a l to o r a l view. Note the r e c r y s t a l l i z e d texture. l o c . T l O O . (X250). 2. Oblique l a t e r a l to o r a l view of platform posterior.F100. (X125). Figs. 3-6 Anchignathodiis minutus Page 73 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 to cusp. F49*. (X100) . 4. Oral view.F49.(XI00). 5. L a t e r a l view. Note lac k of d e n t i t i o n a n t e r i o r to cusp.F49.(X100). 6. Close-up of d e n t i t i o n on F i g . 5. Note the smooth surface f r e e of ornament.F49. (X400). Figs. 7-13 Neogondolella idahoensis subsp. indet. Page 74 7. Oral view of intermediate to mature form.F100. (X125). 8. L a t e r a l view of a n t e r i o r part of element showing d i s c r e t e d e n t i -c l e s . F100. (X125). 9. L a t e r a l view of po s t e r i o r of intermediate form.F100.(X125). 10. Oral view of intermediate form.FlOO. (X125). 11. Oral view of mature form.F100. (X250). 12. Close-up of r e c r y s t a l l i z e d (apatite) texture.F100.(X500). 13. Aboral view of po s t e r i o r of intermediate form.F100.(X125). F i g . 14 Neogondolella idahoensis n.subsp. A - P r i o n i o d e l l a decrescens 14. L a t e r a l view.F49. (X100). Page 105 F i g . 15 Neogondolella idahoensis n.subsp. A - E l l i s o n i a excavata 15. L a t e r a l view..F49. (X100) . P a 8 e 1 0 4 115 Explanation f or Pl a t e 2 A l l f i g u r e s are Scanning E l e c t r o n Micrographs. Figs. 1-4 Neogondolella idahoensis n.subsp. A - Xaniognathus t o r t i l i s 1. Close-up of d e n t i c l e . Note the f i n e s t r i a t e d surface texture.F49. (X400). P a ^ 1 0 3 2. Close-up of denticle.F49. (X400). 3. L a t e r a l view. F49. (X100). 4. L a t e r a l view. F49. (X100) . Fig s . 5-8 Neogondolella idahoensis n.subsp. A - E l l i s o n i a t r i b u l o s a 5. LC element. Note ridge or keel on main cusp.F49. (X100). Page 104 6. LAI element.F49. (X100). 7. LAI element.F54. (X150). 8. LC element.F49. (XI00). Fi g s . 9-19 Neogondolella idahoensis n.subsp. A Page 75 9. Oral view of var. g r a c i l i s . Main cusp and p o s t e r i o r end pointing to the bottom of the page.F49. (X100). P a g e 8 4 10. Oral view of var. intermediatus.F49. (X100). Page 86 11. Oral view of var. intermediatus.F54.(X90). 12. Oral view of var. g r a c i l i s . Note the d i s t i n c t four d e n t i c l e s a n t e r i o r to the cusp.F49.(X100). 13. Oral view of var. intermediatus.F49.(X75). 14. Oral view of var. robustus. Note the serrations on a n t e r i o r 1/3. F49.(X100). P a g e 8 5 15. L a t e r a l view of p o s t e r i o r showing large cusp and lack of brim p o s t e r i o r to cusp: intermediate to mature.F49.(X100). 16. L a t e r a l view of j u v e n i l e . Note the lack of platform on l a t e r a l l y compressed, t r i a n g u l a r o u t l i n e d cusp.F49. (X150). 17. Oblique l a t e r a l view of juvenile.F49. (X150). 18. Oblique l a t e r a l view of intermediate.F49. (X100). 19. Oblique l a t e r a l view of j u v e n i l e . Note the gradual increase i n d e n t i c l e height a n t e r i o r l y and the upturning on the platform margin.F49.(XI00). 117 Explanation f o r Pl a t e 3 A l l f i gures are Scanning Electron Micrographs. Figs. 1-17 Neogondolella idahoensis n.subsp. A Page 75 1. Oral view var. intermediatus. Note 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. Oral view var. intermediatus. F49. (X10Q). Page 86 3. Oral view var. gracilis.F49.(X75). Page 84 4. Oral view var. gracilis.F49-(X75). 5. Oral view var. gracilis.F49.(XI00). 6. Oral view var. gracilis.F49.(X100). 7. Oral view var. robustus.F49.(XI00). Page 85 8. Oral view showing high, 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. Oral view var. robustus.F49.(X100). 10. Aboral view.F54.(X125). 11. Aboral 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 margin from that i n F i g . 10.F54.(X125). 12. Oral view of t h i c k platform margins of mature form.F54.(X85). 13. L a t e r a l view of intermediate form.F49.(X100). 14. L a t e r a l view of intermediate to mature form.F49.(X80). 15. Lateral, view of mature form. Note that the platform reaches the pos t e r i o r of the cusp (unlike that i n Figs. 13, 14). F54.(X80). 16. L a t e r a l view of var. gracilis.F49.(XlOO). 17. L a t e r a l view of var. gracilis.F49.(XlOO). 119 Explanation for P l a t e 4 A l l f i g u r e s are Scanning E l e c t r o n Micrographs. Figs. 1-10 Neogondolella idahoensis n.subsp. A Page 75 1. Oral view of mature to gerontic form of var. lobatus that mimics N. rosenkrantzi i n shape. 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). 2. Oral view of var. intermediatus•F49.(X100). Page 86 3. Oral view of var. intermediatus with a n t e r i o r s e r r a t i o n s . Note the lack of r e t i c u l a t e d ornament on the anterior-most margins of platform. F54.(X75). 4. Close-up of a n t e r i o r part of F i g . 3. F54.(X150). 5. Oral view of mature to gerontic form.F54. (X125). 6. Oral view of mature to gerontic form showing p a r t i a l f u sion of p o s t e r i o r denticles.F54.(X150). 7. Oral view of var. constrictus.F49.(X100). Page 86 8. Oral view of var. lobatus. Note the t h i c k platform margin and d i s c r e t e , very l a t e r a l l y compressed denticles.F54. (X150). Page 87 9. Oral view of intermediate to mature var. intermediatus. Note 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 of the platform.F54.(X150). 10. Oral view of gerontic form with complete fusion of p o s t e r i o r d e n t i c l e s . Note the gradual and progressive fusion of p o s t e r i o r d e n t i c l e s displayed by F i g s . 5, 6 and 10. This demonstrates the synonymy of N. phos-phoriensis with N. idahoensis.F54.(X180). Plate 4 120 121 Explanation f or Plate 5 A l l f i g u r e s are Scanning E l e c t r o n Micrographs. Figs. 1-10 Neogondolella idahoensis n.subsp. A Page 75 1. Close-up of r e t i c u l a t e d ornament. Note the d i s t i n c t but rounded edges of ridges on the outside margin ( l e f t ) , the d i s t i n c t and sharp ridges on the mid-part of the platform margin, and fading and elon-gate ridges near the furrow ( r i g h t ) . F49.(X1500). 2. Close-up of P l . 2, F i g . 14 showing the presence of r e t i c u l a t e d ornament on the d e n t i c l e tip.F49.(X400). 3. Close-up of P l . 2, F i g . 14 showing the presence of r e t i c u l a t e d ornament on the cusp and fused p o s t e r i o r d e n t i c l e s but absence on the furrows.F49.(X400). 4. Aboral view of j u v e n i l e form showing the high, narrow keel and elongate-oval loop. F49.(XI00). 5. Aboral view.F49.(X100). 6. Aboral view of intermediate form.F49. (X100). 7. Aboral view of intermediate to mature form showing low, wide keel and s l i g h t l y t r i a n g u l a r loop.F49.(X100). 8. Aboral view of mature form showing very low and wide keel and t r i a n g u l a r loop. Note the progressive changes of aboral features from Fi g . 5 ( j u v e n i l e ) to F i g . 8 (mature). F49.(XI00). 9. Aboral view of posterior end of mature form.F49.(X150). 10. Close-up of loop i n F i g . 9. Note the truncation of growth lamellae at p o s t e r i o r of loop ( e s p e c i a l l y evident on r i g h t hand side) which accom-panies the t r a n s i t i o n from elongate oval to t r i a n g u l a r shape of loop.F49. (X600). Plate 5 122 123 Explanation f or Plate 6 A l l f i g u r e s are Scanning El e c t r o n Micrographs. Figs. 1-4 Neogondolella n.sp. B Page 92 1. Oral view of intermediate form. Note the very symmetric shape. F83.(X250). 2. Oral view of mature form showing large 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 with coarse s t r i a t e ornament on brim.F83.(X300). 3. Oral view of symmetric mature form.F36.(X250). 4. Oral view of gerontic form with large, c i r c u l a r ( i n cross section) cusp with narrow p o s t e r i o r platform margins and fused d e n t i c l e s . F36.(X250). Fig s . 5, 6 Neogondolella postserrata(?) Page 94 5. Oral view.F87.(X125). 6. Close-up of F i g . 5 showing blunt p o s t e r i o r margin and rectangular nodiform cusp.F87. (X250). Figs. 7-9 Neogondolella serrata(?) Page 90 7. Oral view of intermediate to mature form. Note the degenerate o v e r a l l appearance.F73. (X300). 8. Close-up of p o s t e r i o r end of F i g . 7. F73.(X600). 9. O b l i q u e - l a t e r a l to o r a l view of intermediate form. Note the sharpness of a l l the denticles.F63.(X300). 125 Explanation f o r Pl a t e 7 A l l f i g u r e s are Scanning Electron Micrographs. Figs. 1-8 Neogondolella b i t t e r i n.subsp. C Page 95 1. Aboral view of mid-platform fragment.F47. (X125). 2. Aboral view of po s t e r i o r end.F47. (X125). 3. Oral view of mature form with asymmetric p o s t e r i o r end and large brim.F47. (X250). 4. Oral view of mature form with asymmetric p o s t e r i o r end and large brim.F47.(X250). 5. Oral view. Note the large, c i r c u l a r cusp s i m i l a r to N. n.sp. IS but also the d i s t i n c t asymmetry. F47. (X250). 6. Oral view.F96.(XlOO). 7. Oral view showing p a r t i a l fusion of cusp and posterior-most d e n t i c l e and r e t i c u l a t e ornament on the carina. F96.(XlOO). 8. Oral view of. mature form showing a s l i g h t l y rounded and asymmetric pos t e r i o r end that i s s i m i l a r to the more blunt ended N. rosenkrantzi.F96.(X150) Figs. 9-12 Neogondolella rosenkrantzi n.subsp. D Page 98 9. Oral view of mature form showing the s t r a i g h t , l e s s d i s t i n c t l y asymmetric p o s t e r i o r end as compared to _N. b i t t e r i . Note the r e t i c u l a t e d ornament on the carina.F96.(XlOO). 10. Oral view of mature form w i t h a very t h i c k platform, elongate-oval 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 with furrows and c a r i n a that are almost e n t i r e l y covered with r e t i c u l a t e microornament.F96.(XlOO). 11. Aboral view of platform showing a wide t r i a n g u l a r , and asymmetric loop.F96.(X85). 12. Oral view of mature to gerontic form showing the p o s t e r o - l a t e r a l l y d i r e c t e d furrows and swollen p o s t e r i o r platform margins.F96.(X140) . Plate 7 126 127 Explanation f o r Plate 8 A l l f i g u r e s are Scanning Electron Micrographs. Figs. 1-13 Neogondolella rosenkrantzi n.subsp. _D Page 98 1. Oral view of intermediate form that looks very s i m i l a r to N. i d a -hoensis^ F96.(X85). 2. L a t e r a l view of p o s t e r i o r end of a mature to gerontic form showing large brim, fused carina and downturned l a t e r a l margins.F96.(X100) . 3. Oral view of mature form showing the p o s t e r o - l a t e r a l l y d i r e c t e d furrows and a s l i g h t twist of the platform at the anterior end.F96-(X85). 4. Oral view of mature form with narrow brim and wide but shallow furrows.F96.(X100). 5. L a t e r a l view of intermediate form.F96.(X85). 6. L a t e r a l view of a n t e r i o r end showing the k e e l - l i k e carina (owing to f u s i o n of d e n t i c l e s ) and lac k of platform at anterior-most end: both features are c h a r a c t e r i s t i c for mature to gerontic i n d i v i d u a l s of t h i s subspecies.F96.(X85). 7. L a t e r a l view of intermediate form.F96. (.X85) . 8. L a t e r a l view of F i g . 3 showing k e e l - l i k e a n t e r i o r carina.F96.(X75). 9. Oblique l a t e r a l view of j u v e n i l e form. Note the s i m i l a r i t y to j u v e n i l e s of N. idahoensis n. subsp. A ( P l . 2, F i g . 16).F96.(X225). 10. Oral view of gerontic i n d i v i d u a l with cusp and posterior-most d e n t i c l e ( s ) fused and dire c t e d p o s t e r o - l a t e r a l l y . F 9 6 -(X85). 11. Close-up of ordered r e t i c u l a t e d mieroprnament and flat-topped, smooth denticle.F96.(X900). 12. Close-up of gerontic o r a l surface showing the almost.complete lack of furrows.' Reticulated microornament covers almost the e n t i r e o r a l surface. Note the elongate form of r e t i c u l a t e d ornament where the furrows are normally positioned.F96.(X200) . 13. Aboral surface of intermediate form.F96.(X100). APPENDIX I Schematic of morphological terminology and measured parameters for Neogondolella and measurements (Lj_, L 2, Hi, Wi, W2: a l l i n ym) of specimens from F48 - F54.. aboral view Posterior Parameter Measurements for Sample F48. ecimen L l L2 H V*l *1 W2 1Q4 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 for Sample F49 ecimen L l L2 H V W 1 W l W 2 . h\ (w1+w io 4 1 800 24 Q 140. 5.71 11 72.73 4.Q0 20Q 150 4.20 2 1400 380 280 5.00 13 107.69 4.83 29.0 280 : 10.83 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 47.40./ 3.29. 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 27: 980 350 23Q 4.26 11 89.. 09. 4.45 220 200:; 7.35 28 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: 3.64 131 Parameter Measurements for Sample F49 (cont.) lecimen L l L2 H VW1 W l W 2 % L 1 (Wj+V 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 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 60 1120 320 180. 6.22 14 80.00' 5.09 220 19.0 6.56 132 Parameter Measurements f o r Sample F52 Specimen L l L2 H V * l W l F 2 J2L1(W1+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. oa 4.50 260 240 8.25 133 Paramet er 2 C i m e n L l L2 H. y H 1 760 220 160 4.75 11 2 1180 340 250 4.72 13 3 1160 320 220 5 . 2 7 . 15 4 . 1270 300 210 6.05 16 5 1310 320 190 6.89 14 6 1120 320. 180 6.22 13 7 980 340 190 5.16 11 8 580 280 130 4.46 9 9 720 320 180 4.00 10 10 700 300 160 4.38 9 11 980 280 140 7.00 13 12 1500 380 17Q 8.82 16 13 960 29Q 180 5.33 11 14 860 320 160 5.38 10 15 640 260 130 4.92 9 16 1100 400 170 6.47 11 17 920 350 180 5.11 10 18 620 280 140 4.43 9 19 670 280 160 4.19 9 20 740 240 150 4.93 10 21 700 240 130 5.38 10 22 500 260 120 4.17 8 23 800 300 200 4.0Q 10 24 12Q0 40Q 190 6.32 12 25 840 29.0. 160 5.25 11 26 700 280 130. 5.38 10 27 660 280 120 ., 5.50. 10. 28 980 360 180 5.44 11 29 580 240 120 4.83 9. 30 760 340 150. 5.07 9 .ts for Sample F53 69.09 4 .22 180 150- 3.63 9Q.77 4 .07 290 280 9.69 77.33 4 .46 260 220 7.69 79.38 5 .29 240 200 . 6.60 93.57 5 .04 260 260 8.32 86.15 4 .31 260 240 8.00 89.09 4 .45 220 180 6.80 64.44 3 .63 160 130 4.06 72.00 4 .00 180 160 5.44 77.78 3 .68 190 180 5.55 75.38 4 .45 220 180 5.60 93.75 3 .95 380 310 13.11 87.27 4 .00 240 200 6.38 86.00 4 .30 200 160 5.76 71.11 3 .56 180 140 4.16 100.00 4 .23 260 200 9.20 92.00 4 .18 220 180 7.00 68.89 3 .65 170 140 4.34 74.44 4 .47 150 130 3.92 74.00 4 .63 160 140 3.60 70.00 4 .38 160 140 3.60 62.50 3 .57 140 120 3.38 80.00 3 .64 220 200 6.30 100.00 4 .14 290 280 11.40 76.36 4 .67 180 140 4.64 70.00. 4 .12 17Q 120 4.06 66.00. 3 .67 180 160 4.76 89. Q9 4 .45 220 180 7.20 '.. 64.. 44 4 .14 140 120 3.12 84.44 4 .75 160 120 4 .76 134 Parameter Measurements for Sample F53 (cont.) Specimen L L 2 H I^/II // 1^ /// L /W \ W2 %L O^+wp i o 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 3 6 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 4.69 15 100.00 4.41 340 320 14.52 135 Parameter Heasurements for Sample F54 scimen • L l . # y # Specimen L l // 1 600 9 66.67 201 580 9 64.44 2 : 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 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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