<I>Ewingia</I> Case & Cappetta , 1997 (Chondrichthyes: Rajiformes), preoccupied by <I>Ewingia</I> pearse , 1929 (Insecta: Arachnida)

HEPTRANCHIAS HOWELLII (REED, 1946) (SELACHII: HEXANCHIDAE) IN THE EOCENE OF THE UNITED STATES AND BRITISH COLUMBIA BY BRUCE J . WELTON1

ABSTRACT N o t i d a n i o n h o w o l l i i R e e d , 1946 ( S e l a c h i i : Hexanchidae) from t h e E o c e n e S h a r k R i v e r F o r m a t i o n of New J e r s e y i s r e f e r r e d t o the genus H e p t r a n c h i a s . T e e t h o f H e p t r a n c h i a s from t h e E o c e n e of Vancouver I s l a n d , B r i t i s h Columbia, t h e L i n c o l n Creek Formation of W a s h i n g t o n , and t h e K e a s e y and N e s t u c c a F o r m a t i o n s of Oregon a r e c o n s i d e r e d t o be c o n s p e c i f i c w i t h H e p t r a n c h i a s h o w e l l i i . H. h o w e l l i i i s c h r o n o l o g i c a l l y r e s t r i c t e d t o s t r a t a of N a r i z i a n and l o w e r R e f u g i a n a g e i n t h e n o r t h e a s t e r n P a c i f i c and i s i n t e r p r e t e d t o be a d e e p w a t e r f o r m , o c c u r r i n g i n a s s o c i a t i o n w i t h u p p e r bathyal-lower n e r i t i c invertebrate faunas. INTRODUCTION G i l l (1885) defined t h e family Hexanchidae t o i n c l u d e t h r e e extant g e n e r a , Hexanchus R a f i n e s q u e , 1810, Notorhynchus A y r e s , 1 8 5 5 , and H e p t r a n c h i a s R a f i n e s q u e , 1 8 1 0 . H e x a n c h u s and N o t o r h y n c h u s had a c o s m o p o l i t a n d i s t r i b u t i o n t h r o u g h o u t t h e C e n o z o i c w h e r e a s H e p t r a n c h i a s i s p r e s e n t l y known o n l y f r o m t h e O l i g o c e n e of J a p a n ( A p p l e g a t e and U y e n o , 1968) and t h e E o c e n e of V a n c o u v e r I s l a n d , B r i t i s h C o l u m b i a (Waldman, 1 9 7 1 ) . Reed ( 1 9 4 6 ) d e s c r i b e d N o t i d a n i o n h o w e 1 1 i i , a new E o c e n e s h a r k o f t h e f a m i l y N o t i d a n i d a e from t h e S h a r k R i v e r F o r m a t i o n of New Jersey. She s t a t e d t h a t " I n s i z e , N o t i d a n i o n h o w e l l i i c o m p a r e s f a v o r a b l y w i t h t h e C r e t a c e o u s and E a r l y T e r t i a r y s p e c i e s of Not i d a n i o n microdon ( A g a s s i z ) , Notidanion s e r r a t i s s i m u s ( A g a s s i z ) , and N o t i d a n i o n t e n u i d e n s ( L e r i c h e ) , a l l of w h i c h w e r e o r i g i n a l l y d e s c r i b e d a s s p e c i e s of N o t i d a n u s . I t d i f f e r s from N o t i d a n i o n m i c r o d o n ( A g a s s i z ) and N o t i d a n i o n s e r r a t i s s i m u s ( A g a s s i z ) i n h a v i n g t h e a n t e r i o r d e n t i c l e s much l a r g e r , and i n h a v i n g a d i f f e r e n t number and a r r a n g e m e n t of a c c e s s o r y c o n e s . I t d i f f e r s from N o t i d a n i o n t e n u i d e n s ( L e r i c h e ) i n t h e g r e a t e r t h i c k n e s s of t h e r o o t , more n u m e r o u s a n t e r i o r d e n t i c l e s and f e w e r a c c e s s o r y c o n e s " ( R e e d , 1946: 3 ) . There i s a g r e a t s i m i l a r i t y e l l i i and t e e t h of H e p t r a n c h i a s t h e Eocene of Vancouver I s l a n d , n o t compare h i s specimens w i t h

Museum of P a l e o n t o l o g y , 94720.

b e t w e e n R e e d ' s f i g u r e s of N. howw h i c h Waldman ( 1 9 7 1 ) f i g u r e s from B r i t i s h Columbia. Waldman d i d N. h o w e l l i i ; h o w e v e r , s u b s e q u e n t

U n i v e r s i t y of C a l i f o r n i a ,

1.

Berkeley,

examination of h i s specimens revealed t h a t t h e s e two forms are congeneric and probably c o n s p e c i f i c . Welton (1972, 1973) d e s c r i b e d Heptranchias from the l a t e Eocene of Oregon but r e f r a i n e d from making a s p e c i f i c d i a g n o s i s . Morphologically the Oregon t e e t h d i f f e r only s l i g h t l y from N. h o w e l l i i and I now c o n s i d e r them t o be c o n s p e c i f i c . Heptranchias and Hexanchus were defined by Rafinesque in 1810 for sharks having seven and s i x p a i r s of g i l l openings, r e s p e c t i v e l y . Cuvier (1817, 1829) placed both genera" in synonomy w i t h Notidanus Cuvier, 1817 (= Notidanion Reed, 1946) in the family N o t i d a n i d a e . The name Hexanchus has p r i o r i t y over Notidanus, a problem d i s c u s s e d by Applegate (1965b). He demonstrated t h a t each of t h e s e genera may be d i s t i n g u i s h e d from the o t h e r s on t h e b a s i s of t o o t h morphology, the lower l a t e r a l s in p a r t i c u l a r . Hexanchus may have e i t h e r a smooth (as in the case of j u v e n i l e t e e t h ) or s e r r a t e d symphysial margin on the primary cusp ( F i g . I E ) . Heptranchias has one t o f i v e d i s t i n c t narrow d e n t i c l e s on the symphysial margin of the primary cusp (Fig. 1C), and the t h i r d genus of Hexanchidae, Notorhynchus, d i f f e r s in having small unequal d e n t i c l e s ( F i g . I D ) . The lower l a t e r a l t e e t h in Hexanchus and Notorhynchus e x h i b i t a l a r g e primary cusp and s m a l l e r secondary cusps which d e c r e a s e in h e i g h t along a g r a d i e n t from s c ^ , SC2, sc? . . . t o the l a s t cusp ( F i g . I D , E ) . Heptranchias ( F i g . 1 A-C and PI. 1) resemb l e s o t h e r hexanchids in having a l a r g e primary cusp and d i f f e r s in i t s ungraduated secondary cusp p a t t e r n . The meagerness of the f o s s i l record of Heptranchias i s more a p p a r e n t than r e a l and i s due in p a r t t o s y s t e m a t i c and taxonomic confusion r e g a r d i n g g e n e r i c d i f f e r e n c e s in the Family Hexanchidae. For example, in a paper d e s c r i b i n g t h e elasmobranch faunas of A u s t r a l i a , Pledge (1967: 140) s t a t e d t h a t "Notidanus Cuvier i n cluded Hexanchus and Heptranchias Rafinesque 1810, and Notorhynchus Ayres 1855, a s t h e r e i s no a p p a r e n t g e n e r i c d i f f e r e n c e between the t e e t h of t h e s e l i v i n g g e n e r a . " Of the four figured hexanchid t e e t h ( P l e d g e , 1967: p i . 1, f i g s . 1-3) which he r e f e r s t o Notidanus s e r r a t i s s i m u s A g a s s i z , the lower t o o t h in f i g . 3 belongs t o Heptranchias and the remainder a r e c l e a r l y lower l a t e r a l s of Hexanchus• Jordan (1907) d e s c r i b e d Heptranchias a n d e r s o n i on the b a s i s of four t e e t h from B a r k e r ' s Ranch (Hemingfordian), Kern County, C a l i f o r n i a . L e r i c h e (1908) synonymized Heptranchias andersoni with Notidanion primigenius and Arambourg (1927) r e f e r r e d Heptranchias a n d e r s o n i t o Hexanchus g i g a s . I c o n s i d e r Arambourg's g e n e r i c assignment t o be c o r r e c t but h e s i t a t e t o r e f e r the West Coast s p e c i e s t o a European taxon (Hexanchus g i g a s ) u n t i l the former has been a d e q u a t e l y diagnosed.

2.

lingual

cr symphysial rt A.

commissural B.

F i g u r e 1. A & B, Heptranchias h o w e l l i i (Reed, 1946); h o l o t y p e , ANSP 15078, lower l e f t l a t e r a l t o o t h . C. Heptranchias p e r l o ; LACM F - 4 6 8 , lower l e f t fourth l a t e r a l t o o t h , s c a l e l i n e = 2 mm. D. Notorhynchus m a c u l a t u s ; LACM F - 4 6 1 , lower l e f t f o u r t h l a t e r a l t o o t h , s c a l e l i n e = 4 mm. E. Hexanchus g r i s e u s ; LACM F-107, lower l e f t f o u r t h l a t e r a l t o o t h , s c a l e l i n e = 4 mm. Abbreviat i o n s : cf, crown f o o t ; c r , crown; dn, d e n t i c l e s ; l l r , l o n g i t u d i n a l l i n g u a l r i d g e ; p c , primary c u s p ; r t , r o o t ; s c , secondary c u s p s , numbered one through f i v e .

3.

PLATE 1. Heptranchias p e r l o , LACM F-468, c o l l e c t e d in Suruga Bay, Shizuoka, Japan; upper and lower l e f t t o o t h s e r i e s : lacking L ' in the upper s e r i e s and p o s t e r i o r s in both upper and lower s e r i e s . A b b r e v i a t i o n s : A, a n t e r i o r ; L, l a t e r a l ; M, m e d i a l ; S, symphysial; s c a l e l i n e = 5 mm.

4.

I

5.

I t i s t h e p u r p o s e of t h i s p a p e r t o r e c o n s i d e r t h e s p e c i e s Not i d a n i o n h o w e l l i i R e e d , 1946 and t o d i s c u s s i t s c h r o n o l o g i c , b i o g e o g r a p h i c and p a l e o e c o l o g i c o c c u r r e n c e i n t h e E o c e n e of w e s t e r n O r e g o n , W a s h i n g t o n , and V a n c o u v e r I s l a n d , B r i t i s h C o l u m b i a . ABBREVIATIONS The f o l l o w i n g a b b r e v i a t i o n s a r e u s e d : ANSP, Academy of N a t u r a l S c i e n c e s of P h i l a d e l p h i a ; GSC, G e o l o g i c a l S o c i e t y of C a n a d a , O t t a w a ; LACM, N a t u r a l H i s t o r y Museum of Los A n g e l e s C o u n t y , V e r t e b r a t e P a l e o n t o l o g y S e c t i o n ; NSMT, N a t i o n a l S c i e n c e Museum, T o k y o , J a p a n ; PSU, P o r t l a n d S t a t e U n i v e r s i t y , E a r t h S c i e n c e s Museum, P o r t l a n d , O r e g o n ; UCMP, U n i v e r s i t y of C a l i f o r n i a Museum of P a l e o n t o l o g y , B e r k e l e y ; UW, B u r k e M e m o r i a l W a s h i n g t o n S t a t e Museum, D i v i s i o n of G e o l o g y and P a l e o n t o l o g y , U n i v e r s i t y o f W a s h i n g t o n , S e a t t l e ; C R - 7 , UW-CR-6, l o c a l i t y numbers u s e d i n u n p u b l i s h e d t e x t of J o h n M. A r m e n t r o u t ' s ( 1 9 7 3 ) U n i v e r s i t y o f W a s h i n g t o n P h . D . d i s sertation. The t o o t h t e r m i n o l o g y u s e d i n t h i s p a p e r i s b a s e d i n p a r t upon A p p l e g a t e ( 1 9 6 5 a ) and Compagno ( 1 9 7 0 ) , and i s i l l u s t r a t e d i n F i g . 1 A and B . ACKNOWLEDGEMENTS I am g r a t e f u l f o r c r i t i c i s m s and a d v i c e o f f e r e d by D r . J . T. G r e g o r y a n d Mr. J . H. H u t c h i s o n of t h e D e p a r t m e n t of P a l e o n t o l o g y , U n i v e r s i t y of C a l i f o r n i a , B e r k e l e y . Dr. S h e l t o n P. A p p l e g a t e , A s s o c i a t e C u r a t o r of V e r t e b r a t e P a l e o n t o l o g y of t h e N a t u r a l H i s t o r y Museum of L o s A n g e l e s C o u n t y , o f f e r e d e n c o u r a g e m e n t and a d vice as well as access to the paleontological collection. Dr. J o h n M. A r m e n t r o u t , now of M o b i l O i l C o r p o r a t i o n , D e n v e r , C o l o r a d o k i n d l y s u p p l i e d l o c a l i t y and s t r a t i g r a p h i c d a t a f o r t h e Washi n g t o n specimen of H e p t r a n c h i a s . Mr. Armando S o l i s of t h e N a t u r a l H i s t o r y Museum of Los A n g e l e s C o u n t y p h o t o g r a p h e d t h e t o o t h s e t of Heptranchias perlo (Plate 1). SYSTEMATICS O r d e r HEXANCHIFORMES B e r g , F a m i l y HEXANCHIDAE G i l l ,

1940 1885

Genus H e p t r a n c h i a s R a f i n e s q u e , ( F i g u r e 1A, B, C; P l a t e s

1,

1810 2)

Diagnosis: T e e t h d i f f e r e n t in u p p e r and l o w e r j a w , w i t h u p p e r a n t e r i o r s bearing a s i n g l e commissurally flexed primary cusp lacking symphysial d e n t i c l e s or secondary cusps; teeth strongly d i f f e r e n t b e t w e e n l o w e r M, and L . and b e t w e e n l a t e r a l s and p o s t -

6.

t e r i o r s of both upper and lower s e r i e s ; lower l a b i a l s comb-like w i t h a high primary c u s p , one t o f i v e d e n t i c l e s on the symphysia l edge of the primary c u s p , and four t o nine secondary cusps of ungraduated h e i g h t ; upper l a t e r a l s s m a l l e r than lowers with a s i n g l e primary c u s p , one or two secondary cusps and one or two symphysial d e n t i c l e s ; length of crown foot and d e n t i c l e number i n c r e a s e i n s i z e toward commissure in upper l a t e r a l s ; r o o t s und i v i d e d , t a b u l a r except in p o s t e r i o r s which a r e d o r s o v e n t r a l l y depressed and s m a l l ; lower and upper p o s t e r i o r s s i m p l e , extremely s m a l l , e l o n g a t e w i t h low crown which lacks c u s p s . Type S p e c i e s :

Heptranchias c i n e r e u s Rafinesque, 1810

Heptranchias h o w e l l i i (Reed, 1946), new combination Notidanion h o w e l l i i Reed, 1946, Notulae Naturae Acad. Nat. S c i . P h i l a d e l p h i a 172: 1-3. Revised D e s c r i p t i o n of Holotype: Lower l e f t ?fourth l a t e r a l t o o t h p o s s e s s i n g f i v e d e n t i c l e s on the symphysial edge of the primary cusp near t h e crown b a s e . The d e n t i c l e s i n c r e a s e gradua l l y in s i z e toward the cusp apex. The primary cusp i s twice a s l a r g e as s c , , broad a t i t s base w i t h s t r a i g h t symphysial and comm i s s u r a l e d g e s ; and i s followed by f i v e secondary c u s p s ; sc and s c . a r e e q u a l in h e i g h t , s c , and s c , a r e t a l l e r t h a n t h e l a t t e r two and a l s o equal in h e i g h t , sec i s one t h i r d t h e h e i g h t of s c , . The d e n t i c l e s , primary cusp and secondary cusps a r e i n c l i n e d commissurally a t an a n g l e of a p p r o x i m a t e l y f i f t y d e g r e e s . A hori z o n t a l l i n e i s formed on t h e l i n g u a l t o o t h s u r f a c e by t h e j u n c t i o n of t h e crown foot and r o o t . A rounded t r a n s v e r s e l i n g u a l r i d g e extends the l e n g t h of the r o o t , approximately 1 mm below the crown b a s e , bending s l i g h t l y upward a t t h e commissural end and terminated a t t h e symphysial b o r d e r by a s l i g h t d o r s o v e n t r a l d e p r e s s i o n . The r o o t i s t a b u l a r , somewhat r e c t a n g u l a r and d o r s o v e n t r a l l y narrowed toward the commissural end. In c r o s s s e c t i o n t h e r o o t i s wedge-shaped, t h i c k e s t a t i t s j u n c t i o n w i t h the crown b a s e and t a p e r i n g toward the p o s t e r i o r r o o t b o r d e r . The t o o t h i s f l a t on the l a b i a l s i d e , l i n g u a l l y convex. A l l cusps lack s e r r a t i o n s and a r e "so t h i n a s t o be almost t r a n s l u c e n t ; the t o o t h as a whole i s s l i g h t l y t w i s t e d , but the cusps and d e n t i c l e s a r e s e t s t r a i g h t on the r o o t , a n d a r e n o t a t a l l sigmoid" (Reed, 1946). Measurements: T o t a l h e i g h t 7 mm (measured from the t i p of the primary cusp t o t h e r o o t b a s e , p e r p e n d i c u l a r t o t h e v e n t r a l margin of the r o o t ) ; t o t a l l e n g t h 13 mm (measured from the symphysi a l t o commissural margins a l o n g the crown f o o t ) . D i s c u s s i o n : Heptranchias h o w e l l i i d i f f e r s from H^. e z o e n s i s Applegate and Uyeno, 1968, NSMT 7421, in having narrower cusp b a s e s , a narrower and high primary cusp and more than two dent i c l e s on the symphysial edge of the primary c u s p . A h i g h e r 7.

primary cusp and the p r e s e n c e of no more t h a n t h r e e d e n t i c l e s on t h e lower l a t e r a l t e e t h d i f f e r e n t i a t e s the e x t a n t s p e c i e s H_. p e r l o ( B o n n a t e r r e , 1788) from H. h o w e l l i i . The Hexanchidae have w e l l developed h e t e r o d o n t y , both a l o n g a s i n g l e t o o t h s e r i e s and between upper and lower j a w s , w i t h m e d i a l , s y m p h y s i a l , a n t e r i o r , l a t e r a l , and p o s t e r i o r t o o t h t y p e s (Appleg a t e , 1965a). H e p t r a n c h i a s p e r l o has t h e following d e n t a l formula: IACM F - 4 6 8 , Male: P10 L7 A3 SI SI A3 L7 Pll Left Right Pll

L5

Ml

L5

P14

T o o t h p o s i t i o n s L.. t h r o u g h L . i n t h e l o w e r d e n t i t i o n of H e p t r a n c h i a s p e r l o ( P l a t e 1 ) a r e d i s t i n g u i s h e d by t h e s i z e a n d number o f c u s p s , number a n d a r r a n g e m e n t of d e n t i c l e s , h e i g h t a n d w i d t h r a t i o s , and r o o t m o r p h o l o g y . T e e t h i n t h e f e m a l e s of II. p e r l o b e a r a l a r g e r number of s e c o n d a r y c u s p s t h a n i n m a l e s of t h e s p e c i e s ( B i g e l o w and S c h r o e d e r , 1 9 4 8 ) . C o n f i r m a t i o n of t h e p r e s e n c e o r a b s e n c e of t h i s d i m o r p h i s m i n H. h o w e l l i i and t h e a c c u r a c y of my i n t e r p r e t a t i o n s of t h e t o o t h p o s i t i o n s m u s t a w a i t t h e e x a m i n a t i o n of a d d i t i o n a l s p e c i m e n s when a r r a n g e d i n t o o t h s e t s . OCCURRENCES OF HEPTRANCHIAS HOWELLII Among t h e T e r t i a r y s e l a c h i a n a s s e m b l a g e s from O r e g o n , W a s h i n g t o n , and B r i t i s h C o l u m b i a i n t h e c o l l e c t i o n s a t P o r t l a n d S t a t e U n i v e r s i t y , t h e U n i v e r s i t y of C a l i f o r n i a Museum of P a l e o n t o l o g y a t B e r k e l e y , t h e B u r k e M e m o r i a l W a s h i n g t o n S t a t e Museum, and t h e G e o l o g i c a l S u r v e y of C a n a d a , a r e s i x t e e t h w h i c h a r e r e f e r a b l e t o Heptranchias howellii. Oregon Referred Specimens: PSU F - 4 1 9 , f i r s t l o w e r l e f t l a t e r a l UCMP 5 7 4 5 6 , f o u r t h l o w e r l e f t l a t e r a l t o o t h ( P l a t e 2 B ) .

tooth;

F o r m a t i o n ^ a n d A g e : M i d d l e member of t h e K e a s e y F o r m a t i o n ( S c h e n c k , 1 9 2 7 ; W a r r e n and N o r b i s r a t h , 1 9 4 6 ) , Lower R e f u g i a n (Upper E o c e n e ) . Locality: UCMP l o c . V 6 7 3 0 , 5 ' a b o v e b a s e of a p r o m i n e n t b l u f f , d u r i n g t h e A u g u s t low w a t e r s t a g e , on W s i d e of Nehalem R i v e r , i m m e d i a t e l y S of t h e Burn Road B r i d g e , 0 . 3 m i l e s S of j u n c t i o n o f Oregon S t a t e Highways 4 7 a n d 202 a t M i s t , O r e g o n : NE 1/4 NE 1/4 SE 1/4 S e c . 2 3 , T5N, R5W, B i r k e n f i e l d Q u a d r a n g l e ( 1 5 ' , 1 9 5 5 ) , Columbia County, Oregon. R e f e r r e d S p e c i m e n : PSU F - 4 2 6 , (Plate 2A).

lower r i g h t f i f t h

8.

lateral

tooth

Formation and Age: Nestucca Formation (Snavely and Vokes, 1949), lower N a r i z i a n (Bulimina c o r r u g a t a zone of M a l l o r y , 1959). L o c a l i t y : 10' above beach and approximately 100' below a 400' t h i c k flow of b a s a l t , on N shore of Salmon River a t Cascade Head, 1/4 m i l e from mouth of the Salmon River and the P a c i f i c Ocean and 304' N of Teal Creek, in NW 1/4 SE 1/4 Sec. 14, T5S, R11W, Hebo Quadrangle ( 1 5 ' , 1955), Tillamook County, Oregon. Washington Referred Specimen: UW 60692, a n t e r i o r h a l f ( d e n t i c l e s , primary cusp and s c ^ , s c ? ) of a lower r i g h t l a t e r a l t o o t h w i t h mold of sc3 - sc5. Formation and^ Age: Lincoln Creek Formation, Echinophoria d a l l i z o n e , lower Refugian. L o c a l i t y : UW-B0268, Canyon River S e c t i o n , t r a v e r s e u p - s e c t i o n a l o n g the Canyon River from the n o r t h in Sec. 18, T21N, R6W, toward the s o u t h to Sec. 36, T21N, R6W, Mt. Tebo and G r i s d a l e Q u a d r a n g l e s , ( 1 5 ' , 1953), Mason and Grays Harbor C o u n t i e s , Washi n g t o n ; Loc. CR-7, g l a u c o n i t i c s i l t y s a n d s t o n e ; 3 ' t h i c k bed w i t h thanatocoenose fauna, in g l a u c o n i t e sand 6' s t r a t i g r a p h i c a l l y above Loc. UW-CR-6, j u s t above deep pool a t SE c o r n e r of meander bend, 4700' W, 800' N of SE c o r n e r , Sec. 18, T21N, R6W (Personal communication, J . Armentrout, 1974). Vancouver I s l a n d , B r i t i s h Columbia Referred Specimens: GSC No. 21734 A and GSC No. 21734 B, two lower l a t e r a l t e e t h . Formation and Age: J e l e t z k y ' s " D i v i s i o n C" of Unit 2 of the Tert i a r y beds of the Hesquiat P e n i n s u l a , Vancouver I s l a n d (Waldman, 1971), lower Refugian (Sigmomorphina s c h e n c k i zone) (Cameron, 1971). L o c a l i t y : "West c o a s t of Vancouver I s l a n d , B r i t i s h Columbia, e a s t s i d e of Hesquiat P e n i n s u l a , 0 . 6 m i l e s (1 km) s o u t h of Hesquia t v i l l a g e , 20 yards (18 m) from t h e beach below h i g h water mark (Waldman, 1 9 7 1 ) . " Specimen Number PSU F-419 PSU F-426 UCMP 57456 UW 60692 GSC 21734A

Measurements T o t a l Height 3.4 mm 7.5+ mm 9.9 mm 9.1+ mm

GSC 21734B

9.

T o t a l Length 6.0+ mm 15.4+ mm 17.0+ mm 6.1+ mm 19.0 mm (Waldman, 1971) 25.0 mm (Waldman, 1971)

CHRONOSTRATIGRAPHIC DISTRIBUTION OF HEPTRANCHIAS HOWELLII H. h o w e l l i i i s a m i n o r b u t c h a r a c t e r i s t i c e l e m e n t of t h e E o c e n e d e e p w a t e r f i s h f a u n a s from w e s t e r n Oregon and W a s h i n g t o n and s o u t h w e s t e r n B r i t i s h Columbia. Its chronostratigraphic d i s t r i b u t i o n i s p r e s e n t l y r e s t r i c t e d t o s t r a t a of N a r i z i a n - R e f u g i a n a g e ( F i g . 2 ) . H e p t r a n c h i a s h o w e l l i i f i r s t appears in the Bulimina c o r r u g a t a z o n e w i t h i n t h e u p p e r p a r t o f t h e m i d d l e member of t h e N e s t u c c a F o r m a t i o n ( s t r a t i g r a p h i c d i v i s i o n b a s e d upon u n p u b l i s h e d d a t a , p e r s o n a l c o m m u n i c a t i o n , A. D . C a l l e n d a r , 1974) and i t s l a t e s t o c c u r r e n c e f a l l s w i t h i n t h e S i g m o m o r p h i n a s c h e n c k i z o n e of l o w e r Refugian a g e . This l a s t b i o s t r a t i g r a p h i c zone i s represented in s t r a t i g r a p h i c D i v i s i o n " C " of J e l e t z k y ( 1 9 5 4 ) a t H e s q u i a t P e n i n s u l a on V a n c o u v e r I s l a n d , B r i t i s h C o l u m b i a , w i t h i n t h e l o w e r L i n c o l n C r e e k F o r m a t i o n o f s o u t h w e s t e r n W a s h i n g t o n ( p e r s o n a l commun i c a t i o n , J . M. A r m e n t r o u t , 1973) and p e r h a p s i n t h e u p p e r m i d d l e member o f t h e K e a s e y F o r m a t i o n , C o l u m b i a C o u n t y , O r e g o n ( F i g . 2 ) . BIOGEOGRAPHY AND PALEOECOLOGY Upper b a t h y a l t o l o w e r n e r i t i c s e d i m e n t s of l a t e E o c e n e a g e w e r e d e p o s i t e d a l o n g t h e w e s t e r n m a r g i n of a t e c t o n i c a l l y a c t i v e t r e n c h i n w e s t e r n O r e g o n , W a s h i n g t o n , and B r i t i s h C o l u m b i a ( S n a v e l y and W a g n e r , 1 9 6 3 ) . R e g i o n a l s u b s i d e n c e i s c o r r e l a t e d w i t h t h e s u b s e q u e n t a p p e a r a n c e of b a t h y a l and n e r i t i c m o l l u s c a n , f o r a m i n i f e r a l and s e l a c h i a n f a u n a s . The s i m i l a r i t y o f i n f e r r e d d e p o s i t i o n a l environments between t h e N e s t u c c a , Keasey, Lincoln C r e e k and V a n c o u v e r I s l a n d s e d i m e n t s ( J e l e t z k y ' s D i v i s i o n "C") and s u g g e s t e d c h r o n o s t r a t i g r a p h i c c o r r e l a t i o n b e t w e e n t h e s e f o r m a t i o n s ( F i g . 2) s u p p o r t s t h i s v i e w . Heptranchias h o w e l l i i appears t o be a geographically limited and n a r r o w l y a d a p t e d member of t h e b a t h y a l - n e r i t i c f a u n a i n t h e North P a c i f i c Ocean. H e p t r a n c h i a s p e r l o o c c u r s t o d a y i n w a t e r s of t h e n o r t h A t l a n t i c , o f f S o u t h A f r i c a , A u s t r a l i a , and J a p a n . I t is generally cons i d e r e d t o b e a b o t t o m f e e d i n g d e e p w a t e r s h a r k and h a s b e e n r e p o r t e d a t d e p t h s of 3 8 0 t o 4 6 0 m e t e r s o f f P o r t u g a l and from d e e p w a t e r o f f Cuba ( B i g e l o w and S c h r o e d e r , 1 9 4 8 : 9 1 ) . If H e p t r a n c h i a s i s indeed a deep w a t e r genus then i t i s not s u r p r i s i n g t h a t i t should appear in the aforementioned d e p o s i t s . Moore and Vokes ( 1 9 5 3 ) c i t e d b o t h f a u n a l and l i t h o l o g i c c h a r a c t e r i s t i c s of t h e K e a s e y F o r m a t i o n a s i n d i c a t i v e of a d e p t h of d e p o s i t i o n b e t w e e n 600 and 3 , 0 0 0 f e e t . A s p e c i f i c d e p t h of 365 m was p r o p o s e d f o r t h e K e a s e y F o r m a t i o n a t M i s t , Oregon (UCMP V6730) by Z u l l o et_ a l . ( 1 9 6 4 ) . W e l t o n ( 1 9 7 3 ) l i s t e d a s e l a c h i a n f a u n a f o r t h e K e a s e y F o r m a t i o n a t M i s t , Oregon t h a t i n c l u d e d N o t o r h y n c h u s , I s u r u s , O d o n t a s p i s , C e n t r o p h o r u s , S q u a t i n a , and P r i s t i o p h o r u s • W i t h t h e e x c e p t i o n of I s u r u s , a l l of t h e a b o v e t a x a a r e k n o w n , o n t h e b a s i s of e x t a n t s p e c i e s , t o i n h a b i t d e p t h s e q u a l t o o r g r e a t e r

10.

California

o c

Benfhic

Foraminiferal Stages

Colifornio Benthic Foraminiferal

Zones

Offer Kleinpell, 1936 Cuihmon 8

Washington Benfhic Forominiferol Offer

Zones

Row, 1966

Hesquiat

Peninsula,

Southwestern

British Columbia

Columbia County ,

Washington

Vancouver Island, ofW

Armentrout, 1973

Si-myisoo, 1 9 4 4

Oregon after

Worren el o l .

Yaqulno

Bay

S e c t i o n , Oregon after

Snovely et a)..

1945-1946

Mallory. 1959

after Cameron, 1971

Uvigerinella

UJ

sparsicostata

z:

Upper

Zemorrian Nye Mudstone Scappoose

LLJ

Lincoln

o

Uvigerina gallowayi

Lower

Formation

Zemorrian

Yaquina F o r m a t i o n

-J

o

Uvigerina

Cassidulina

vtcksburgensis

golvinensis

Division C

P i t t s b u r g Bluff Formation

Siltstone of Alsea

Formation

REFUGIAN Uvigerina

Sigmomorphina

cocoaensis

schencki

Keasey

Formation

UJ LU O O LU

Amphimorphina

Nestucca

jenkinsi Not

NARIZIAN Bulimina

Differentiated

Cowlitz

Formation

Cowlitz Formation

corrugata

Figure 2. Correlation chart of stratigraphic units discussed in text.

Formation

12.

PLATE 2. A. Heptranchias h o w e l l i i , PSU F-426, Nestucca Format i o n , Tillamook County, Oregon; s c a l e l i n e = 2 mm. B. Heptranc h i a s h o w e l l i i , UCMP 57456, Keasey Formation, Columbia County, Oregon; s c a l e l i n e = 2 mm.

13.

t h a n 365 m. Armentrout ( p e r s o n a l communication, 1973) i n t e r p r e ted t h e Lincoln Creek Formation a t UW l o c a l i t y B-0268 a s d e p o s i t ed in middle n e r i t i c depths of 30 t o 100 fathoms with open marine c i r c u l a t i o n . The Nestucca Formation of Oregon, a t Cascade Head, may have been d e p o s i t e d in waters from 600 t o 6,000 f e e t in d e p t h . This i s based upon molluscan and f o r a m i n i f e r a l evidence ( p e r s o n a l communication, A. D. C a l l e n d a r , 1974). Cameron (1971) i n t e r p r e t s J e l e t z k y ' s s t r a t i g r a p h i c D i v i s i o n "C" on Hesquiat P e n i n s u l a , Vancouver I s l a n d , B r i t i s h Columbia, a s a deep water d e p o s i t . LITERATURE CITED A g a s s i z , L. , 1843. Recherches s u r l e s poissons f o s s i l e s . 4 ° , NeuchStel and S o l e u r e , 3 : 1-390. A p p l e g a t e , S. P . , 1965a. Tooth terminology and v a r i a t i o n i n s h a r k s w i t h s p e c i a l r e f e r e n c e t o the sand shark C a r c h a r i a s t a u r u s Rafinesque. Los Angeles Co. Mus. Contr. S c i . 86: 1-18. , 1965b. A c o n f i r m a t i o n of the v a l i d i t y of Notorhynchus pect i n a t u s ; the second record of t h i s upper Cretaceous cowshark. B u l l . So. C a l i f . Acad. S c i . 64 ( 3 ) : 122-125. A p p l e g a t e , S. P . , and T. Uyeno, 1968. The f i r s t d i s c o v e r y of a f o s s i l t o o t h belonging t o the shark genus Heptranchias w i t h a new P r i s t i o p h o r u s s p i n e , both from the Oligocene of Japan. B u l l . Nat. S c i . Mus. 11 ( 1 ) : 195-200, 1 p i . Arambourg, C . , 1927. Les poissons f o s s i l e s d'Oran. Mater. Carte Geol. A l g e r i e , ( I ) P a l e o n t . , ( 6 ) : 222-223. Armentrout, J . M., 1973a. S t r a t i g r a p h i c r e l a t i o n s h i p s of the Narizian-Refugian boundary in Washington. Geol. Soc. Amer. A b s t . with Programs 5 ( 1 ) : 4 . , 1973b. Molluscan paleontology and b i o s t r a t i g r a p h y of the Lincoln Creek Formation, Late Eocene - Oligocene, southwestern Washington. Unpublished Ph.D. D i s s e r t a t i o n , U n i v e r s i t y of Washington, 2: 479 pp. Bigelow, H. B . , and W. C. Schroeder, 1948. F i s h e s of the western North A t l a n t i c . Mem. Sears Found. Mar. R e s . , P t . 1: 1-576. Cameron, B. E. B . , 1971. T e r t i a r y s t r a t i g r a p h y and microfaunas from t h e Hesquiat-Nootka a r e a , west c o a s t of Vancouver Island (92E). Geol. Surv. Canada Paper 7 1 - 1 : 91-94. Campagno, L. J . V., 1970. Systematics of the genus H e m i t r i a k i s ( S e l a c h i i : C a r c h a r h i n i d a e ) , and r e l a t e d g e n e r a . Proc. C a l i f . Acad. S c i . 38 ( 4 ) : 63-98. Cushman, J . A . , and R. R. Simonson, 1944. Foraminifera from the Tumey Formation, Fresno County, C a l i f o r n i a . J o u r . Paleon. 18: 186-203, p i s . 30-34. Cuvier, G., 1817. Le rggne animal d i s t r i b u e d ' a p r e s son o r g a n i s a t i o n . 1 s t e d . , P a r i s , 1: 540 pp. , 1829. Le regne animal d i s t r i b u e d ' a p r e s son o r g a n i s a t i o n . 2nd ed. , P a r i s , 5 v o l s . Garman, S . , 1913. The Plagiostoma ( s h a r k s , s k a t e s , and r a y s ) .

14.

Mem. Mus. Comp. Z o o l . (Harvard) 3 6 : 1-528. Hickman, C. S . , 1972. Review of the b a t h y a l gastropod genus Phane r o l e p i d a (Homalopomatinae) and d e s c r i p t i o n of a new s p e c i e s from the Oregon Oligocene. V e l i g e r 15 ( 2 ) : 107-112. J e l e t z k y , J . A . , 1954. T e r t i a r y rocks of the Hesquiat-Nootka a r e a , west c o a s t of Vancouver I s l a n d , B r i t i s h Columbia. Geol. S u r v . Canada Paper 53 ( 1 7 ) : 1-65. J o r d a n , D. S . , 1907. The f o s s i l f i s h e s of C a l i f o r n i a w i t h s u p p l e mentary notes on o t h e r s p e c i e s of e x t i n c t f i s h e s . Univ. C a l i f . P u b i s . Geol. 5 ( 7 ) : 95-144, p i s . 11-12. K l e i n p e l l , R. M., 1938. Miocene s t r a t i g r a p h y of C a l i f o r n i a . T u l s a , O k l a . : Amer. Assoc. P e t r o l . G e o l . : 450 p p . , 22 p i s . L e r i c h e , M., 1908. Observations de l e s Squales neogenes de l a C a l i f o r n i e . Ann. Soc. Geol. Nord. 3 7 : 302-306. M a l l o r y , V. S . , 1959. Lower T e r t i a r y b i o s t r a t i g r a p h y of the C a l i f o r n i a c o a s t r a n g e . T u l s a , O k l a . , Amer. Assoc. P e t r o l . G e o l . : 416 pp. Moore, R. C . , and H. E. Vokes, 1953. Lower T e r t i a r y c r i n o i d s from n o r t h w e s t e r n Oregon. U. S. Geol. Surv. Prof. Paper 233-E: 113-147, p i s . 14-24. P l e d g e , N. S . , 1967. F o s s i l elasmobranch t e e t h of South A u s t r a l ia and t h e i r s t r a t i g r a p h i c d i s t r i b u t i o n . T r a n s . Roy. Soc. So. A u s t . 9 1 : 135-160. Rau, W. W., 1966. S t r a t i g r a p h y and Foraminifera of the Satsop River A r e a , Southern Olympic P e n i n s u l a , Washington. Wash. Div. Mines Geol. B u l l . 53: 66 p p . Reed, M. D . , 1946. A new s p e c i e s of f o s s i l shark from New J e r s e y . Notulae Naturae Acad. Nat. S c i . P h i l a d e l p h i a 172: 1-3. Schenck, H. G., 1927. Marine Oligocene of Oregon. Univ. C a l i f . P u b l . B u l l . Dept. Geol. S c i . 16 ( 1 2 ) : 449-460. Snavely, P. D . , J r . , and H. E. Vokes, 1949. Geology of the c o a s t a l a r e a between Cape Kiwanda and Cape Foulweather, Oregon. U. S. Geol. Surv. P r e l i m . Map 97, Oil and Gas I n v e s t i g a t i o n s Ser. Snavely, P. D . , J r . , and H. C. Wagner, 1963. T e r t i a r y g e o l o g i c h i s t o r y of western Oregon and Washington. Washington Div. Mines G e o l . , Report of I n v e s t i g a t i o n s No. 22: 1-25. Waldman, M., 1971. Hexanchid and o r t h a c o d o n t i d shark t e e t h from the lower T e r t i a r y of Vancouver I s l a n d , B r i t i s h Columbia. Canadian J o u r . E a r t h S c i . 8 ( 1 ) : 166-170. Warren, W. C , and H. N o r b i s r a t h , 1946. S t r a t i g r a p h y of the upper Nehalem River b a s i n , n o r t h w e s t e r n Oregon. Amer. A s s o c . P e t r o l . Geol. B u l l . 3 0 : 213-237. Welton, B. J . , 1972. F o s s i l sharks in Oregon. The Ore Bin, Oregon Dept. Geol. Mineral I n d u s t r i e s 34 ( 1 0 ) : 161-170, 1 p i . , 1973. Oligocene s e l a c h i a n s from the Keasey Formation a t M i s t , Oregon. Geol. Soc. Amer. Abst. w i t h Programs 5 ( 1 ) : 121. Z u l l o , V., R. F. K a r r , J . W. Durham, and E. C. A l l i s o n , 1964. The echinoid genus S a l e n i a in the e a s t e r n P a c i f i c . Paleont. 7 ( 2 ) : 331, p i . 56.

15.

PALEOBIOS, No. 1 7 ,

1974.

i