STRATIGRAPHIC MICROPALEONTOLOGY OF ATLANTIC BASIN AND BORDERLANDS
FURTHER TITLES IN THIS SERIES 1. A.J. Boucot EVOLUTION AND EXTINCTION RATE CONTROLS
2. W.A. Berggren and J.A. van Couvering THE LATE NEOGENE - BIOSTRATIGRAPHY, GEOCHRONOLOGY AND PALEOCLIMATOLOGY O F THE LAST 1 5 MILLION YEARS IN MARINE AND CONTINENTAL SEQUENCES 3. L.J. Salop PRECAMBRIAN OF THE NORTHERN HEMISPHERE
4. J.L. Wray CALCAREOUS ALGAE 5 . A. Hallam (Editor) PATTERNS OF EVOLUTION, AS ILLUSTRATED BY THE FOSSIL RECORD
This book is produced by a photographic offset process directly from the manuscript. Thus. the publisher is not responsible for any errors appearing in the book.
Developments in Palaeontology and Stratigraphy, 6
STRATIGRAPHIC MICROPALEONTOLOGY OF ATLANTIC BASIN AND BORDERLANDS Edited by
F.M. SWAIN Department of Geology, University of Delaware, Newark, Del. ( U . S . A . ) and Department of Geology and Mineralogy, University of Minnesota, Minneapolis, Minn. (U.S.A.)
ELSEVIER SCIENTIFIC PUBLISHING COMPANY 1977 Amsterdam - Oxford - New York
ELSEVIER SCIENTIFIC PUBLISHING COMPANY 335 Jan van Galenstraat P.O. Box 211, Amsterdam, The Netherlands
Distributors f o r the United States and Canada. ELSEVIER NORTH-HOLLAND INC. 52, Vanderbilt Avenue New York, N.Y. 10017
Library of Congress Cataloging in Publicalion Data
Main e n t r y under t i t l e : S t r a t i g r a p h i c rricropaleontology of A t l a n t i c b a s i n and border l a n d s . (Developments i n palaeontology and s t r a t i g r a p h y ;
5) Papers p r e s e n t e d a t a symposium supported by and convened a t t h e U n i v e r s i t y of Delaware, J u n e 14-16,
1976. Bibliography: p. InclJides indexes. 1. Eicropaleontology--Worth A t l a n t i c Oceany--~o~t~ Congresses. 2. ~ ~ c r o ~ a l e o ~ ’ t o l o g y A-nerica-Conpeesses. 5 . Geology, S t r a t i g r a p h i c - - C o n g r e s s e s . 1. Geology--North A t l a n t i c Ocean--Congresses. 5. G e o l o p - - N o r t h America--Congresses. I. Swain, 11. Delaware. F r e d e r i c k M o r n l l , 1916U n i v e r s i t y , Newark. 111. S e r i e s . QE719.S83 560’ .921 77-915 ISBN
0-444-41554-8
0 Elsevier Scientific Publishing Company, 1 9 7 7 .
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the publisher, Elsevier Scientific Publishing Company, P.O.Box 330,Amsterdam, The Netherlands
Printed in The Netherlands
V
PREFACE A symposium on the stratigraphic micropaleontology of the Atlantic basin and margins was convened at the University of Delaware, June 14-16, 1976. The symposium was attended by 56 people, and 27 papers were presented. The volume represents the published proceedings of the symposium and consists of 26 papers by 28 authors. Although a serious attempt was made to provide at least minimum coverage of all major micropaleontologicd groups in and around the Atlantic, these efforts were not successful in the case of diatoms, Paleozoic calcareous Foraminifera, coccoliths and Paleozoic and Cenozoic palynology. The volume includes summaries and discussions of the stratigraphic and geographic distribution of Paleozoic, Mesozoic and Cenozoic Foraminifera, Radiolaria and Ostracoda; Paleozoic and Mesozoic conodonts; Paleozoic Chitinozoa and acritarchs; Mesozoic dinoflagellates, and palynomorphs; and Cenozoic silicoflagellates. The discussions of individual papers represent transcripts of tapes made at the meeting, written questions submitted at the meeting, and subsequent modifications of questions and answers by questioners and authors, through correspondence. Several participants in the symposium were unable t o submit final manuscripts or plan t o publish their papers elsewhere; G.L. Williams and J.P. Bujak, Geological Survey of Canada (Cenozoic palynology ), Stefan Gartner, Texas A. and M. University (Cenozoic coccoliths), F.M. Gradstein, Geological Survey of Canada (Jurassic Foraminifera), and Piero Ascoli, Geological Survey of Canada (Biostratigraphy of Scotian Shelf). The papers on South Atlantic Cretaceous Ostracoda and Paleogene and Neogene Foraminifera contain more details on localities than most of the other papers. I t was decided t o retain this information because some of it is not easily accessible t o workers in the Northern Hemisphere. It was originally planned that a summary chapter be prepared on Atlantic stratigraphic micropaleontology, with the collaboration of all the authors. Limitations on the length of the book have prevented this being done. ACKNOWLEDGEMENTS
I am greatly indebted t o the Graduate School and the College of Arts and Sciences of the University of Delaware for support of the symposium and to Dr. John C. Kraft, Chairman of the Geology Department who made the arrangements for support. Michelle Mayrath prepared illustrations for my ostracode and acritarch papers, and also provided invaluable assistance in editorial matters. Teresa Grilli typed or re-typed many of the manuscripts. Patricia Barlow, Barbara Frank, and Sarah Cornell also assisted in secretarial and editorial work. F.M. SWAIN
This Page Intentionally Left Blank
VII
CONTENTS Preface . . . . . . . . . . . . . . . . . . . . . . . . Early Paleozoic Ostracoda of Eastern Canada. M.J. COPELAND . . . . . Early Paleozoic Ostracoda of the Atlantic Margin. other than Eastern Canada. F.M. SWAIN . . . . . . . . . . . . . . . . . . . . . Paleozoic Smaller Foraminifera of the North American Atlantic Borderlands. JAMES E CONKIN and BARBARA M CONKIN . . . . . . . . . Late Paleozoic Ostracodes of Western Europe and North America: A Review. LUIS C SANCHEZ DE POSADA . . . . . . . . . . . . . . Early Paleozoic Conodont Biostratigraphy in the Atlantic Borderlands. STIG M. BERGSTROM . . . . . . . . . . . . . . . . . . . Late Paleozoic and Triassic Conodont Biostratigraphy : Correlations around the Expanding Atlantic Ocean. DAVID L CLARK . . . . . . . . . . Notes on Paleozoic Acritarchs from the Atlantic Margins. F.M. SWAIN . . . Paleobiogeography of Chitinozoa. D W. ZALUSKY . . . . . . . . . Paleozoic Radiolaria: Stratigraphic Distribution in Atlantic Borderlands. B.K. HOLDSWORTH . . . . . . . . . . . . . . . . . . Autunian and Carnian Palynoflorules: Contribution to the Chronology and Tectonic History of the Moroccan Pre-Atlantic Borderland. HAROLD L . COUSMINER and WARREN MANSPEIZER . . . . . . . . . . Mesozoic Foraminifera - '8,stern Atlantic. RICHARD K . OLSSON . . . . Jurassic Ostracoda of the Atlantic Basin. R.H. BATE . . . . . . . . . Cretaceous Ostracoda of the North Atlantic Basin. JOHN W NEALE . . . Cretaceous Ostracoda - South Atlantic. ALWINE BERTELS . . . . . . Mesozoic Radiolaria from the Atlantic Basin and its Borderlands. HELEN P FOREMAN . . . . . . . . . . . . . . . . . . . . . Jurassic Palynostratigraphy of Offshore Eastern Canada. JONATHAN P. BUJAK and GRAHAM L WILLIAMS . . . . . . . . . . . . . Comparison of Lower and Middle Cretaceous Palynostratigraphic Zonations in the Western North Atlantic. DANIEL HABIB . . . . . . . . . Upper Cretaceous Dinoflagellate Zonation of the Subsurface Toms River Section Near Toms River. New Jersey. RICHARD AURISANO and DANIEL HABIB . . . . . . . . . . . . . . . . . . . . North Atlantic Cenozoic Foraminifera. W.A. BERGGREN . . . . . . . Paleogene Foraminifera - South Atlantic. ALWINE BERTELS . . . . . Neogene Foraminifera - South Atlantic. ALWINE BERTELS and MARLY MADEIRA-FALCETTA . . . . . . . . . . . . . . . . Cenozoic Ostracoda - North Atlantic. M.C. KEEN . . . . . . . . . Cenozoic Marine Ostracoda of the South Atlantic. W.A. VAN DEN BOLD . Cenozoic Radiolarians of the Atlantic Basin and Margins. RICHARD E CASEY and KENNETH J. McMILLEN . . . . . . . . . . . . Atlantic Cenozoic Silicoflagellates. Potential for Biostratigraphic and Paleoecologic Studies. RICHARD E . CASEY . . . . . . . . . . . . North American Microtektites. Radiolarian Extinctions and the Age of the Eocene-Oligocene Boundary. B.P. GLASS and MICHAEL J . ZWART . . Index t o Authors . . . . . . . . . . . . . . . . . . . . . Index t o Genera and Species . . . . . . . . . . . . . . . . .
.
.
* . v . . 1 19
.
.
.
.
.
.
.
.
.
.
49 61
. .
85
. . . . . .
111 131 151
.
. 167
. . 185 . . 205
. . . . .
. 231 . 245 . 211
. 305 .
321
. .
341
. 369 . 389 . 411 . . .
. 441 . 461 .
495
. .
521
.
. 545
. . 553 . . 569 . . 511
This Page Intentionally Left Blank
1 EARLY PALEOZOIC OSTRACODA OFEASTERN CANADA
M . J. Copeland Geological S u r v e y of Canada, Ottawa, Ontario, K1A OE8 Abstract Pre-Middle Devonian OStrQCodQo c c u r in three northeast-trending s t r u c t u r a l provinces of eastern Canada. They comprise biogeographically a n d stratigraphical ly distinctive leperditicopid, palaeocopid and podocopid assemblages dominated by Ordovician hollinids in the n o r t h e r n (Anticosti Basin) province a n d by SiluroDevonian beyrichiids in all three provinces. Correlation is possible only between endemic North American Early Silurian beyrichiid assemblages of the n o r t h e r n and central (Appalachian Belt) provinces; the s o u t h e r n ( F u n d y Belt) province displays the typical Beyrichienkal k fauna of n o r t h e r n Europe. Introduction S o continuous sequence of pre-Middle Devonian palaeocopid or podocopid
ostracode faunas h a s been d e s c r i b e d from e a s t e r n Canada. T h i s is d u e i n p a r t to facies variation and discontinuities i n the geological r e c o r d b u t it is anticipated that future study will reveal a more complete zonation than is now possible. Distinct depositional, temporal, geographic and tectonic control is evident, however, i n the ostracode assemblages presently known.
These faunas a r e
contained i n t h r e e generally parallel s t r u c t u r a l provinces comprising a n older platformal sequence i n Anticosti Basin and two y o u n g e r , shallow water sequences, those of the "Appalachian Belt" i n Gasp6 Peninsula and s o u t h e r n Quebec and the "Fundy Belt" of Maritime Canada and coastal New England ( F i g u r e 1 ) . Assemblages of North American aspect occur on either s i d e of the n o r t h e r n limit of Acadian orogenic deformation. The n o r t h e r n , undeformed St. Lawrence Platform
-
Antieosti Basin faunal sequence reflects a n initially provincial Middle
Ordovician ostracode assemblage, modified somewhat b y Late Ordovician boreal elements, and a subsequently endemic North American Early Silurian ostracode fauna of Appalachian aspect. T h i s endemic fauna, b e a r i n g the only palaeocopid ostracodes presently known to provide correlation a c r o s s the Acadian Front (Figure 2 ) , also o c c u r s i n basal s t r a t a of the deformed Silurian sequence i n Gasp6 Peninsula and m a r k s initiation of the typical northeastern North American Siluro-Devonian Appalachian Belt ostracode succession. It differs completely from the more stratigraphically restricted European beyrichiid fauna of t h e Fundy Belt to the south. These faunas a r e separated i n central New Brunswick b y a
2
EASTERN CANADA Early Paleozoic Sedimentary Basins
1. Port-au-Port Peninsula 4. 7. 10. 13.
Anticosti Island Dalhousie Arisaig E a s tpor t
2. 5. 8. 11. 14.
Table Point 3. Forillon Peninsula 6. Lake Matapedia 9. Portapique River 12. Coast of JIaine (general) 15.
Mingan Islands Mt. Albert area Rloose River Synclinorium Jones Creek Newbury port
Figure 1. Early Paleozoic structural provinces of eastern Canada and northeastern United States, with localities mentioned. wide zone of Silurian turbidites that may represent a marine trough that effectively separated contemporaneous ostracode faunas of North American and European aspect. The time-restricted Late Silurian-Early Devonian Fundy Belt assemblage, of distinctive north European Beyrichienkalk affinity, is contained in a segment of the European continental plate, left behind after collision with the North American plate had resulted in closure of the proto-Atlantic Ocean and its subsequent reopening during the Mesozoic.
3
1
NORTH
I
I
UNDETORMCD
ST.
LAWRCMCE
LOWLAND and Q Y t b W !
AMTICOSTI MAS111
Ionfarlo
8 * a
I
AMERICAN
-
DCFORYCO
APPALACHIAN
I V
MELT
EUROPEAN
A C A D I A M OROGCMV
TUNDV
MELT
, Y
Figure 2 . Faunal and tectonic relationships of ostracode assemblages recognized in eastern Canada History of Investigations Ostracoda have been recorded from relatively few localities in Anticosti Basin (Fig. 1, locs. 1-4). Chief among the early investigators were Billings (1865, 1866) and Jones (1858, 1890a,b, 1891).
During the present century, Ulrich and
Bassler (1923), Bassler (in Twenhofel, 1 9 2 8 ) and Copeland (1970a, b, 1973, 1974a) presented a Late Ordovician and Early Silurian ostracode zonation for Anticosti Island, and Berdan ( i n Whittington and Kindle, 1963) reported on a Middle Ordovician fauna from western Newfoundland (Fig. 1, loc. 2 ) .
Additional
information on younger Early Paleozoic ostracode faunas may be forthcoming when samples of strata from beneath the Gulf of St. Lawrence between Anticosti
4
Island, Quebec (Fig. 1, loc. 4 ) and Port-au-Port Peninsula, Newfoundland (Fig. 1, loc. 1) a r e studied. Middle Ordovician ostracode faunes from St. Lawrence Lowland and related areas surrounding the Precambrian shield of eastern Canada a r e recorded in Kay (1934), Carter (1957) and Copeland (1965, 1970a, 1976, in p r e s s ) , This is probably the most extensively distributed Early Paleozoic ostracode fauna in northern North America a s it i s also reported from Yukon Territory and Districts of Mackenzie and Franklin (Copeland, 1974b, in p r e s s ) , Within the Appalachian Belt the earliest report of Devonian ostracodes was by Jones (1889) from collections made along the southern shore of Chaleur Bay in northern New Brunswick (Fig. 1, loc. 7 ) .
This fauna was redescribed and
numerous additional species were recorded by Clarke (1909) and Copeland (1962) from the vicinity of Campbellton and Dalhousie, New Brunswick. In northern Maine, south of Quebec City, Berdan (in Boucot, 1961) reported a small beyrichiid fauna from the Moose River Synclinorium (Fig. 1, loc. 9) that has proved slightly older than an Early Devonian fauna from Forillon Peninsula (Fig. 1, loc. 5) that was identified independently by Berdan and Copeland and reported in Burk (1964) and Boucot (1965). Because of its European aspect, ostracode faunas from the Fundy Belt (Fig. 1, locs. 10-15) attracted the attention of several early workers during the midnineteenth century. Honeyman (1859, 1864), Dawson (186G, et seq. ) , Hall (1860) and Jones (1870, 1881a, b ) each recorded part of this Beyrichienkalk fauna from Xrisaig, Nova Scotia (Fig. 1, loc. 1 0 ) . In this century, McLearn (1924) and Copeland (1960, 1964) defined the position of this fauna within the thick stratigraphic sequence present in that area. Only recently have additional localities been reported in Nova Scotia and New Brunswick (Fig. 1, locs. 11. 1 2 ) , and Berdan (1966, 1971, in Brookins et a l . , 1973) recorded a longer ranging ostracode fauna from coastal Maine (Fig. 1, locs. 13, 1 4 ) extending southwestward to Massachusetts (Fig. 1, loc. 15). r h z Ostracode Fauna The earliest ostracode fauna of St. Lawrence Lowland-Anticosti Basin is Whiterockian (Llanvernian) of the Table Head Formation of western Newfoundland.
' E o l e p e r d i t i a ' of the general 'Eoleperditia' bivia (White) group i s most significant in that it represents a circum-cratonal North Americar. platform fauna that occurs
5 also in Nevada and southwestern District of .<,&enzie (Copeland, 197413). On I
Mingan Islands (Twenhofel, 1938) and between Quebec and Montreal, a small, relatively unknown ostracode fauna has been reported. There appears to be little similarity between this and other Chazyan ostracode faunas with which i t has been correlated. Middle Ordovician ostracodes of Wildernessian-Barneveldian (Caradocian) age are widespread within the platformal facies north and west of St. Lawrence River (Plate 1). These may be generally designated a s the mid-continental Decorah fauna, after the area in Minnesota and Iowa from which they were described by Kay (1934) and later workers.
Ostracodes of this assemblage occur in two
temporal subassemblages, with older Wildernessian elements in Minnesota, Michigan and southern Ontario, which could be termed 'southern', typified by the genera 'Aparchites', Dicranella, Eurychilina and Ceratopsis; and younger, Barneveldian elements in northern Ontario, Foxe Basin and Baffin Island, of 'boreal' aspect, typified by the genera Oepikella, Ocpikium, Distobolbina, Steusloffina and Levisulculus.
This younger fauna shows distinct north
European affinities, but may be slightly older than its first appearance i n Scandinavia, and marks initiation of a modified North American-European fauna. Late Ordovician faunas a r e best known from Anticosti Island (Fig. 3, Plate 1) in strata of the Vaurkal and Ellis Bay formations.
There, also, a
mixing of North American and Baltic faunas is apparent. Typical North American hollinaceans such a s Tetradella and Anticostiella occur with European Carinobolbina and Foramenella. The upper limit of this Maysvillian-Richmondian (Ashgillian) fauna i s established by the last occurrence of tetradellid ostracodes, which marks the close of the Ordovician in northeastern North America. O n Anticosti Island a conformable sequence of Late Ordovician to Early Silurian strata is present. We have been unable to establish an ostracode succession across this interval as about fifty to sixty-five feet ( 1 5 - 2 0 m ) of strata, barren of palaeocopid ostracodes separate the youngest Ordovician tetradellids and the oldest Silurian beyrichiids. This is the presumed glacial or Cherokee Discontinuity (Dennison and Head, 1975) postulated by Berry and Boucot (1973) , among others, to explain a faunal hiatus during which time glacio-eustatic changes in sea-level caused near complete destruction of all faunas throughout northern North America.
6
SILURIAN LLANDOVERY
ORDOVICIAN ASHGILL
7' WENLOCK
VAUREAL EtLlS BAY BECSCIE GUN RIVER JUPITER CHICOllE CRASPEDOBOLBININAE
19
ZYGOBOLBINAE
3 m
$
SUBFAMILY UNCERTAIN BOLLllDAE RlCHlNlOAE AECHMINIDAE KIRKBYELLIDAE QUADRIJUGATORIDAE SIGMOOPSIOAE HOLLINIDAE TETRAOELLIOAE EURYCHlLlNlDAE CHlLOBOLBlNlOAE
-
-I I
I
4 I
I
I
I
I
I
I
= a r
5
E
1
Figure 3. Stratigraphic distribution of palaeocopid ostracodes, Anticosti Island, Quebec. The transition between Ordovician and Silurian marks one of the most distinct lines of extinction and reestablishment of Early Paleozoic ostracode faunas in eastern North America. The sudden appearance of beyrichiids with the early Niagaran brachiopod Virgiana in the upper membcr of the Becscie Formation is startling. It i s unlikely that this beyrichiid fauna arose from a loculate tetradellid; it i s far more plausible to assume a eurychilinid o r piretellid ancestry. However, the beyrichiids Zygobursa from Anticosti Island and Zygocosta from southern
Ontario mark the earliest o c c u r r e n c e of this fauna i n e a s t e r n North America (Plate 2 ) . The Ulrich and Baesler zygobolbine succession ( 1923) that was cstablished with the appearance of Zygobursa (Copeland, 19708) continued to well within the Llandoverian on Anticosti Island a n d , i n the Appalachian Belt and platform facies from Maryland to southern Ontario, to t h e top of the Wenlockian with
Drepunellina clarki. T h i s fauna is known from nowhere else i n the world; indeed only one zygobolbid g e n u s i s known to occur elsewhere than i n e a s t e r n North America. Other beyrichiids occur with the zygobolbids i n the Gun River and Jupiter formations but the top of the Anticosti Island succession is marked b y
Zygobolba decora, zone fossil to the top of the Lower Clinton Group i n the Appalachian Belt (Fig. 2 ) . Relatively little h a s been published on the ostracode micropaleontology of the Appalachian Belt i n Gasp6 Peninsula.
The earliest assemblage (Fig. 1, loc. 6 ) ,
Zygobolba decora (Billings) i n the Awantjish Formation, is similar to that of the upper J u p i t e r Formation of Anticosti Island.
Deeper water, graptolite-bearing
clastic strata of Wenlockian a g e occur t h e r e a n d it i s only i n the Ludlovian, calcareous Sayabec Formation (Fig. 1, loc. 8) that a n ostracode fauna, h e r e termed kloedenellid, initiated, i n e a s t e r n Canada, what may b e called the Appalachian 'false Kloedenia' assemblage (Plate 3 )
.
The older or kloedenellid
fauna is typified b y primitiopsids, kloedenellids and some t h l i p s u r i d s as well a s
a few smaller beyrichiids.
T h i s fauna i s also p r e s e n t i n the Hardwood Mountain
Formation, late Ludlovian, of northwestern Maine (Fig. 1, loc. 9 ) .
The younger
o r 'false Kloedenia' fauna o c c u r s throughout shallow water Late Cayugan to Helderbergian (Pridolian-Emsian) s t r a t a of the Appalachian Belt ( B e r d a n e t al. ,1969) This fauna contains s u c h distinctive g e n e r a a s Kloedeniopsis, Pintopsis,
Cornikloedenia, Welleria and Zygobeyrichia and t h l i p s u r i d s of many t y p e s . It is not yet a s fully defined i n e a s t e r n Canada a s that demonstrated b y Berdan for the Cobleskill, Coeymans-Manlius, Becraft-Port Ewen-Glenerie and Schoharie formations of New York State b u t it does occur i n t h e St. Albans Formation
of Forillon Peninsula (Fig. 1, loc. 5 ) , Dnlhousie Beds of n o r t h e r n New Brunswick (Fig. 1, loc. 7) and i n s t r a t a of the Grand G r h e Formation of Gasp6. Although faunas of Llandoverian to Ludlovian a g e s h a v e b e e n reported from the Fundy Belt, the earliest datable ostracodes from t h i s province a r e b e y r i c h i i d s
8
typical of, and in most cases conspecific with those of the shallow water benthonic Beyrichienkalk of the Baltic Province and Downtonian of Great Britain (Plate 3 ) . True Kloedenia and Frostiella occur in the Stonehouse Formation of northeastern Nova Scotia (Fig. 1, loc. 1 0 ) with Londinia only in the lower part of the formation and Nodibeyrichia only in the upper part. Londinia and Frostiella occur in the Jones Creek Formation of southern New Brunswick (Fig. 1, loc. 1 2 ) and the Leighton Shale Member of the Pembroke Formation near Eastport, Maine (Fig. 1, loc. 13). Also near Eastport, Nodibeyrichia has been found in thc overlying Hersey Shale Member of the Pembroke Formation, thus equating with the upper Stonehouse Formation. These faunas are considered a s Pridolian and no younger marine ostracodes a r e as yet identified from that part of the Fundy Belt in Canada. In the Eastport Quadrangle of Maine, however, Berdan has discovered a most interesting later ostracode fauna that i s presently being studied. Equivalence with the Gcdinnian of Podolia and Germany is quite possible for this fauna. Provincialism of Ostracode Faunas Questions arise a s to the distinct faunal separation of the North American (St. Lawrence Lowland-Anticosti Basin-Appalachian) and European (Fundy) provinces.
We can speculate very little on the Ordovician faunal history of the
Fundy Belt but do know that the provincial ostracode faunas of northeastern North American and northern European aspect were in contact from at least the late stages of the Middle Ordovician. A s the continents continued to approach, the Anticosti Basin, sheltered to the north and east by the Canadian Shield, remained undeformed by the Acadian Orogeny.
This stable, shallow, platformal area may
have received marine sediments and supported a rich ostracode fauna almost a s long as the deformed Appalachian Belt. This will only be known i f post midLlandoverian calcareous marine strata a r e found in the Gulf of St. Lawrence area between Anticosti Island and Newfoundland.
It would seem logical that an even more complete physical connection may have existed between platformal North American and European faunal elements during the latest Ordovician and earliest Silurian due to eustatic fall of sea level
a s a result of glacial activity. The ancestral, benthonic beyrichiid stock could have been able to migrate through shallow waters to inhabit shores of both approaching continents. In this case it might appear that a west-to-east migration along the Appalachian-Caledonian belt took place as North American beyrichiids
9
seem to have appeared earlier than their European counterparts. Return to normal sea level d u r i n g the later Llandoverian isolated these rapidly evolving faunas, which were separated, possibly b y a remnant s u b s i d i n g t r o u g h of d e e p water, relict of the proto-Atlantic Ocean, a c r o s s which the l a r g e , cruininate, benthonic beyrichiids could not migrate.
T h i s trough today i s r e p r e s e n t e d b y the thick
sequence of Middle and Late Silurian turbidites extending t h r o u g h the central parts of New Brunswick and Maine that i s covered i n e a s t e r n New Brunswick b y continental strata of Devonian and younger Paleozoic age References Bassler, R . S. , 1928. Ostracoda: in Twenhofel, W. H . , Geology of Anticosti Island: Geol. S u r v . C a n . , JIem. 154, p. 340-350. Berdan, J.XI., 1966. Baltic Ostracodes from Maine: U . S. Geol. S u r v . , Prof. Paper 55O-A, p. 111. _ _ _ _ _ _ , 1 9 7 1 . Silurian to Early Devonian ostracodes of European aspect from the Eastport Quadrangle, Maine: Geol. Soc. A m e r . , ( a b s . ) Northeastern Section, p. 18. _ - - _ _ ~, B e r r y , W . B. N . , Boucot, A. J . , Cooper, G . A . , Jackson, D . E . , Johnson, J . G . , Klapper, G . , Lenz, A. C . , hlartinsson, A. Oliver, W . A . , J r . , Rickard, I,. V. and Thorsteinsson, R . , 1969. Siluro-Devonian boundary i n North America: Geol. Soc. Amer. Bull., v. 84, p. 275-284. Berry, W , B. N. and Boucot, A. J . , 1973. Glacio-eustatic control of Late Ordovician-Early Silurian platform sedimentation and faunal changes: Geol. Soc. Amer. Bull., v. 84, p. 275-284. Billings, E . , 1865. Palaeozoic Fossils. Vol. 1, Containing descriptions and figures of new o r little known species of organic remains from the Silurian rocks: Geol. S u r v . C a n . , p. 299, 300. _ ~ _ _ _ ,_ 1 8 6 6 . Catalogues of Silurian fossils of the Island of Anticosti, with descriptions of some new g e n e r a and species: Geol. S u r v . Can. Boucot, A. J . , 1961. S t r a t i g r a p h y of the hloose River S!,mclinorium, Maine: U . S . Geol. S u r v . , Bull. 1111-E. _ _ _ _ _ _ , 1965. Silurian s t r a t i g r a p h y of Gasp6 Peninsula, Quhbec: Bull. Am. Assoc. Petrol. Geols., v. 49, n. 1 2 , p. 2295-2316. Brookins, D . G . , Bcrdan, J . N . nnd Stewart, D . B . , 1973. Isotopic and paleontologic evidence for correlating t h r e e volcanic sequences i n the Rlaine coastal volcanic belt: Geol. Soc. Amer. B u l l . , v . 84, p. 1619-1628. Burk, C . F . , J r . , 1964. Silurian s t r a t i g r a p h y of Gasp6 Peninsula, Qukbec: Bull. A m . -4ssoc. Petrol. Geols., v . 48, n. 4, p. 437-464. Carter, G. F. E. , 1957. Ordovician Ostracoda from the St. Lawrence Lowlands of Qukbec: unpubl. Ph. D. thesis, McGill Univ. , Montreal. Clarke, J . M. , 1909. Early Devonic history of New York and eastern North America: N . Y . State hIus., Mem. 9, pt. 2. Copeland, &I. J., 1960. Ostracoda from the Gpper Silurian Stonehouse Formation, Arisaig Nova Scotia, Canada: Palaeont., v . 3 , pt. 1, p. 93-103. - _ _ _ _ , 1962. Ostracoda from the Lower Devonian Dalhousie b e d s , n o r t h e r n New Brunswick: Geol. S u r v . C a n . , Bull. 91, p. 18-51, pls. V-X. ~
10 Copeland, ill. J . , 1964. Stratigraphic distribution of Upper Silurian Ostracoda, Stonehouse Formation, Nova Scotia: Geol. Surv. Can., Bull. 1 1 7 , p. 1-13, pl. 1. _ _ _ - _ _ , 1965. Ordovician Ostracoda from Lake Timiskaming, Ontario: Geol. Surv. Can., Bull. 127. _ _ _ _ _ _ , 197Oa. Two new genera of beyrichiid Ostracoda from the Niagaran (Middle Silurian) of Eastern Canada: Geol. Surv. Can., Bull. 187, p. 1-7, pl. I, figs. 1, 2. _ _ _ _ - _ , 1970b. Ostracoda from the Vaurbal Formation (Upper Ordovician) of Anticosti Island, Qu6bec: ibid., p. 15-29, pls. IV-V. _ _ _ _ _ _ , 1973. Ostracoda from the Ellis Bay Formation (Ordovician), Anticosti Island, Qubbec: Geol. Surv. Can., Paper 72-43. - _ _ - _ _ , 1974a. Silurian Ostracoda from Anticosti Island, Qubbcc: Geol. Surv. Can., Bull. 241. _ _ _ _ _ _ , 197413. Middle Ordovician Ostracoda from southwestern District of LIackenzie: Geol. Surv. Can., Bull. 244. _ _ _ - _ _ , 1976. Leperditicopid ostracodes as Silurian biostratigraphic indices: Geol. Surv. Can., Paper 76-IB, p. 83-88. Early Paleozoic Ostracoda from southwestern District of -. _ _ _ _ , in press. Mackenzie and Yukon Territory: Geol. Surv. Can., Bull. _ _ _ - - _ , in press. Ordovician Ostracoda from southeastern District of Franklin: i n , Bolton, T . E . , Sanford, B . V . , Copeland, ill. J. Barnes, C . R . and Rigby, J . K. , Gcology of Ordovician rocks, Melville Peninsula and region, southeastern District of Franklin: Geol. Surv. Can., Bull. 269. Dnwson, J . IV. , 1860. (1868, 1878, 1891). The geology of Nova Scotia, New Brunswick and Prince Edward Island or Acadian Geology: London. Dennison, J . M . and Head, J . W. , 1975. Sea level variations interpreted from the Appalachian Basin Silurian and Devonian: Am. J. Sci., v. 275, p. 1089-1120. Hall, J . , 1860. Description of new species of fossils from Silurian rocks of Nova Scotia; Can. Nat. Geol., v. 5, p. 144-159. Honeyman, D . , 1859. Abstract of a paper on the fossiliferous rocks of Arisaig: Trans. Nova Scotia Lit. and Sci. S O C . ,p. 19-29. _ _ _ _ _ _ , 1864. On the geology of Arisaig, Nova Scotia: Quart. J. Geol. SOC. London, v. 20, p. 333-345. Jones, T. R . , 1858. On the Palaeozoic bivalve Entomostraca of Canada: Geol. Surv. Can., Figures and descriptions of Canadian organic remains, Dec. 111, p. 91-102, pl. XI. -. .- - _ _ , 1870. Notes on some Entomostraca from Arisaig: i n , Honeyman, D . , Notes on the geology of Arisaig, Nova Scotia: Quart. 3 . Geol. SOC.London, v. 2 6 , p. 492. _ _ _ _ _ - , 1881a. Notes on some Palaeozoic Entomostraca: Nova Scotian Inst. Nat. S c i . , Proc. and T r a n s . , v. 5, n. 3, p. 313, 314. _ _ _ _ _ _ , 1881b. Notes on some Palaeozoic bivalved Entomostraca: Geol. RIag., dec. 2 , v . 8, p. 337-347. ___ , 1889. Notes on the Palaeozoic bivalved Entomostraca. No. XXVII. On some North-American (Canadian) species: Ann. Mag. Nat. Hist., ser. 6, n. 3, p. 373-387. _ _ _ _ , 1890a. On some Palaeozoic Ostracoda from North America, Wales and Ireland: Quart. J . Geol. SOC.London, v. 46, p. 1-31. - - - _ _ _ , 1890b. On some Devonian and Silurian Ostracoda from North America, France and the Bosphorus: Quart. J. Geol. SOC.London, v . 46, p. 534-536. -~
11 Jones, T. R . , 1891. On some Ostracoda from the Cambro-Silurian, S i l u r i a n , and Devonian rocks: ContT-ih. Can. hlicro-Pal. 111, Geol. Nat. Hist. S u r v . Canada, p. 59-99. Kay, G. R.1. , 1934. Rlohawkian Ostracoda: species common to Trenton faunules from the Hull and Decorah formations: J . Paleontol., v . 8, n . 3, p. 328-343. RlcLearn, F. €1. , 1 9 2 4 . Palaeontology of the Silurian r o c k s of Arisaig, Nova Scotia: Geol. S u r v . C a n . , 4Iem. 137. Twenhofel, W. H. , 1938. Geology and paleontology of the Rlingan Islands, Quebec: Geol. SOC. Amer. Sp. P a p e r s , n. 11, p. 65-67. Ulrich, E . 0. and B a s s l e r , R. S . , 1923. Ostracoda: in Rlaryland Geol. S u r v . , Silurian volume, p. 500-704. Whittington, H. B. and Kindle, C. H . , 1963. Middle Or,dovician Table Head Formation, western Newfoundland: Geol. SOC. Amer., Bull., v. 74, p. i45-758.
Explanation of Plates (GSC-Geological S u r v e y of Canada; USNM-United States National Rluseum) PLATE 1: (1-19 hliddle Ordovician, 20-27 Late Ordovician) I . Not-ochilina nora Copeland, x20, GSC 17056, 2. Oepikella labrosa Copeland, x10, GSC 17048d, 3. T e tr a d e ll a ulrichi Kay, x60, USNM 216123, 4. Ceratopsis quadrifida ( J o n e s ) , x20, USNRI 216115, 5. Scofieldia bilatcralis (Ulrich) , x10, GSC 18654, 6. Bassleratia typa Kay, x28, GSC 18657, 7. Bulticella s p . , x28, GSC 18655, 6. Euprimilia labiosa ( U l r i c h ) , x10, GSC 18652, 9. Dicranella marginata Ulrich, x16, GSC 17078, 10. Distobolbina teicherti Copeland, x50, GSC 41919, 1 1 . Skusloffina ulrichi Teichert, x20, USNM 216128, 12. Thoinasatia falcicosta Kay, x28, GSC 48245, 13. Krausella rawsoni Roy, x20, USNRI 2 1 6 1 2 9 , 14, 1 7 , T e t r a d e l l a buckensis Guber, x40, x50, GSC 41924, 41926, 1 5 . Levisulculus rnichiganensis Kesling, x60, USNhl 216120, 16. Glyrnmatobolbina? spinosa Copeland, x20, USNR? 216131, 16. .Monoceratella d e c o r a l a Copeland, x37, v c n t r a l view, USNM 216094, 19. Eurychilina s u b r a d i a t a Ulrich, x20, GSC 17035c, 20. Junesites setnilunatus ( J o n e s ) , x40, GSC 31568, 21. Euprimitia gamachei Copclnnd, x43, GSC 31556, 22. Foramenella phippsi Copeland, x43, GSC 31526, 23. T e t r a d e l l a anticostiensis Copeland, x50, GSC 31554, 24. Platybolbina shaleri Copeland, x43, GSC 31538, 25. Anticostiella e l l i s r n s i s Copeland, x43, GSC 3 1 5 2 9 , 26. Foramenella phippsi Copeland, x30, GSC 31411, 27. T e t r a d e l l a thomasi Copeland, x43, GSC 31539.
12 PLATE 2: (1-4, 7-29 Silurian, Anticosti Island, 5, 6 Devonian) 1. Conbathella inornata Copeland, x30, GSC 32470, 2. Conbathella biporata Copeland, x32, GSC 32518, 3. Conbathella equilateralis Copeland, x32, GSC 32504, 4. Ulrichia (Ulrichia) verticalis Copeland, x33, GSC 32922, 5. Venzavella s p . , x37, GSC 48246, 6. Kirkbyella ( B e r d a n e l l a ) obliqua Coryell and Cuskley, x50, GSC 48247, 7, 8. Anticostiella pustulosa Copeland, x33, GSC 32564, 32565, 9. Eustephanella? jupiterensis Copeland, x33, GSC 66763, 10. Punctobeecherella punctata Copeland, x33, GSC 33025, 11. Zygobolba twenhofeli Ulrich and Bassler, x l i , GSC 33087, 12, 23. Zygobolba dccora Ulrich and Bassler, x15 and x20, GSC 34698, 33126, 13, 19. Bolbineossia ( B r e v i b o l b i n e o s s i a ) berdanae Copeland, x16. 5 and x15, GSC 3 2 9 4 4 , 32951, 14. Bolbibollia papillosa Copeland, x33, GSC 33057, 1 5 , Bolbibollia labrosa Ulrich and B a s s l e r , x24, GSC 33043, 16. Zygobolba robusta Illrich and B a s s l c r , x17, GSC 33117, 17, 18. Zygobursa praecursor Copeland, x25 and x26, GSC 24389, 24390, 20, 24, 29. Apatobolbina whiteavesi Copeland, x33, x16 and x33, GSC 32952, 32954, 32960, 21. Craspedobolbina (Mitrobeyrichia) bolloni Copeland, x22, GSC 33012, 22. Zygobolba anticosliensis Ulrich and B a s s l e r , x15. GSC 33123, 25, 27. Bolbineossia (Bolbineossia) pineaulti Copeland, x30, GSC 32398, 32392, 26, 28. Anticostibolbina jupiterensis Copeland, x30, GSC 32415, 32416. PLATE 3: ( l - 4 , 6-9 Silurian, Arisaig, 5 Silurian, Pinto, Rlaryland, 1 1 , 15 Silurian, Lakc Alatapedia, 19-23 Devonian, Dalhousie, 10, 12-14, 16-18 Devonian, Forillon Peninsula) 1 . 2. .\facrypsilon salterianum ( J o n e s ) , x16, GSC 14566, a, 3. Kloedenia wilkensiana ( J o n e s ) , x13, GSC 1 4 5 1 3 , 4. Beyrichia ( N o d i b e y r i c h i a ) pustulosa Hall, x9, GSC 14503, 5. Pintopsis tricornis (Ulrich and B a s s l e r ) , x16, USNM 142251, 6. Hemsiella rnaccoyiana mclearni Copeland, x15, GSC 14561, 7, 8. Londinia arisaigensis Copeland, x10, GSC 14562, 14563, 9. Hemsiella maccoyiana sulcata ( R e u t e r ) , x16, GSC 14511, 10. Kloedeniopsis s p . , x20, GSC 48248, 1 1 . Garniella concentrica Berdan, x40, GSC 48249, 1 2 . Dizygopleura s p . , x40, GSC 48250, 13. T h l i p s u r o p s i s inaequalis (Ulrich and B a s s l e r ) , x50, GSC 48251, 14. Kloedenella s p . , x30, GSC 48252. 15. Lciocyamus s p . , x40, G S C 48253, 16. Janusella biceratina Roth, x20, G S C 4 8 2 5 4 , 17. Neothlipsura s p . , x52, GSC 48255, 18. Eucraterellina sp. cf. E . oblong0 (Ulrich and Bassler) , x55, GSC 48256, 19. Mesomphalus magnificus Copeland, x15, GSC 14537j, 20. Kloedeniopsis retifera (Ulrich and B a s s l e r ) , x15, GSC 14540d. 21. Ariklocdenia n e w b r u n s w i c k e n s i s (Copeland), x13. 5, GSC 14541d, 22. T h l i p s u r a cf. T . v - s c r i p t a (Jones and Holl), x30, GSC 14518, 23. S t w p u l i t e s dalhousiensis Copeland, x30, GSC 14523.
PLATE I
13
PLATE 2
PLATE 3
15
16
Discussion D r . B. K. H o l d s w o r t h : To what e x t e n t i s t h e a p p a r e n t p r o v i n c i a l sm o f Ordov i c i a n o s t r a c o d e s perhaps a f u n c t i o n o f environment and f a c i e s r a t h e r t h a n one o f geographic s e p a r a t i o n ? D r . M. J . Copeland: N o r t h American M i d d l e O r d o v i c i a n o s t r a c o d e s a r e q u i t e d i f f e r e n t f r o m t h e i r European c o u n t e r p a r t s . T h i s i s l e s s o b v i us i n L a t e O r d o v i c i a n faunas, a t l e a s t t h o s e I have s t u d i e d i n e a s t e r n and A r c t i c Canada. Undoubtedly environment and f a c i e s have a g r e a t e r a f f e c t on some faunal groups t h a n on o t h e r s . T h i s i s c e r t a i n l y e v i d e n t i n those groups t h a t depend on a p e l a g i c s t a g e o f t h e i r d i s t r i b u t i o n . The l a r g e , heavy S i l u r o - D e v o n i a n b e y r i c h i a c e a n s o f e a s t e r n N o r t h America on which Jean Berdan and I base most o f o u r concept of p r o v i n c i a l i s m by geographic separ a t i o n were, however, b e n t h o n i c , s h a l l o w w a t e r f a c i e s d w e l l e r s . T h e i r c r u m i n a l t y p e o f dimorphism c o u l d have been a response t o t h i s h i g h energy t y p e o f environment and p r o v i d e d p r o t e c t i o n f o r b o t h t h e eggs and j u v e n i l e s , p r e c l u d i n g a p e l a g i c s t a g e i n t h e i r development. D i s t r i b u t i o n o f such forms were t h e r e f o r e l i m i t e d t o m i g r a t i o n t h r o u g h s h a l l o w w a t e r env i r o n m e n t s . The appearance o f b e y r i c h i i d s a t a b o u t t h e same t i m e i n b o t h t h e N o r t h American and European assemblages c a n n o t have been c o i n c i d e n t a l . There must have been an i n i t i a l , s h a l l o w w a t e r c o n n e c t i o n between t h e cont i n e n t s i n t h e L a t e O r d o v i c i a n o r E a r l y S i l u r i a n a c r o s s which t h e p r i m i t i v e b e y r i c h i a c e a n s t o c k , whatever i t was, m i g r a t e d and, we t h i n k , t h e subsequent development o f a m a r i n e b a r r i e r such as a t r o u g h t h a t e f f e c t i v e l y d i v i d e d t h e fauna and p e r m i t t e d independent development o f N o r t h American and European assemblages i n s h a l l o w w a t e r s on e i t h e r s i d e . Dr. Jean Berdan: The b e y r i c h i a c e a n o s t r a c o d e s o f t h e Hardwood Mountain Form a t i o n o f w e s t e r n Maine a r e o f A p p a l a c h i a n a f f i n i t i e s and r e p r e s e n t , acc o r d i n g t o A. J . Boucot, an environment o f s h a l l o w w a t e r d e p o s i t i o n poss i b l y around i s l a n d a r c s . The b e y r i c h i a c e a n o s t r a c o d e s o f t h e E a s t p o r t area i n e a s t e r n Maine a r e e n t i r e l y d i f f e r e n t ; t h e y a r e r e l a t e d t o those o f t h e European B e y r i c h i e n k a l k b u t a l s o r e p r e s e n t a v e r y s h a l l o w w a t e r e n v i ronment a s s o c i a t e d w i t h v o l c a n i c s . Both t y p e s o f o s t r a c o d e s a l s o o c c u r i n p l a t f o r m environments and t h e d i f f e r e n c e s between them a r e t h e r e f o r e considered p r o v i n c i a l . Dr. J . E. Conkin: What i s t h e age o f t h e P r i d o l i a n beds and t h e i r presumed e q u i v a l e n t , t h e Downtonian, on t h e b a s i s o f o s t r a c o d e s ? Copeland: I t h i n k most o s t r a c o d e workers would c o n s i d e r them t o be L a t e S i 1u r i a n . D r . I . G. Sohn: Murray, you have Kirkbyacea? i n one o f y o u r s l i d e s . What genus was t h a t ? Copeland: A d o u b t f u l Roundyella? sp. f r o m n e a r t h e base o f t h e C h i c o t t e Formation o f A n t i c o s t i I s l a n d . T h i s i s a t b e s t a q u e s t i o n a b l e d e t e r m i n a t i o n based on r e l a t i v e l y few specimens. One comment c o n c e r n i n g o s t r a c o d e s f r o m t h e Lower M i s s i s s i p p i a n Banff Formation. Green (1963, Research C o u n c i l o f A l b e r t a , B u l l . 11) f o u n d t h a t many o f t h e s i l i c i f i e d o s t r a c o d e s o b t a i n e d by e t c h i n g i n d i l u t e h y d r o c h l o r i c a c i d d i f f e r e d f r o m t h o s e o f t h e n o n - s i l i c i f i e d fauna. I f y o u ' r e worki n g i n Nevada on t h e s i l i c i f i e d fauna, I s t r o n g l y suggest y o u d o n ' t e t c h everythinq. Sohn: We have known t h a t f o r years.(Sohn, 1950, Geol. SOC. America B u l l . , v. 61, p. 1504). T h i s i s something we d i s c u s s e d a t t h e Pau o s t r a c o d e conv e n t i o n . The s i l i c i f i e d fauna t h a t J o r d a n has i n Germany, Jean Berdan has from t h e Devonian and I have f r o m t h e M i s s i s s i p p i a n seem t o have a g r e a t d e a l i n common, namely, a l l t h e ornaments and s p i n e s a r e preserved, which you do n o t o b t a i n when you b e a t them o u t o f t h e o t h e r r o c k s . Copeland: I w a s n ' t t h i n k i n g o f t h e m o r p h o l o g i c a l aspect, I was t h i n k i n g o f t h e taxonomic. I n l i m e s t o n e o f t h e B a n f f Formation, and I e x p e c t i n many o t h e r s i m i l a r r o c k u n i t s , s p e c i e s o f o s t r a c o d e s a r e p r e s e r v e d e i t h e r as
17
calcareous o r s i l i c e o u s remains, b u t , u s u a l l y n o t b o t h . A l s o , numerous specimens o f o t h e r f o s s i l groups such as t r i l o b i t e s and b r a c h i o p o d s may s u r v i v e a c i d d i s i n t e g r a t i o n b u t t h e o s t r a c o d e s do n o t , even though t h e y w e r e v i s i b l e i n hand specimens. Why t h e r e i s t h i s p r e f e r e n t i a l s i l i c i f i c a t i o n I d o n ' t know. l t m u s t be due t o t h e o r i g i n a l c o m p o s i t i o n and s t r u c t u r e o f t h e s h e l l combined w i t h t h e t y p e o f a c i d t h a t i s used. I n some s t u d i e s , t o a r r i v e a t a complete l i s t o f t h e c o n t a i n e d o s t r a c o d e s , chemical and p h y s i c a l . you have t o combine b o t h methods
-
This Page Intentionally Left Blank
19
EARLY PALEOZOIC OSTRACODA OF THE ATLANTIC MARGIN, OTHER THAN EASTERN C A N A D A by F. M. Swain University of D e l a w a r e , Newark, D e l a w a r e 19711; a n d University of Minnesota, Minneapolis, Minnesota 55455 Abstract C a m b r i a n Archaeocopida and Ordovician a n d Silurian o s t r a c o d e s a r e b e t t e r developed in t h e m a r g i n a l a r e a s of t h e N o r t h A t l a n t i c t h a n those of t h e South A t l a n t i c w h e r e r o c k s of t h o s e a g e s a r e only s p a r s e l y repre sen ted. The m o s t useful a s s e m b l a g e s b i o s t r a t i g r a p h i c a l l y a r e t h o s e of t h e middle Ordovician a n d middle Silurian of b o t h t h e e a s t e r n a n d w e s t e r n North Atlantic.
Les A r c h a e o c o p i d a c a m briens et l e s o s t r a c o d e s o r d o v i c i e n s e t siluriens s o n t m i e u x &olu&s d a n s les rfgions d e bord d e I'Atlantique du Nord q u e d a n s 1'Atlantique du Sud, o u l e s r o c h e r s d e ces k r e s n e s o n t que peu reprdsentes. L e s a s s e m b l a g e s l e s plus u t i l e s b i o s t r a t i g r a p h i q u e m e n t s o n t c e u x de mi-ordovicien et mi-silurien d e s rggions est et o u e s t d e I ' A t l a n t i q u e du Nord.
Introduction I m p o r t a n t e a r l y c o n t r i b u t i o n s to knowledge of e a r l y P a l e o z o i c O s t r a c o d a w e r e m a d e in Europe by J o n e s a n d Holl (18691, B a r r a n d e (18721, K r a u s e (18911, R i c h t e r (1869) a n d G u r i c h (1896) and in t h e e a s t e r n United S t a t e s by Ulrich (1890, I89l>, a n d Whitfield (1890) to m e n t i o n a, few. During t h e e a r l y d e c a d e s of t h e t w e n t i e t h c e n t u r y major European w o r k s included t h o s e by Bonnema (1909) a n d K u m m e r o w (1924). In t h e United States s i g n i f i c a n t works included t h o s e of Ulrich and Bassler (1908, 1913a, 1913b, 19231, S w a r t z (1932, 1933, 19361, a n d K a y (1934, 1940). In Europe m a j o r p a p e r s on Ordovician and Silurian o s t r a c o d e s in Scandinavia a n d t h e B a l t i c region w e r e published by Hessland (19491, Henningsmoen (1953a, 1953b,' 1954a, 1954b, 1 9 5 4 ~ )J a a n u s s o n (1957) a n d Sarv (1959). In t h e U n i t e d S t a t e s p a p e r s d e a l i n g w i t h Ordovician a n d
20
Silurian o s t r a c o d e s include: S w a r t z a n d W h i t m o r e (1956); Swain (1953, 1957, 1962). In s u b s e q u e n t y e a r s m a n y p a p e r s h a v e a p p e a r e d o n European Ordovician a n d Silurian o s t r a c o d e s as discussed below, b u t only a f e w in t h e U n i t e d S t a t e s . For t h e l i t e r a t u r e on C a n a d i a n f a u n a s see Copeland (this volume). This a r t i c l e will briefly r e v i e w t h e major e a r l y P a l e o z o i c o s t r a c o d e f a u n a s of t h e A t l a n t i c margins o t h e r t h a n t h o s e of C a n a d a which a r e d e a l t w i t h by Dr. Murray Copeland, herein. Cambrian Archaeocopids o c c u r t h r o u g h o u t t h e C a m b r i a n S y s t e m in t h e United Kingdom; In t h e e a r l y C a m b r i a n , A l u t a , Beyrichiona, Bradoria, Dielymella, a n d Indiana a r e represented-bold a n d Pocock, 19 34; Cobbold, 1936). T h e middle C a m b r i a n contains, in a d d i t i o n , Entomidiella?, Hipponicharion, a n d S v e a l u t a (Cobbold a n d Pocock, 1934; Jones, 1856; Taylor a n d R u s h t o n X T h e l a t e C a m b r i a n h a s yielded C y c l o t r o n , F a l i t e s , Hesslandona, a n d f u r t h e r to t h e preceding: Vestro o t h i a (A. W. A. Rushton, w r i t t e n c o m m u n i c a t i o n , 6 F e b r u a r y
+
E l s e w h e r e in t h e A t l a n t i c borderlands, a r c h a e o c o p i d s a r e poorly known. A f e w o c c u r in t h e e a r l y a n d middle C a m b r i a n of m a r i t i m e C a n a d a (Copeland t h i s volume) a n d New York (Ulrich a n d Bassler, 1931). Ordovician The distribution of Ordovician O s t r a c o d a a r o u n d t h e "ProtoAtlantic" O c e a n is shown in F i g u r e 1. The c o n t i n e n t a l r e c o n s t r u c t i o n i s f r o m Smith, Briden, a n d D r e w e r y (1973). Early Ordovician Early Ordovician C a n a d i a n (Trernadocian? + Arenigian) o s t r a c o d e s a r e poorly known f r o m e a s t e r n N o r t h A m e r i c a , b u t isochilinids such as I.' g r e g a r i a (Whitfield) (Table 1, fig. 2-11) Isochilina s e e l y i I, fig. ( W h i t f i e l a (Table I, fig. 2-12) a n d I. c r i s t a t a (Whitfie1-e 2-10) and e o l e p e r d i t i i d s a r e r e p r e s e n t e d in c a r b o n a t e f a c i e s (Whit field, 1890, Bassler -and K e l l e t t , 1934). Undescribed leperditellids a n d o t h e r s m a l l e r s p e c i e s o c c u r in t h e c a l c i t i c facies (Swain, 19571, b u t widespread e a r l y Ordovician d o l o m i t e s a r e b a r r e n of o s t r a c o d e s . Isochilinids also o c c u r in t h e Arenigian of n o r t h e r n G r e e n l a n d (Poulsen 1929, 1934).
21
In Sweden (Hessland, 19491, a diverse early Arenigian ostracode assemblage includes Conchoides minuta Hessland (Table I, fig. 2-5) and related species, Glossomorphites tenuilimbata (Hessland) (Table I), and related forms, Primitiella brevisulcata Hessland (Table I, fig. 2-17), Aulaco sis spp. (Table I, gifProtallinella rewin ki (Bock) Table I, fig. 2-18), and others. The Arenigian of Norway Asaphus Series) has yielded a variety of beyrichiacean ostracodes in a mainly shaly facies (Henningsmoen, 1954a). In Wales and central England entomidellids, beyrichiids, and tetradellids have been recorded from Arenigian beds (Table I, fig.2). Additional early Ordovician ostracodes not mentioned above a r e listed in Table 1 and illustrated in figure 1 (Henningsmoen, 1954a; Ulrich and Bassler, 1908; Jones, 1884). An important early Ordovician ostracode fauna has been described from the Estonian S.S.R. (Sarv, 1959)
T---e
g__pf
22
Figure 2 1. 2.
3. 4. 5.
6. 7.
8. 9. 10.
11. 12.
13. 14. 15.
16. 17.
18. 19. 20.
Aulacopsis b i f i s s u r a t a Hessland. R i g h t valve, x23, e a r l y Ordovician,’ Leskusanget, Sweden (Hessland, 1949). Aulacopsis m o n o f i s s u r a t a Hessland. R i g h t valve, x23, e a r l y Ordovician, S t e n b e r g , Sweden (Hessland, 1949). Beyrichia b a r r a n d i a n a Jones. L e f t valve, x3, e a r l y Ordovician, Mynyddgarw, Wales ( J o n e s , 1885). C e r a t o c y p r i s longispina Hessland. R i g h t valve, x23, e a r l y Ordovician, L e s k u s a n g e t , Sweden (Hessland, 1949). Conchoides m i n u t a Hessland. R i g h t valve, x23, e a r l y Ordovician, S t e n b e r g , S w X c H e s s l a n d , 1949). C t e n e n t o m a p l a n a Hessland. R i g h t valve, x23, e a r l y Ordovician, Leskusanget, Sweden (Hessland, 1949). Entomidella m a r r i i Jones. R i g h t valve, x1.5, e a r l y Ordovician (distorted), N a n t l l e , Wales (Jones, 1884). Eurychilina d o r s o t u b e r c u l a t a Hessland. R i g h t valve, x23, e a r l y Ordovician, Silberberg, Sweden. (Hessland, 1949). Glossopsis d e p r e s s o l i m b a t a Hessland. R i g h t valve, x23, e a r l y Ordovician, Leskusanget, Sweden (Hessland, 1949). Isochilina c r i s t a t a (Whitfield). R i g h t v a l v e x8, e a r l y Ordovician, L a k e C h a m p l a i n , V e r m o n t (Swain, 1957). Isochilina g r e g a r i a (Whitfield). R i g h t valve, x6, e a r l y Ordovician, L a k e C h a m p l a i n , Vermont (Swain, 1957). Isochilina s e e l y i (Whitfield), L e f t valve, x6, e a r l y Ordovician, L a k e C h a m p l a i n , Vermont (Swain, 1957). Laccochilina d o r s o p l i c a t a Hessland. R i g h t valve, x23, e a r l y Ordovician, Silberberg, Sweden. Nanopsis n a n e l l a (Moberg and Segerberg). R i g h t valve, x38, e a r l y Ordovician, S l e m m e n s t a d , Norway (Henningsmoen, 1954). Ogmoopsis n o d u l i f e r a Hessland. R i g h t valve, x23, e a r l y Ordovic i a n , Silverberg, Sweden (Hessland, 1949). P i n n a t u l i t e s p r o c e r a (Kurnmerow). R i g h t valve, x23, e a r l y Ordovician, L e s k u s a n g e t , Sweden (Hessland, 1949). P r i m i t i e l l a b r e v i s u l c a t a Hessland. R i g h t valve, x23, e a r l y Ordovician, Leskusanget, Sweden, (Hessland, 1949). Protallinella grewingki (Bock). R i g h t valve, x23, e a r l y Ordovic i a n , S t e n b e r g , Sweden (Hessland, 1949). Steusloffia a c u t a (Krause). L e f t valve, x15, e a r l y Ordovician? d r i f t boulders, Mark Brandenberg, G e r m a n y (Ulrich a n d Bassler, 1908). Steusloffia polynodulifera Hessland. L e f t valve, x23, e a r l y Ordovician, Born-Dadran, Sweden (Hessland, 1949).
23
Middle Ordovician The o s t r a c o d e s of t h e C h a z y a n (Llanvirnian a n d Llandeilan) of t h e e a s t e r n United S t a t e s a r e i e p r e s e n t e d by a lower Bullatella kauff manensis Zone and a n upper M o n o c e r a t e l l a t e r e s Zone= figs. 3-6, 21,) Swain 1957, 1962). Many o t h e r leperditiids, a p a r c h i t i d s , opikellids, leperditellids, eurychilinids, budnianellids a n d p r i m i t i v e cypridids a r e also r e p r e s e n t e d (Swain 1957, 1962) s o m e of which a r e listed in Table 2 a n d i l l u s t r a t e d in f i g u r e 3. An e x t e n s i v e silicified Chazyan f a u n a is r e p r e s e n t e d in Virginia ( K r a f t , 1962) a n d in New York (Swain, 1962). Black R i v e r a n (early C a r a d o c i a n ) o s t r a c o d e s a r e poorly known i n t h e e a s t e r n United S t a t e s a l t h o u g h w e l l r e p r e s e n t e d in t h e c e n t r a l United S t a t e s (Kay 1934, 1940; Swain, Cornell, a n d Hansen, 1961). Trentonian (middle C a r a d o c i a n ) shaly l i m e s t o n e s of New York a r e
24
Table 1. Early Ordovician Ostracoda, Atlantic Margins E, Eastern Atlantic; W , Western Atlantic Canadian Species Beyrichiona triceps Nanopsis nanella Primitia? sp. Aulacopsis spp. Conchoides minuta Glossomorphites tenuilimbata Primitiella brevisulcata PTotallinella grewingki Ceratocypris longispina Eurychilina dorsotuberculata Glossopsis depressolimbata Laccochilina dor soplica ta Ogmoopsis nodulifera Pinnatuli tes procer a Steuslof f ia cf. polynodulif e r a Beyrichia barrandiana Conchopr im i t ia eos C t e n t o m a plana Glossmor phi tes sp. Isochilina cris ta ta Isochilina gregar ia Isochilina seelyi Rigidella e r r a t i c a Steusloffia a c u t a Tetradella? sp. Isochilina a r c t i c a
* GB, G r e a t Britian; Gr , Greenland
N, Norway, S , Sweden; US, United States;
exemplified by Bassleratia typica Kay (Table 2, fig.2-31, Bollia subaeauata Ulrich (Table 2. fie. 3-41. Primitiella constricta Ulrich ( T a b l e g . 3-27), Thomasatia T a l c i c o k a (Table 2, f m d other forms (Kay 1934, 1940; Swain et a m In limestone facies of c e n t r a l and southern Sweden, t h e Llanvirnian (Aseri Stage) contains among others, Piretia geniculata Jaanusson (Table 2, fig. 3-23) and Laccochilina (L.) bulbata Jaanusson (Table 2, fig. 3-19). The overlying early Llandeilzn (=ae Stage) is typified by Eu rimites effusus Jaanusson (Table 2, fig. 2-12) and Actinochilina (Jaanusson,l957). The succeeding Uhaku S t a g e spp. Table 2, 1)3-
+-
25
Table 2. Middle Ordovician Ostracoda of Atlantic Margins E, Eastern Atlantic; W , Western Atlantic Mohawkian Chazyan Species LOC. Llanv. Lland. Bullatella kauffmanensis US --W-Laccochilina bulbata S --E-S -- E-Piretia geniculata Budnianella shenandoense US -------W Elliptocyprites parallela US -------W Eogra p hiodac t ylus eos us -------W Eurychilina strasburgensis US -------W Krausella variata us -----__W--? Primitiella champlainensis US -------W Actinochilina spp. Balticella spp. Conchopr imites? spp. Euprimites effusus Eupr imi tes bur sell us Euprimites suecicus Hesperidella spp. Parapyxion spp. Primitiella spp. Pyxion sp. Sigmobolbina sigmoidea Steusloffia linnarssoni Tallinella dimorpha Tvarenella spp. Ulrichia? bipunctata Monoceratella t e r e s Euprimites locknensis Plstybolbina spp. Steusloffia c o s t a t a Tetradella complicata Tallinella trident Ullerella triplicata Ctenobolbina spp. Bolbina spp.
.
Tr B.R. Early Caradoc.
26
contains, among others Laccochilina (L.) auci ranosa Jaanusson (Table 2, fig. 3-20), Euprimites bursellus Jaanusson Table 2 , fig. 3-13), Tallinella dimor ha Jaanusson (-fig. 3-32), Steusloffia linnars- Jaanusson Table 2, fig. 3-31), and soni Jaanusson (Table 2, fig. 3-29). In t h e l a t e t w o facies assemblages occur: a carbonate facies, with species of (Table 2, fig. 3-36), Euprimites 2, fig. 3-17) and Balticella a mudstone f a c i e s with Conchoprimites? (Table 2, fig. 3-71, Actinochilina (Table 2, fig. 3-1) and Parapyxion (Table 2); both f a c i e s a r e represented by Euprimites locknensis Jaanusson (Table 2, fig. 3-14), and Steusloff i a costata Jaanusson (Table 2, fig. 3-20) (Jaanusson 1957). A large middle Ordovician fauna is present in t h e Kuckruse and other formations of Estonia (Bonnema, 1909; Sarv, 1959).
E_B__
-+
Figure 3 1. 2. 3.
4. 5. 6. 7. 8.
9. 10. 11.
12.
Actinochilina suecica (Thorslund). Right valve of holotype, x19, middle Ordovician, Ludibundus beds, Kinnekulle, Sweden (Jaanusson, 1957). Balticella deckeri (Harris). Right valve, x13, middle Ordovician, Strasburg Junction, Virginia (Kraf t, 1962). L e f t valve, holotype, x23, middle Bassleratia typa Kay. Ordovician, Northumberland County, Ontario (Kay, 1934). Bollia subae u a t a Ulrich. L e f t valve, x23, middle Ordovician, Church, Iowa Kay, 1940). Budnianella shenadoense Kraf t. Right valve, holotype, x20, middle Ordovician, Tumbling Run, Virginia (Kraf t, 1940). Bullatella kauff manensis (Swain). Right valve, x16, middle Ordovician, Marion, Pennsylvania (Swain, 1957). Conchoprimitia leperditioidea Thorslund. Right valve, x15, middle Ordovician, Siljan, Sweden (Jaanusson, 1957). Ctenobolbina minor kuckersiana (Bonnema). L e f t valve, xll, middle Ordovician, early Caradocian, Kukruse, Estonia (Bonnema, 1909). Ctenbolbina o r n a t a latimarginata (Bonnema). L e f t valve, xll, middle Ordovician, early Caradocian, Kukruse, Estonia (Bonnema, 1909). Elliptocyprites longula Swain. Right valve, holotype, x36, middle Ordovician, Lake Champlain, Vermont (Swain, 1962). Eographiodactylus eos K r a f t . Left valve, x26, middle Ordovician, Tumbling R u n T r g i n i a (Kraft, 1962). Euprimites efiusus Jaanusson. a, L e f t valve, of heteromorph, x19, b, l e f t valve of heteromorph, middle Ordovician, Oland Seby, Sweden (Jaanusson, 1957.)
-*
27
13.
14. 15. 16.
17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30.
31. 32.
Euprimites bursellus Jaanusson. L e f t v a l v e of h e t e r o m o r p h , x19, middle Ordovician, O s t e r g o t l a n d , Sweden (Jaanusson, 1957.) Euprimites locknensis (Thorslund). R i g h t v a l v e of h e t e r o m o r p h , x19, middle Ordovician, J a m tland, S w e d e n (Jaanusson, 1957). Euprimites s u e c i c u s (Thorslund). L e f t valve, of h e t e r o m o r p h , x19, middle Ordovician, J a m t l a n d , Sweden (Jaanusson, 1957). Eur‘ychilina s t r a s b u r g e n s i s K r a f t . L e f t v a l v e of f e m a l e , x13, middle Ordovician, S t r a s b u r g J u n c t i o n , Virginia ( K r a f t , 1962). Hesperidella e s t h o n i c a (Bonnema). R i g h t v a l v e of h e t e r o m o r p h , x28, middle Ordovician, Ludibundus L i m e s t o n e , T v a r e n a r e a , Sweden (Jaanusson, 1957). Krausella v a r i a t a K r a f t . R i g h t valve, x13, middle Ordovician, Strasburg J u n c t i o n , Virginia ( K r a f t , 1962). L a c c o c h a i n a ( L a c c o c h a i n a ) b u l b a t a Jaanusson. L e f t valve of tecnomorph, x19, middle Ordovician, O s t e r g o t l a n d , Sweden (Jaanusson, 1957). Laccochilina (Laccochilina) paucigranosa Jaanusson. L e f t v a l v e of h e t e r o m o r p h , x19, middle Ordovician, e r r a t i c boulder Erken No. 10, S o u t h Bothnia, Sweden (Jaanusson, 1957.) Monoceratella t e r e s T e i c h e r t . L e f t valve, x13, middle Ordovician, S t r a s b u r g J u n c t i o n , Virginia ( K r a f t , 1962). Parapyxion s i b o v a t u m (ThGslund). L e f t valve, x23, middle Ordovician Ludibundus beds, Vasterogotland, Sweden (Jaanusson, 1957). P i r e t i a g e n i c u l a t a Jaanusson. L e f t valve of h e t e r o m o r p h , x26, middle Ordovician, O s t e r g o t l a n d , Sweden (Jaanusson, 1957). P r i m i t i a s i m p l e x (Jones): R i g h t valve, x6, middle Ordovician, n e a r Coimbra. P o r t u g a l (Jones, 1855). P r i m i t i a ? bicornis (:ones). L e f t valve, x6, middle Ordovician, Shropshire, England (Jones, 1855). P r i m i t i e l l a c t a m p l a i n e n s i s Swain. R i g h t valve, x16, middle Ordovician, L a k e C h a m d a i n , V e r m o n t (Swain, 1962). Prirnitiella’ c o n s t r i c t a Ulrich. L e f t valve, x23, middle Ordovician, Cannon Falls, Minnesota (Kay, 1940). Pyxion c a r i n a t u m -(Hadding). R i g h t valve, x19, middle Ordovician, Scania, Sweden (Jaanusson, 1957). Sigmobolbina s i g m o i d e a Jaanusson. a, L e f t v a l v e of t e c n o m o r p h , x19; b, d i a g r a m m a t i c d r a w i n g of h e t e r o m o r p h of s p e c i e s , x30, middle Ordovician, Oland, Sweden (Jaanusson, 1957). R i g h t valve, xll, middle Steusloffia costata (Linnarsson). Ordovician, Ludibundus l i m e s t o n e , Siljan d i s t r i c t , Sweden (Jaanusson, 1957). Steusloffia- linnarsoni (Krause). L e f t valve, xll, middle Ordovician, Oland,-(Jaanusson, 1957). Tallinella d i m o r p h a Opik. R i g h t valve, xll, middle Ordovician, Siljan d i s t r i c t , Sweden (Jaanusson, 1957).
29
36
37
38
39
A f e w middle Ordovician ostracodes of primitiid and tetradellid types a r e known from France, Spain, and Portugal (Bassler and Kellett 1934). A g r e a t many species have also been described from drift boulders presumed t o be middle Ordovician in northern Germany (Kummerow, 1921). Ostracoda a r e s c a r c e in t h e United Kingdom prior to t h e Caradocian. Primitiids and tetradellids a r e common in t h e Caradocian and Ashgillian of Wales (Jones, 1855; Harper, 1947; A.W.A. Rushton, written communication, 6 February 1976). L a t e Llandeilan and early Caradocian beds in San Juan Province, Argentina contain Ctenobolbina?, Parenthatia, Balticella, and Anisochilina, all of which are reminiscent of Scandinavian and middle Appalachian Assemblages (Baldis and Rossi d e Garcia 1972). Figure 3. (Continued) 33. Tallinella trident Henningsmoen. L e f t valve, x16, middle Ordovician, Sandsvaer, Norway (Henningsmoen, 1953). 34. Tetradella complicata Salter. Right valve, x9, middle Ordovician, early Llandeilan, Pembrokeshire, WaledHarper, 1947). 35. Thomasatia falcicosta Kay. Right valve, x23, middle Ordovician, Cannon Falls, Minnesota (Kay, 1940). 36. Tvarenella c a r i n a t a (Thorslund). Right valve of tecnomorph, x19, middle Ordovician, Sodermanland, Sweden (Jaanusson, 1957). 37. Uhakiella coelodesma Opik. Right valve of male, x15, middle Ordovician, early Caradocian, Purtsejogi River, Estonia. 38. Ullerella triplicata Henningsmoen. L e f t valve, x13, middle Ordovician, Ringerike, Norway (Henningsmoen, 1953). 39. Ulrichia bipunctata (Jones and Holl). - Right valve, XU, middle Ordovician, Llandeilan, Brecknockshire, Wales (Jones and Holl, 1869).
30
T a b l e 3. L a t e O r d o v i c i a n O s t r a c o d a of A t l a n t i c Margins E, E a s t e r n A t l a n t i c ; W , W e s t e r n A t l a n t i c Eden-Mays Species Ceratopsis chambersi Ctenobolbina c i l i a t a Tetradella aff. quadrilirata Aparachites maccoyi Macronotella spp. Pla tybol bina cf p l a n a Primitia s a n c t b a t r i c i i
.
Rich-Gam
LOC.
us us us
GB N N GB
L a t e Ordovician T h e r e a r e s p a r s e late Ordovician Edinian, Maysvillian, a n d Richmondian ( L a t e C a r a d o c i a n a n d Ashgillian) o s t r a c o d e f a u n a s in c l a s t i c l i t h o f a c i e s of t h e e a s t e r n U n i t e d s t a t e s (Butts, 1940). A f e w s p e c i e s a r e l i s t e d in T a b l e 3 a n d i l l u s t r a t e d in f i g u r e 4, b u t in g e n e r a l t h e s e f a u n a s a r e poorly known.
Figure 4
1. 2.
3. 4.
5. 6. 7.
8.
A p a r c h i t e s m a c c o y i i J o n e s a n d Holl. R i g h t valve, x8?. l a t e Ordovician, C a r a d o c i a n (Bala), C h a i r of Kildare, Ireland ( J o n e s a n d Holl 1868). C e r a t o p s i s c h a m b e r s i (Miller). L e f t valve, i n t e r n a l mold, x9, l a t e ? Ordovician, Martinsburg Shale, R o a n o k e C o u n t y , Virginia. (Butts, 1941). C t e n b o l b i n a c i l i a t a (Emmons). L e f t valve, i n t e r n a l mold, x9, l a t e ? (Ordovician, M a r t i n s b u r g Shale, R o a n o k e C o u n t y , Virginia. (Butts, 1941) M a c r o n o t e l l a cf. M. r a e l o n a (Steusloff). R i g h t valve, x8, l a g e 0r d o v i c i a n , Ask e r y N o r w ay Henn i ngs m oe n , 195 4 ) Platybolbina cf. P. e l o n ata (Krause). L e f t valve, XU, late Ordovician, Oslo, Norway Henningsmoen, 1954). Platybolbina cfi P. l a n a ( K r G s e ) . . L e f t valve, x15, late Ordovician, Oslo, N2rway Henningsmoen, 1954). P r i m i t i a s a n c t i p a t r i c i i J o n e s a n d Holl. R i g h t valve, x15, late Ordovician, C a r a d o c i a n (Bala), C h a i r of Kildare, Ireland. T e t r a d e l l a -sp. aff. T. q u a d r i l i r a t a (Hall a n d Whitfield). R i g h t valve, x9, l a t e ? Ordovician, M a r t i n s b u r g Shale, R o a n o k e C o u n t y , Virginia. (Butts, 1941).
e_Tg
-+5
.
31
A hiatus occurs between middle Ordovician and l a t e Ordovician ostracode faunas in Norway, t h e Tretaspis beds being nearly barren (Henningsmoen, 1954b). The overlying Dalmanitina beds (Ashgillian) contain a varied fauna cf. P. plana Krause (Table 3, fig. 4-61, P. (Ta3le 3, fig. 4-51, Macronotella spp. (TaEle Ashgillian deposits of t h e United Kingdom have yielded a dozen or more species including Aparchites (Salter) (Table 3, fig. 4-11 and Primitia sanctipatricii Salter fig. 4-7). The l a t e Ordovician of Estonian S.S.R. has a maior ostracode fauna (Sarv, 1959). Other regions bordering t h e Atlantic'have f e w or no recorded l a t e Ordovician ostracodes. Silurian The distribution of Silurian Ostracoda around t h e "ProtoAtlantic" Ocean plotted on t h e map for t h e Lower Devonian of Smith, Briden, and Drewery (1973) is shown in Figure 5. Early Llandoverian (Medinan) ostracodes a r e poorly developed in eastern United S t a t e s owing to unfavorable sandy facies but a f e w leperditiids occur, i.e. Leperditia cylindrica Hall (Bassler and Kellett, 1934). L a t e Llandoverian C -C beds (Clinton Group) of t h e Applachian region United S t a t e 3 ape abundantly supplied with ostracodes and have been divided into t h e following zygobolbid zones in
32
Figure 5. Map of Lower Devonian, S-pole stereographic projection, a f t e r Smith, Briden, and Drewery (1973), showing Silurian ostracode distribution about t h e "Protoatlantic" Ocean. Circles and dots show interpretations of south polar positions as determined from various places, with size of circle proportional to degree of uncertainty. t h e Rose Hill Shale and other Clinton equivalents. The succeeding Wenlockian beds (Rochester Shale) contain another zone, t h e Drepanellina clarki(Tab1e 4, fig. 6-11). Ludlovian beds (McKenzie and Wills Creek Formations) have abundant small kloedenellids such as: Dizygopleura acuminata Ulrich and Bassler (Table 4, fig. 6-10), Eukloedenella umbonata Ulrich and Bassler (Table 4, fig. 6-12), and Kloedenella scapha Ulrich and Bassler (Table 4, fig 6-15) PAdolian shaly limestones have large Kloedenia (Table 4, fig. 6-16), Pintopsis (Table 41, Welleria (Table 4, f i g m e l l e r i o p s i s (Table 4, fig 6-32), and Lophokloedenia (Table 4, fig. 6-18), which a r e indigenous to t h e American Appalachians at t h e species level but have a number of European affinities (Berdan 1962, 1970, 1971a; Martinsson 1970; Swartz and Whitmore, 1956).
33
Table 4a. E a r l y Silurian O s t r a c o d a of A t l a n t i c Margins E, E a s t e r n A t l a n t i c ; W , W e s t e r n A t l a n t i c
Species Leperdi t i a cylindr i c a Zygobolba e r e c t a Zygobolba a n t i c o s t i e n s i s Zygobolba d e c o r a Zygobolbina e m a c i a t a Mastigobolbina l a t a Zygosella post i c a Bonnemaia rudis Mastigobolbina t y p u s P a r a e c h m i n a spinosa Arcuaria sineclivula Beyrichia p a u c i t u b e r c u l a t a Craspedobol bina a r m a ta Neobeyrichia r e g n a n s Nodibeyrichia scissa Platybolbina? bulbosa Primitiella? p a r a l l e l a
LOC.
us us us us us us us us us us
B N? N? B? GB,B N? N?
Medinian Early Niagaran Llandover ian Early Late
--w-?-w-? ?-W-? ?-w-?
?-w-?
?-W-? ?-W-? ?-W-? ? - W? ?-W ? E - - - - - - -? ? _ - - - - - - - -E - - - - - - -? ?---------E------? ? _ - - - - - - - - E - - - - - - -? ? - - - - - - - - - E - - - - - - -? ? - - - - - - - - - E - - - - - - -? ? - - - - - - - - - E - - - - - - -?
_________
T a b l e 4b. Middle Silurian O s t r a c o d a of A t l a n t i c Margins E, E a s t e r n A t l a n t i c ; W, W e s t e r n A t l a n t i c
SDecies Apatobolbina p l a t y g a s t e r Beyrichia kloedeni Craspedobolbina v a r i o l a t a Drepanellina c 1a r k i Microcheilinella variolaris Be yr ic hia ringer ikensis Craspedobolbina p e r c u r r e n s Dizygopleura a c u m i n a t a Entomozoe? marstoniana Eukloedenella u m b o n a t a Hemsiella a n t e r o v e l a t a Hemsiella m a c c o y a n a Kloedenella s c a p h a Neobeyrichia n u t a n s Primitiopsis planif r o n s Signetopsis s e m i c i r c u l a r i s Macrypsilon s a l t e r i a n u m Welleria obliqua
LOC. N GB GB,N
us B N B
us GB us B GB
us
B N N GB
us
34
Table 4c. L a t e Silurian Ostracode of Atlantic Margins E, Eastern Atlantic; W , Western Atlantic Pridolian (Downtonian) Species LOC. Middle and L a t e Cayugan B ? _ - - -E _ - - - ? Am phitoxis cur va ta Frost iella groenvalliana US,GB,N Hemsiella aff. maccoyana US,B Kloedenia crassipunctata us Le perdi t ia scalar is us Londinia arisaigensis US,GB Londinia aff. arisaigensis 5,US Londinia kiesowi B Lophokloedenia manliensis us Macrypsilon aff. salterianum us Nodibe yr ichia tubercula ta B Pintopsis spp. US,B Richteria? sp. us Sleia equestris G B, B, US W eller ioDsis iersevensis us ~~
++ +*
In Norwav (Oslo region) t h e Llandoverian is represented by Be richia- (Eobi richia) sfi. (Table 4, figs. 6-4), Table 4, figs. 6-26 Primitiella? spp. (Table moen, 1 9 5 4 ~ ) . The Wenlockian contains f e w ostracodes, among t h e m A atobolbina platygaster Kummerow (Table 4, fig. 6-2) (Hennings.c)The Ludlovian contains abundant Primitio sis spp. (Table 4, fig. 6-28), Si net0 sis spp. (Table 4, and Neobeyrichia rin erikensis Henningsmoen) (Table 4, fig. 6-5) (HenningThe Ludlovian of northwestern Europe is generally smoen, 1954c characterized by "amphitoxidine" ostracodes: Londinia sp. (Table 4, fig. 6-21), Lo hoctenella spp. (Table 41, and S l e i a m b l e 4, fig. 6-30) fauna occurs in l a t e Wenlockian? and Ludlovian (Martinsson 1962 beds (Berdan, 1967, 1970, 1971; Martinsson, 1970). The l a t e r Ludlovian (Leintwardinian and Whitcliffian Stages) in northern England comprises zones based on Nodibeyrichia (Table 4, fig. 6-24), Neobe richia (Table 4, fig. 6-23), Cyptopholobus, Juviella, and Hemsiella Table 4, fig. 6-14] (Shaw, 1971). The succeeding D o w n t o w n i a n m a o7 l i a n ) is characterized by t h e kloedenid Frostiella groenvalliana Martinsson (Table 4, fig. 6-13) (Shaw, 1971). A similar fauna occurs in New York, Maine and Nova Scotia (Martinsson, 1970; Berdan, 1970, 1972).
6
_E_T--
-?-
An interesting undifferentiated Silurian ostracode assemblage characterized by entomids was long ago recorded from Sardinia (Canavari, 1899, 1900). The entomids a r e generally believed to b e pelagic ostracodes (Rozhdestvenskaja, 1971) and although t h e family is known from t h e Ordovician, t h e Sardinian Silurian entomids seem to represent one ot t h e earliest pelagic assemblages.
35
Elsewhere in t h e A t l a n t i c basin, e x c e p t in e a s t e r n C a n a d a which is discussed herein by Dr. Copeland, Silurian o s t r a c o d e s a r e only slightly known; a f e w f o r m s h a v e been described f r o m F r a n c e (Tromelin and Lebesconte 1876) and f r o m Brazil (Clarke, 1869) (Table
4). Figure 6 1.
2. 3.
4. 5. 6. 7.
8.
9. 10.
11.
12. 13. 14.
15. 16. 17.
Amphitoxis c u r v a t a Martinsson. L e f t valve, f e m a l e , x45, Pridolian, Snoder Gotland (Martinsson, 1962). Apatobolbina p l a t y g a s t e r (Kummerow). L e f t valve, xll, Wenlockian, Ringerike, Norway (Henningsmoen, 1954). Beyrichia (Beyrichia) cf. B. kloedeni M'Coy). L e f t valve, x23, Wen I ock i a n , S h ro p shi r e , E "21 a w t in sso n , 19 6 2). Beyrichia (Eobeyrichia) p a u c i t u b e r c u l a t a Henningsmoen. L e f t valve, x14, Llandoverian, Gunneklev, Norway (Henningsmoen, 1954). Beyrichia (Neobeyrichia) ringerikensis Henningsmoen. Left valve, x7, Ludlovian, Oslo, Norway (Henningsmoen, 1954). Bonnemaia rudis Ulrich and Bassler. Right valve, x6, Niagaran, Powell Mountain, Tennessee (Ulrich and Bassler, 1923). Craspedobolbina a r m a t a Heningsmoen. L e f t valve, xll, Llandoverian, Oslo, N o r w m e n n i n g s m o e n , 1950). Craspedobolbina (Mitrobeyrichia) percurrens M a r t insson. Left valve of f e m a l e , x30, Ludlovian, Follingbo, Sweden (Martinsson, 1962). Craspedobolbina (Mitrobeyrichia) v a r i o l a t a Martinsson. Left valve of f e m a l e , x30, Wenlockian, Djupvik, Sweden, (Martinsson, 1962). Dizygopleura a c u m i n a t a Ulrich a n d Bassler. L e f t valve, x12, l a t e Niagaran, Cumberland, Maryland. (Ulrich and Bassler, 1923). Drepanellina c l a r k e i Ulrich and Bassler. L e f t valve of m a l e , x9, Niagaran, Cumberland, Maryland. (Ulrich and Bassler, 1923). Eukleodenella u m b o n a t a Ulrich and Bassler. L e f t valve, e x f o l i a t e d posteriorly, xl2, l a t e Niagaran, Flintstone, Maryland, (Ulrich and Bassler, 1923). Frostiella groenviliiana Martinsson. L e f t valve of f e m a l e , x23, Downtonian-Pribolian, Scania, Sweden (Mart insson, 1962). Hemsiella maccoyana. (Jonesj. L e f t valve of h e t e r o m o r p h , x30, Ludlovian, Kirkby Moor Flags, England (Shaw, 1971). Kloedenella s c a p h a Ulrich - a n d Bassler. L e f t valve, xl5, L a t e Niagaran, G r e a t Cacapon, Maryland (Ulrich and Bassler, 1923). Kloedenia c r a s s i p u n c t a t a - S w a i t z a n d Whitmore. L e f t valve of f e m a l e , X15, Cayugan, Austin's Glen, New York (Swartz a n d Whitmore, 1956). Leperditia scalaris p r a e c e d e n s Ulrich a n d Bassler. L e f t valve,
36
Figure 6 (continued) 18.
19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37.
x5, Cayugan, Pinto, Maryland (Ulrich and Bassler, 1923). Lophokloedenia manliensis (Weller). L e f t valve of male, x15, Cayugan, Tristate, New York (Swartz and Whitmore, 1956). Mastigobolbina la t a (Hall). Right valve of male, x6, Niagaran, New Hartford, New York (Ulrich and Bassler, 1923). Mastigobolbina typus Ulrich and Bassler. Right valve of female, x5, Niagaran, Six Mile House, Maryland (Ulrich and Bassler, 1923). Londinia kiesowi (Krause). Right valve of female, x23, Pridolian, e r r a t i c boulder, c e n t r a l Baltic a r e a (Martinsson, 1962). Macrypsilon sa'lterianum (Jones). Right valve, x30, Ludlovian, Kirkby Moor Flags, England (Shaw, 1971). Neobeyrichia (NeobeGichia) nutans (Kiesow). L e f t valve of female, x23, Ludlovian, Ham marudden, Sweden (Mart insson, 1962). Nodibeyrichia scissa Martinsson. L e f t valve of female, x23, Llandoverian, Hulte, Sweden (Martinsson, 1962). Paraechmina- s inosa (Hall). L e f t valve, x15, Niagaran, Rochest e r . New York Ulrich and Bassler. 19231, Platybolbina? bulbosa Henningsm'oen. L e f t valve, xll, Llandoverian, Oslo, N a H e n n i n g s m o e n , 1954). Primitiella? cf. P. parallels Kummerow. L e f t valve, xll, Llandoverian, G u n n a l e v , Norway (Henningsmoen, 1954). Primitiopsis cf. P. lanifrons Jones. L e f t valve, xll, Ludlovian, Oslo region, Norway Henningsmoen, 1954). Si net0 sis semicircularis (Krause). Right valve, x17, Ringerike, Norway, Henningsmoen, 19 54). Sleia e uestris Martinsson. L e f t valve of female, x34, Pridolian, Sles, Gotland Martinsson, 1962). Welleria obliqua Ulrich and Bassler. Right valve of female, x9, Cayugan, Hancock, Maryland (Ulrich and Bassler, 1923). Welleriopsis jerseyensis (Weller). Holotype l e f t valve of male, x14, Cayugan, Nearpass Quarries, New Jersey (Swartz and Whitmore 1956). Zygobolba anticostiensis Ulrich and Bassler. Right valve of male, x6, Niagaran, Anticosti Island, Quebec (Ulrich and Bassler, 1923). Zygobolba decora (Billings). Right valve of female, x23, Niagaran, Anticosti Island, Quebec (Martinsson, 1962). Zygobolba e r e c t a Ulrich and Bassler. L e f t valve of male, x6, Niagaran, Cherrytown, Pennsylvania (Ulrich and Bassler, 1923). Zygobolbina e m a c i a t a Ulrich and Bassler. Right valve of female, x6, Niagaran, Cove Gap, Pennsylvania (Ulrich and Bassler, 1923). Zygosella postica Ulrich and Bassler. Right valve of male, x6, Niagaran, Cove Gap, Pennsylvania (Ulrich and Bassler, 1923).
+-
_g_qT_ -p-7
-+
38
Figure 6 (continued)
39
Summary and Conclusions Early Paleozoic ostracodes, a r e plentifully represented in t h e marginal northern Atlantic region, but a r e poorly known or generally sparse in O C C U ~r r e n c e in t h e southern Atlantic, where early Paleozoic rocks a r e restricted in distribution. Although ostracodes occur in small numbers in most of t h e marine facies of t h e north Atlantic margins, they a r e most abundant and biostratigraphically useful in t h e time-absorbent, slowly-deposited rock units and a r e sparse in thick clastics or massive carbonate units. Cambrian archaeocopids, although poorly known, appear to have limited biostratigraphic possibilities in t h e Atlantic region. Early Ordovician ostracodes a r e presently or potentially useful as assemblage zone faunas in t h e North Atlantic region, but differ in important respects on either side of t h e Atlantic probably owing to minimal transoceanic connections during t h a t Epoch. The middle Ordovician radiation of northwestern European ostracodes t h a t took place from Europe to t h e present western North Atlantic and in some instances to t h e southwestern Atlantic w a s a striking event and probably represents increasing proximity of not only trans-protoatlantic plates, but connections with Gondwanaland as well. In l a t e Ordovician t i m e unfavorable epicontinental facies obscured ostracode faunal interrelationships but t h e y apparently remained similar in t h e North Atlantic area. The hindrance posed by unfavorable facies to knowledge of ostracode faunas continued through t h e early Silurian, but in t h e l a t e Llandoverian and Wenlockian well developed ostracode zones appeared in both t h e eastern United S t a t e s and p a r t of western Europe. The Ludlovian faunas on t h e other hand a r e not well matched in t h e United States and western Europe, perhaps owing to arid-facies controls. The Pridolian-Downtonian Age marks a return t o trans-protoatlantic similarities of ostracodes in t h e United S t a t e s and western Europe.
40
References Cited Baldis, Bruno and Rossi d e Garcia, Elsa, 1972, Algunos ostracodes der Llandeiliense - Caradociense d e la Republica Argentina. Revista Espanola de Micropaleontologia, v. 4, no. 1, p. 19-22. Barrande, Joachim, 1872, Systeme Silurien du c e n t r e d e la Bohkme. Ire partie. Recherches pal6ontologiques. Supplement a u vol. I, Trilobites. Crustaces divers et Poissons. Plates in s e p a r a t e atlas. Prague and Paris, (fide -Bassler and Kellett, 1934). Bassler, R. S., and Kellett, Betty, 1934, Bibliographic Index of Paleozoic Ostracoda. Geol. SOC. Amer. Spec. Paper 1, 500 p. Berdan, J. M., 1967, Baltic ostracodes from Maine. (in) Geol. Survey Research, 1966. U.S. Geol. Survey Prof. Paper 550-A, p. A l l l . , 1970, American ostracode zonation: in Correlation of t h e North American Silurian rocks. Geol. SOC. Arne,: Spec. Paper, No. 102, p. 39-40, 1970. , 1971, Some ostracodes from t h e Schoharie formation (lower Devonian) of New York. in Paleozoic perspectives; a paleontological t r i b u t e to G. Arthur-Cooper, Smithson. Contrib. Paleobiol., No. 3, p. 161-174, illus., 1971. , 1972, Brachiopoda and Ostracoda of t h e Cobleskill Limestone (Upper Silurian) of Central New York. U.S. Geol. Surv., Prof. Paper, No. 730, 44 p., illus. (incl. sketch map), 1972. Bonnema, J. H., 1909, Beitrag zur Kenntnis der Ostrakoden der Kuckersschen Schicht (C2). Mitt. Min. Geol. Inst. Univ. Groningen, vol. 2, pt. 1, p. 1-84, pls. 1-8. Leipzig and Groningen. Butts, C., 1940, Geology of t h e Appalachian Valley in Virginia. Geol. Survey Bull. 52, p. 333-335. Canavari, M., 1899, Ostracodi Siluriani di Sardegna. SOC. Toscana Sci. Nat. Pisa., Pr. Verb., v. 11, 1899, art. 5 , p. 150-153. , 1900, Fauna dei calcari nerastri con Cardiola ed Orthoceras di Xea San Antonia in Sardegna I. Palaeont. ital., 1899, v. 5 , p. 187-210, 2 tav. Pisa. Clarke, J. M., 1899, The Paleozoic faunas of Para, Brazil, I. The Silurian fauna of t h e Rio Trombetas. 11. The Devonian Mollusca of t h e s t a t e of Para. Mus. Nac. Rio d e Janeiro, Arch. v. 10 (author's English edition), p. 1-100, pls. 1-8, 1900, Albany. Cobbold, E. S., 1936, The Conchostracea of t h e Cambrian a r e a of Comley, Shropshire with a note on a new variety of Atops reticulatus (Walcott). Geol. SOC. London, Q. J. no. 367, v. 92, pt. 3, p. 221-235, 2 PIS. , and Pocock, R. W., 1934, The Cambrian a r e a of Rushton (Shropshire). Roy. SOC. London. Philos. Tr. B501, v. 223, p. 305-409. Gurich, George, 1896, Das Palaeozoicum im Polnischen Mittel gebirges. Russ. Kais. Min. Ges., St. Petersburg, Verh., ser. 2, v. 32, p. 374-392, pl. 10, fig. 15, pl. 14, pl. 15 (part).
41
Harper, J. C., 1940, The upper Valentian ostracode f a u n a of Shropshire. Ann. Mag. Nat. Hist., v. 11, p. 385-400. , 1947, Tetradella complicata (Salter) and some Caradoc species of t h e genus. Geol. Mag., v. 84, p. 345-353, pl. 10. Henningsmoen, Gunnar, 1953a, Classification of Paleozoic straighthinged ostracods. Norsk Geol. Tidsskrift, v. 31, p. 185-288, 12 t e x t figs. , 1953b, The Middle Ordovician of t h e Oslo region, Norway, 4. Ostracoda. Norsk Geol. Tidsskrift, v. 32, p. 35-56, 5 pls., 1 t e x t fig. , 1954a, Lower Ordovician Ostracoda from t h e Oslo region, Norway. Norsk. Geol. Tidsskrift, v. 33, nos. 1, 2, p. 41-68, 2 pls. , 195433, Upper Ordovician ostracods from t h e Oslo region, Norway. Norsk Geol. Tidsskrift, v. 33, nos. 1, 2, p. 69-108, pls. 1-6. , 1954c, Silurian ostracods from t h e Oslo region, Norway. 1. Beyrichiacea with a revision of t h e Beyrichiidae. Norsk. Geol. Tidsskrift, v. 34, no. 1, p. 15-71, 8 pls., 5 t e x t figs. Hessland, Ivar, 1949, Investigations of t h e Lower Ordovician of t h e Siljan District, Sweden. Bull. Geol. Inst. Uppsala, v. 33, 408 p., 26 pls. Jaanusson, Valdar, 1957, Middle Ordovician ostracodes of c e n t r a l and southern Sweden. Bull. Geol. Inst. Uppsala University, v. 37, p. 176442, 15 pls. Jones, T. R., 1855, Notes on t h e Paleozoic bivalved Entomostraca, No. 1, Some species of Beyrichia of t h e upper Silurian limestones of Scandinavia. Ann. Mag. N a t . Hist. Ser. 2, v. 16, p. 31-92, pl. 5. , 1856, Notes on t h e Paleozoic bivalved Entomostraca, No. 3, Some species of Leperditia. Ann. Mag. N a t . Hist., ser. 2, v. 17, p. 81-101, pis. 6, 7. , 1884, Notes on t h e Palaeozoic bivalved Entomostraca, No. 17, Some North American Leperditiae and allied forms. Ann. Mag. Nat. Hist. Ser. 5, v. 14, p. 339-347. , and Holl, H. B., 1869, Notes on Palaeozoic bivalved Entomostraca, No. 9, Some Silurian species. Ann. Mag. Nat. Hist., ser. 4, vol. 3, p. 211-227, pls. 14, 15. Kay, G. M., 1934, Mohawkian Ostracoda: species common to Trenton faunules from t h e Hull and Decorah Formations. Jour. Paleontology, V. 8, p. 328-343, PIS. 44-46. , 1940, Ordovician Mohawkian Ostracoda: lower Trenton Decorah fauna. Jour. Paleontology, v. 14, p. 234-269, pls. 29-34. Kraft, J. C., 1962, Morphologic and systematic relationships of some middle Ordovician Ostracoda. Geol. SOC. Amer. Mem. 86, 104 p., 19 pl., 15 t e x t figs. Krause, Aurel, 189 1, Beitrag zur Kenntniss der Ostrakoden-Fauna in Silurian Diluvialgeschieben. Deutsch. Geol. Ges., Zeitschr., v. 43, p. 488-521, PIS. 29-33.
42
Kummerow, E., 1924, Beitrage zur Kenntnis der Ostracoden und Phyllocar iden aus nordischen Diluvialgeschieben. Preuss. Geol. Landes., Jahrb., 1923, vol. 44, p. 405-433, pls. 21,22. Lundin, Robert F., 197 1, Possible paleoecological significance of Silurian and early Devonian ostracode faunas from midcontinental and northeastern North America (with discussion): (in) Oertli, H. J., ed., Paleoecology of ostracodes, Cent. Rech. Pau, Bull., vol. 5 , (suppl.), p. 853-868. Martinsson, Anders, 1962, Ostracods of t h e Family Beyrichiidae f r o m t h e Silurian of Gotland. Bull. Geol. Inst. Univ. Uppsala, v. 41, p. 1369, 203 text-figs. , 1970, Correlation with Europe: (in) Correlation of t h e North American Silurian rocks, Geol. SOC. Amer., Spec. Paper, No. 102, p. 41-44, 1970. Poulsen, C., 1929, The Cambrian, Ozarkian, and Canadian faunas of northwest Greenland, Medd. om Gronland, v. 70 (Jubilaeumsekpeditionen Nord om Gronland 1920-23, no. 21, p. 308316, pl. 21. , 1934, The Silurian faunas of North Greenland. 1. The f a u n a of t h e Cape Schuchert Formation. Medd. om Gronland, v.72, (Jubilaeumsekpeditionen Nord O m Gronland, 1920-23) p. 1-46, pl. 1-3. Rozhdestvenskaja, A. A., 197 I , Ostracods and paleogeographic conditions of their distribution in a l a t e Devonian basin in t h e east of t h e Russian Platform. (in) Oertli, H. J., ed., Paleoecologie Ostracodes, Bull. C e n t r e Rech. Pau. - SNPA, v. 5, suppl., p. 763-768. Sarv, L., 1959, Ordovician ostracods in t h e Estonia SSR. Eesti Nsvteaduste Akademmia Geoloogia Instidundi Unrimused, v. 4, 206 p., 32 pls. Shaw, R. W. L., 1971, Ostracoda from t h e Underbarrow, Kirkby Moor and Scout Hill Flags (Silurian) near Kendal, West morland. Palaeontology, vol. 14, pa$ 4, p. 595-611, illus., 1971. Smith. A. G.. Briden, J. C., and Drewery, G. E., 1963, Phanerozoic worid maps. Palaeontol. Assoc. Spec. Paper. 12;~. 1-42. Swain, F. M., 1953, Ostracoda from t h e Camden C h e r t , western Tennessee. Jour. Paleontology, v. 27, p. 237-284, pls. 37-39, 21 t e x t figs. , 1957, Early Middle Ordovician Ostracoda of t h e e a s t e r n Stratigraphic d a t a and description of United States, Part I. Leperditiidae, Aparchitidae, and Leperditellidae. Jour. Paleontology, V. 31, p. 528, 590, pis. 59-62. , 1962, Early Middle Ordovician Ostracoda of t h e e a s t e r n United States, P a r t 11. , Cornell, J. and Hansen, D., 1961, Ostracoda of t h e Decorah Shale of Minnesota. Jour. Paleontology, v. 45, p. 519-545. 1933, Dimorphism and orientation in ostracods of t h e Swartz, F. M., Family Kloedenellidae from t h e Silurian of Pennsylvania. Jour. Paleontology, v. 7, p. 231-260, pls. 28-30.
43
, 1939, Keyser Limestone and Helderberg Group, (in) B. Willard et al., The Devonian of Pennsylvania, Penn. Geol. Survey 4th Ser. Bull. G 19, p. 29-72. , and Whitmore, F. C., Jr., 1956, Ostracoda of t h e Silurian Decker and Manlius limestones in New Jersey, and e a s t e r n New York. Jour. Paleontology, v. 30, p. 1029-1091, pls. 103-110, 3 t e x t figs. Tromelin, G. and Lebesconte, P., 1876, Essai d'un catalogue raisonne des fossiles Siluriens des Departments d e Maine-et-Loire, d e l a Loire infe'rieure et du Morbihan. Assoc. Francaise I'Avanc. Sci., C.R., 4th Session, 1875, p. 623, Paris. Tschigova (Chizhova), V. A., 197 1, Geographical distribution of ostracods in t h e European sea basin a t Famennian time: (in) Oertli, H. J., ed., Paleoecology of ostracodes, Cent. Rech. Pau, Bull., vol. 5, (suppl), p. 755-76 1. Ulrich, E.O.,1890, 1891, New and l i t t l e known American Paleozoic Ostracoda. Cincinnati SOC. Nat. Hist., Jour., v. 13, p. 104-137, (ISSO), p. 173-211, (1891), 8 pis. , and Bassler, R. S., 1908, New American Paleozoic Ostracoda. Preliminary revision of t h e Beyrichiidae, with descriptions of new genera. U.S. Nat. Mus., Pr., i. 35, p. 277-340, figs. i61, PIS. 37-44. , 1923, Maryland Geological Survey, Silurian volume, 794 p., 27 figs., 67 pls. Paleozoic Ostracoda: their morphology, classification and occurrence, p. 27 1-39 1. Systematic paleontology of Silurian deposits (Ostracoda) p. 500-704, pls. 36-65, Baltimore. , 1931, Cambrian bivalved C r u s t a c e a of t h e Order Conchostraca. U.S. Nat. Mus. Proc., v. 78, P a r t 4, p. 1-130, pl. 1-10.
44
Additional References Not Cited Abushik, A. F., 1967, The Importance of ostracods in drawing t h e Siluro-Devonian boundary in t h e European part of t h e USSR. (in) Oswald, D. H., ed., Internat. Sympos. on Devonian System, Calgary, Alberta, Alta. SOC. Pet. Geol., v. 2, p. 875-884. Brookins, Douglas G; Berdan, J e a n M.; and Stewart, David B., 1973, Isotopic and Paleontologic Evidence for correlating t h r e e volcanic sequences in t h e Maine coastal volcanic belt. Geol. SOC. Am., Bull., v. 84, no. 5, p. 1619-1628. Guber, A. L. and Jaanusson, Valdar, 1964, Ordovician ostracodes with posterior domiciliar dimorphism. Bull. Geol. Inst. Univ. Uppsala, v.42, no. 53, 43 p., 6 pls. Gurevich, K. Ya., 1971, (Silurian-Devonian boundary and S t a g e Subdivision of t h e Lower Devonian in t h e Lvov Paleozoic Depression): Meyhdunar. Simp. Granitsa Silura Devona, Biostratigr. Silura, Nizhega Srednego Devona, Tr., no. 3, v. I, p. 79-85. Krandievsky, V. S., 1963, Fauna ostracod siluriiskikh vidkladiv Podillya, Akad. Nauk. Ukrain. RSR, Inst. Geol. Nauk, Kiev, 148 pp. , 1971, (The Silurian-Devonian Boundary of Volyn-Podolia based on Ostracods and Graptolites): Mezhdunar. Simp. Granitsa Silura Devona, Biostratigr. Silura, Nizhego Srednego Devona, Tr., no. 3, v. 1, p. 108-113, (inc. Engl. sum.). Le Fevre, J., 1967, (Succession of Ostracod and Conodont Associations in t h e Silurian Lower Devonian, and Eifelian from Several Sections in France and t h e Sahara). (in) Colloque sur l e devonien inferieur et ses limites (Rennes, 16-24 Septembre 19641, Fr., Bur. Rech. Geol. Minieres, mem., no. 33; p. 373-389. Martinsson, A., 1963, Kloedenia and related Ostracode genera in t h e Silurian and Devonian of t h e Baltic a r e a and Britain. Geol. Inst. Univ. Uppsala, Bull. 42, p. 1-63. , 1965a, The Siluro-Devonian ostracode genus Nodibeyrichia and faunally associated kloedeniinen. Geol Foren. Forhandl., v. 87, p. 109-138. , 1965b, Remarks on t h e Silurian ostracode genus Craspedobolbina from t h e Baltic a r e a and Britain. Geol. Foren. Stockholm, Forh. v. 87, pt. 3, no. 522, p. 314-325, illus., 1965. , 1967, The succession and correlation of ostracode faunas in t h e Silurian of Gotland. Geol. Foren. Forhandl., v. 89, p. 350-386. , 1968a, An Appalachian species of t h e Silurian Ostracode Genus Craspedobolbina. Geol. Foren. Stockholm, Forh., v. 90, pt. 2, no. 633, p. 302-208, illus., 1968. , 1968c, The Appalachian species of t h e Silurian Ostracode Genus Craspedobolbina. Geol. Foren. Forhand, Stockholm, v. 90, p. 302-308, 6 text-figs.
45
Petersen, L. E.; and Lundin, R. F.., 1974, Thlipsura martinssoni, a new ostracode species from t h e Silurian of England. J. Paleontol., v. 48, no. 2, p. 357-360. Pranskevichus, A. A., 1970, Silurian ostracod assemblages of t h e south Baltic Region and their correlation value: Acad, Sci. USSR, Dokl., Earth Sci. Sect., vol. 192, p. 83-85. , 1971, (New ostracodes of Wenlockian a g e in t h e southern Baltic region): (in) Paleontologiya i stratigrafiya Pribaltiki i Belorussii, sbornik 111, p. 51-60 (incl. Engl. sum,), illus., Lit. NauchnoIssled. Geologorazc. Inst. Vilnius., 1971. , 1971, (Paleontological characteristics of lower Silurian in t h e southern Baltic region based on studies of Ostracods): (in) Paleontologiya i stratigrafiya Pribaltiki i Belorussii, shomik 111, p. 6170 Inst., Vilnius, 1971. , 1972, (Silurian ostracods of t h e southern Baltic sea): Liet. Geol. Mokslinio Tyrimo Inst., Tr,, no. 15, 180 p. (incl. Lith., Engl. sum.), illus. (Incl. sketch map), 1972. ,1972, (New Silurian rishonids f r o m t h e South Baltic area): (in) Novyye vidy drevnikh rasteniy i bespozvonochnykh SSSR, p. 272275, illus., Akad. Nau, SSSR, Nauch. Soviet Probl., Moscow, 1972. ,1972, (Silurian Ostracoda of t h e south Baltic region): Vilnius, Geol., Inst., Darb., no. 15, 280 p. (incl. Lith., Engl. sum.), illus. (incl. sketch map), 1972. , 1972, Ostracods from t h e Upper Silurian of t h e Southern Baltic region: Geol. Foeren. Stockh., Foerh., v. 94, p a r t 3, no. 550, p. 439-447, ilus., 1972. , 1972, New ostracods of t h e Llandoverian of Lithuania. Geol. Foren. Stockh., Foerh., v. 94, p a r t 3, no. 550, p. 435-438, illus., 1972. , 1973, New l a t e Silurian ostracodes from t h e South of t h e Baltic region: Paleontol. J., v. 7, no. 1, p. 32-42, illus., 1973. , 1973, (New l a t e Silurian ostracods from t h e South of t h e Baltic region): Paleontol. Zh., no. I, p. 39-47, illus., 1973. , 1973, New l a t e Silurian ostracodes from t h e south of t h e Baltic region): Paleontological Journal, v. 7, no. I, p. 32-41, pls. 3, 4. Robardet, M.; Henry, J. L.; Nion, J.; et al., 1972, (Pont d e Daer (Caradocian) Formation in t h e Domfront and Sees Synclines, Normandy): SOC. Geol. Nord., Ann., v. 92, no. 3, p. 117-137 (incl. Engl. sum), illus, 1972. Sarv, L., 1973, (Zonation of t h e Silurian in t h e section of t h e Kalvariya Well based on ostracods): Eesti NSVstead. Akad., Toim., Keemia Geol., v. 22, no. 2, p. 88-91, illus., 1973. Schallreuter, R U, 1967, (New ostracods f r o m Ordovician boulders). Geologie (Berlin), v. 16, no. 5, p. 615-631 (incl. Engl. Russ. sum.) illus., 1967.
46
, 1968, (Ordovician Podocopida; Beecherellidae). Neues Jahrb. Geol. Palaontol., Abh., v. 131, no. 1, p. 82-96 (incl. Engl. sum.), illus., 1968. , 1969, Neue Ostracoden aus ordovizischen Geschieben, 11. Sonderdruck aus Geol. Jg. 18, H.2, Akad-Verlag Berlin, p. 203-215, 3 PIS. , 1971, Ostrakoden aus Ojlemyrgeschrieben (Ordoviz) N. jb. GeoLPalaont, Mh, Jg. 1971, H. 7, p. 423-431, 1 pl. , 1971 (Asymmetric Ordovician Ostracods): Neues Jahrb. Geol. Palaeontol., Monatsh, no. 4, p. 249-260 (incl. Engl. sum.), illus., 1971. ,1971, (Ostracods from t h e Ojlemyr flint, Ordovician): Neues Jahrb. Geol. Palaeontol., Monatsh, no. 4, p. 423-431 (incl. Engl. sum.), illus., 1971. , 1972, (Drepanellacea (Ostracoda, Beyrichicopida) f r o m t h e middle Ordovician Backstein Limestone boulders: 4, L a t e r o hores hystrix n. sp., Pedomphalella germanica n. sp. and Easc midtella fragosa): Dtsch. Ges. Geol. Wiss., Ber., Reihe A. Geol. Palaontol., v. 17, no. I, p. 139-145, illus., 1972. Shaw, R. W. L., 1971, The faunal stratigraphy of t h e Kirky Moor f l a g s of t h e type a r e a near Kendal, Westmorland: Geol. J., v. 7, p a r t 2 , p. 359-380, illus. (incl. geol. m a p 1:63, 3601, 1971. Sidaravichene, N.V., 1971, (New Ostracods from t h e middle and upper Ordovician of Lithuania), (in) Paleontologiya i stratigrafiya Pribaltiki i Belorusii, sbornik 111, p. 23-36 (incl. Engl. sum.), illus. (incl. sketch map), Lit. Naerchno-Issled. Geologorazv. Inst., Vilnius, 1971. Whitfield, R. P.! 1890, Observations on some imperfectly known fossils from t h e calciferous sandrock of Lake Champlain and descriptions of several new forms. Am. Mus. N a t Hist. Bull. 2, 56-80., pl. 13, figs. I6. Zbikowska, B., 1973, (Upper Silurian Ostracods from t h e Leba Elevation, northern Poland): (in) P r a c e zwiazane y problematylca struktur wg lebnych Polski, A c t a Geol. Pol., v. 23, no. 4, p. 607-644, (incl. Engl. sum.), illus., 1973. Zenger, D. H., 1971, Uppermost Clinton (Middle Silurian) Stratigraphy and Petrology, East-Central New York: N.Y. State Mus. Sci. Serv., Bull., no. 417, 58 p. illus. (incl. sketch map), 1971.
+
47
Discussion S . Bergstr8m: I w a s most i n t e r e s t e d t o l e a r n a b o u t t h e B a l t i c a f f i n i t i e s of t h e o s t r a c o d e s of t h e San J u a n F o r m a t i o n o f A r g e n t i n a , a n d r e f e r -
Dr.
ence t o i t a s b e i n g o f L l a n v i r n i a n a n d L l a n d e i l a n a g e . The c o n o d o n t f a u n a o f t h i s u n i t a l s o shows B a l t i c a f f i n i t i e s b u t , a s n o t e d by S e r p a g l i (1974), i t s u g g e s t s a l a t e A r e n i g i a n r a t h e r t h a n y o u n g e r a g e o f t h e f o r m a t i o n . Can t h e S a n J u a n o s t r a c o d e s be u s e d t o d a t e t h e f o r m a t i o n , and i f s o , d o t h e y show c o r r e l a t i o n w i t h L l a n v i r n i a n and L l a n d e i l a n s t r a t a i n n o r t h w e s t e r n E u r o p e r a t h e r than with Arenigian u n i t s ? Swain: The few s p e c i e s o f o s t r a c o d e s from t h e S a n Juan Formation are n o t d e f i n i t i v e i n a g e , b u t a r e more s u g g e s t i v e o f m i d d l e t h a n o f e a r l y Ordo v i c i a n . Reconstructions of c o n t i n e n t a l p o s i t i o n i n t h e O r d o v i c i a n by S m i t h e t a 1 . ( 1 9 6 3 ) s h o w t h a t s h a l l o w m a r i n e m i g r a t i o n s b e t w e e n t h e B a l t i c area and w e s t e r n S o u t h America c o u l d r e a s o n a b l y h a v e occurred. D r . M . C . Keen: I n y o u r a b s t r a c t you m e n t i o n e d b r a c k i s h water o s t r a c o d s i n t h e L u d l o v i a n . When d i d b r a c k i s h water o s t r a c o d s f i r s t a p p e a r ? How d o you r e c o g n i s e them and a r e t h e r e a n y f r e s h - w a t e r ones? Swain: J e a n B e r d a n and I h a v e d i s c u s s e d t h i s s e v e r a l times. C e r t a i n e l o n g a t e , s u l c a t e (eukloedene l l i d o s t r a c o d e s o f t h e S i l u r i a n may r e p r e s e n t e i t h e r b r a c k i s h o r h i g h - s a l i n i t y forms; t h e y occu r i n f o s s i l i f e r o u s r e d b e d s (Bloomsburg) a s d e s c r i b e d by H o s k i n s from c e n t r a l P e n n s y l v a n i a . D r . J . Berdan: I would a g r e e w i t h t h a t , a n d a l s o I think t h a t the difference i n t h i s context, b e t w e e n t h e A p p a l a c h i a n f a u n a , d o m i n a t e d by k l o e d e n e l l i d s , and t h a t t o t h e w e s t , s u c h a s t h e Henr y h o u s e Formation where k l o e d e n e l l i d s a r e n o t abu n d a n t , i n d i c a t e s t h a t t h e r e w a s something p e c u l i a r about t h e environment i n t h e Appalachian area a t t h a t t i m e . Swain: One o f t h e p r o b l e m s i s t h a t t h i s w a s a t i m e d u r i n g which e v a p o r f t e c o n d i t i o n s o c c u r r e d i n t h e A p p a l a c h i a n r e g i o n , so i t i s d i f f i c u l t t o know w h e t h e r t h e s e a r e h i g h o r low s a l i n i t y f o r ms. A s f a r as f r e s h w a t e r forms are concerned, w e have i n d i c a t i o n s of f r e s h w a t e r s p e c i e s i n t h e C a r b o n i f e r o u s and p e r h a p s i n t h e Devonian.
--
48
Berdan: I think they are earlier and appear in the middle or late Silurian. Murray Copeland has more information. Dr. J. E. Conkin: Does the Pridolian represent a pulsation (transgression) of the seas onto the continents? If so what is the bearing of this on the Silurian-Devonian boundary? Swain: The result of a late Silurian transgression in the Appalachians, as well as a return to a more favorable benthic environment seems to have been the production of a transitional Siluro-Devonian population of ostracodes and other invertebrates in such units a s the Keyser Limestone.
49
PALEOZOIC SMALLER FORAMINIFERA OF THE NORTH AMERICAN ATLANTIC BORDERLANDS by James E. Conkin, University of Louisville, Lou. Ky. 40208 and Barbara M. Conkin, Jefferson Community College, Lou. Ky. 40202 Abstract Species of Paleozoic smaller Foraminifera, combined . with physical evidence of diastrophism (paracontinuity), may be used in precise placement of time-stratigraphic boundaries in the eastern United States as with the Devonian-Mississippian and KinderhookianOsagean boundaries. Evolutionary lineages a r e recognized in t h e Devonian and Mississippian and certain stratigraphic intervals within the Lower and Middle Paleozoic are delimited by foraminifera1 zones; nevertheless, even though the stratigraphic value of Paleozoic smaller Foraminifera has been demonstrated, their full potential remains to be realized. Agglutinated Foraminifera frequently occur in shale and limestone containing silt or fine sand; however, their distribution varies in diverse lithologies within single genetic units. Sparry bioclastfc calcarenites rarely contain agglutinated Foraminifera, but often bear calcareous forms. Earliest textulariinid Foraminifera, consisting of simple, free or attached, tubular, globular, spherical, or hemispherical chambers, arose by mid-Ordovician. Tubular t o hemitubular attached forms appeared in late Ordovician. Apertured planoconvex, tubular, irregularly coiled and planispirally coiled forms and globular forms with various projections, arose in early Silurian. Multichambered Highly forms with enlarging chambers began in late Devonian. organized, attached hemitubular, and well-organized multichambered forms, appeared in early Mississippian and evolved into more complex multichambered forms (including attached forms) in Pennsylvanian and Permian. Microgranular calcareous (fusulininid) foraminiferans arose in Middle Devonian and increased in number and complexity in Mississippian, Pennsylvanian, and Permian. Calcareous porcelaneous (miliolinid) tests appeared in early Pennsylvanian and calcareous hyaline (rotaliinid) tests evolved in early Permian.
50
\
Les esp\eces d e petits foraminiferes pale*ozoiques, unies 3 l a preuve physique du diastrophism (paracontinuit&), peuvent Ae t r e utilisdes dans l'emplacement pr6cis des frontikres d e temps-stratigraphiques dans l a rdgion est des Etats-Unis c o m m e a v e c l e s frontikres Les lignages d6vonien-mississippiens et kinderhookien-osagkens. 4volutionnistes se sont reconnus dans l e dkvoni/en et l e mississippien. Certains intervalles stratigraphiques dans l e paleozoique infdrieur et l e mi-palebzoique sont delimitds par d e s zones foraminif6rales; n6anmoins, malgre' la valeur stratigraphique dCyontre'e des petits foraminifkres pale'ozoiques, leur potentialit6 r e s t e a S t r e rCalise'e. Des foraminifkres agglutings souvent se trouvent dans d e s schistes argileux et d e s calcaires contenant d e s sables fins ou d e s vases. Cependant, leur distribution se varie dans d e diverses lithologies l'int6rieur d e simples unit& g6nCtiques. I1 est :are q u e les sparry calcarenites bioclastiques contiennent des foraminiferes agglutin6s mais souvent donnent des formes calcaires. Par l e mi-ordovicien, les plus anciens foraminif&es textulariinid apparaissaient. 11s contenaient des chambres, soit simples, soit libres ou lides, soit tubulaires, g!obulaires, sphgriques o u he'misph6riques. Des for mes lids tubulaires a l'hdmi-tubulaires apparaissaient pendant l'ordovicien r6cent. Des formes planoconvexes \ apertures, ainsi que tubulaires, lov& irre'gulikrement et loves planispiralement et d e s for mes globulaires projections diverses apparaissaint pendant l e d6but du silurien. Des f o r m e s multichambrks a v e c d e s chambres aggrandissantes commensaient pendant l e dgvonien re'cent. Des f o r m e s multichambrds l i i s , he'mitubulaires, assez complexes et organisds apparaissaient pendant l e d6but du mississippien et se sont Cvoluks e n des f o r m e s multichambr6s plus complexes (y compris l e s formes lids) pendant l e pennsylvanien et l e permien. Des foraminifkres microgranulaires calcaires (f usulininid) apparaissaient dans l e midevonien et se sont augment& e n nombre et complexitd pendant l e mississippien, le pennsylvanien et l e permien. Des tests calcaires porcelanaires (miliolinid) apparaissaient pendant l e d6but du pennsylvanien. Des tests calcaires hyalines (rotaliinid) se sont kvoluds pendant l e ddbut du permien.
>
51
Introduction Agglutinated and microgranular, porcelaneous, and hyaline calcareous (excluding fusulinid, endothyrid, and tournay ellid) foraminiferans may be considered %mailer" Foraminifera; however, even these may require sectioning for identification. Paleozoic smaller Foraminifera from the North American Atlantic borderlands (Texas to New York and Ontario), have been studied for nearly 50 years, but the quality of the work is uneven. Reviews of North American late Devonian-early Mississippian agglutinated Foraminifera (Conkin and Conkin, 1967 and 1970) and smaller Foraminif era of the pre-Pennsylvanian (Conkin and Conkin, 1973a) stressed their stratigraphic significance. Agglutinated Foraminifera occur abundantly in shale (Conkin, 1961, p. 234) which contains silt and/or fine-grained sand, as well as in limestone (even sublithographic) if fine-grained detrital material is present, but they may be unevenly distributed in diverse lithologies within a single genetic unit; limestone lenses and impure ironstone concretions may contain many specimens, while surrounding shales, silty shales or shaly siltstones bear essentially none. Sparry bioclastic calcarenites rarely contain agglutinated forms, but calcareous foraminiferans are frequently found therein. The dominance of agglutinated (textulariinid) Foraminifera in early Paleozoic resulted from their early evolution and pioneer occupation of shallow marine benthonic niches; with evolution in the Devonian of calcareous Foraminifera, competition ensued and increased significantly from Carboniferous onward, with calcareous groups progressively replacing agglutinated forms in "normal epicontinental" marine environments. Paleozoic Genera Of Smaller Foraminifera In preparing generic keys and ascertaining ranges of genera (Text-figs. 1-31, we have exercised our best judgment in determining the validity of records of stratigraphic occurrence and taxonomic assignment. The keys serve to clarify generic identities and relationships. We consider several genera recorded from the Paleozoic t o be junior synonyms (in parentheses) of the following: Calcitornella (ADterrinella. Calcivertella and Plummerinella), Hemisphaerammina (Fairliella; Metamorphina, and l T W e b b i n r n , A m movertella (Ammodiscella), Oryctoderma (Crithionina), Ordovicina (Croneisella, Gastroam mina, and Shidelerella), Nodosinella - Climacammina Mono enerina), Palaeotextularia (Paratextularia), &:ml Saccam mina (Proteonina, Raibosammina and T v Lunucam mina (Geinitzina), and Tholosina (Amphicervicis). Other genera are so ill understood '(Agathamminoides, Kerionammina, Tuberitina, Stacheia, and Stacheoides) that no useful statement can be made. Some genera have been recorded erroneously from the Paleozoic (Astrammina, Astrorhiza, Bullapora I and
52
ORDOVICIAN
SILURIAN
IARLY IMIDDLEI LATE
EARLY IliIDDLEI LATE
DEVONIAN EARLY lHIDDLE[ LATE
’-ASCHEMONELL
EUDOPALMULA
1
I
TEXT-FIGURE
1.
GENERIC RANGES OF PALEOZOIC SMALLER
53
IIISSISSIPPIAY EARLY
I MIDDLE^
LATE I
ORTHOVERTELLA
AMVOVERTELLA
PREPEILOPSIS PLACOPSILIIUA
F O W I N I F E R A IN THE NORTH AMERICAN ATLANTIC BORDERLANDS.
54
Schizammina); and still others have been incorrectly assigned t o the Foraminif era (Thuram minoides, a radiolarian, and Weikkoella, a charophyt e). Ranges And First Occurrences Of Some Genera Of Foraminifera Text-figure 2 presents the stratigraphic ranges of genera of Foraminifera in the Paleozoic of the North American Atlantic borderlands arranged systematically. Text-figure 3 presents t h e genera arranged according to their first occurrences (see Text-figure 1 for illustrations of genera). The agglutinated Foraminifera are represented by the suborder Textulariina, with two superfamilies, the Ammodiscacea (globular, tubular, or irregular tests with one or more apertures) and the Lituolacea (multilocular, rectilinear, enrolled or uncoiled, or trochospiral coil modified to biserial, tests). The suborder Fusulinina (with a nonlamellar, microgranular, calcite wall) is represented by two superfamilies, the Parathuramminacea (with globular or tubular tests with one or more apertures) and the Endothyracea (with multiple, rectilinear, enrolled, or biserial chambers and single to multiple apertures). The suborder Miliolina (with an imperforate, porcelaneous, calcite wall) is represented by the superfamily Miliolacea (comprising forms with a proloculus and tubular second chamber, planispirally or variously coiled) and the suborder Rotaliina (calcareous, hyaline, perforate, lamellar tests) is represented by the superfamily Spirillinacea. The first textulariinid Foraminif e r a had simple tests consisting of free or attached, tubular, globular, spherical, or hemispherical chambers; by Middle Ordovician, the tubular Bathysiphon, H perh e ammina, and Rhabdammina, the box-like planoconvex Hemisphaeram mina and tubular to hemitubular attached To1 pammina and the globular t er simple forms appeared Blastammina arose in late Ordovocian. in early Silurian: the globular Psammosphaera and Sorosphaera and planoconvex -Webbinelloidea, . as well as the tubular, irregularly coiled GlomospirE Lituotuba and the planispirally coiled Ammodiscus. Barly Sil- ' A globular forms with various projections are Saccammina, La enarrimina, Thurammina, and Pseudastrorhiza. The first multicham ere agg utinated forms with chambers enlarging in size a r e found in late Devonian: the attached multichambered Oxinoxis and the free
+
b
my-
TEXT-FIGURE
2.
GENERIC RANGES OF PALEOZOIC SMALLER FORAMINIFERA, ARRANGED ACCORDING TO
SYSTEMATIC CLASSIFICATION.
SOLID LINE INDICATES RECOROED OCCURRENCES; DOTTED LINE,INFERRED OCCURRENCES. ARROW INDICATES RANGE ABOVE PALEOZOIC. 55
TEXT-FIGURE
3. GENERIC RANGES OF PALEOZOIC SMALLER FORAMINIFERA &KANGED ACCORDING TO THEIR F I R S T OCCURRENCES.
56
Reophax (first abundant in early Mississippian). The attached, spiral, tubularTrepeilopsis is first found in late Devonian. The organized attached hemitubular form, Ammovertella, appeared in early Mississippian when the well-organized multichambered Trochammina and Ammobaculites also appeared. In early Pennsylvanian other complex agglutinated for m s appeared : Haplophrag m oides, Text ularia , Bigenerina, and Mooreinella. Finallv, in late Pennsylvanian, the Y attached, multichambered P1acopsilina"arose. The earliest known calcareous foraminiferan, the complex, multichambered Semitextularia, first appeared in Middle Devonian. Two simpler calcareous forms are the two-chambered (proloculus and tubular second chamber) rectilinear Earlandia and the planispiral Cornuspira which appear in middle Mississippian. More complex calcareous genera are found in late Mississippian: Palaeotextularia, Cli macam mina, and Tetra taxis. A number of new calcareous forms appeared in Early Pennsylvanian: Nodosinella, Polytaxis, Valvulinella, Globivalvulina, Brad h a , Aeathammina. Calcitornella. and SDirillina. Textularia. Hemiaor lus, a i d Rectocornuspira appeared in middlePennsylvanian, and, in late Pennsylvanian two additional multichambered genera, Glyphostomella and Lunucammina. Finallv. in earlv Permian, a comdex, multichambered rotaliinid calcarGous genus; Patellina,- was introduced.
-41
Zonation By Smaller Foraminifera And Devonian Early Mississippian Evolutionary Lineages
We restrict our zonation to the pre-Chesterian portion of the Paleozoic and include only those Foraminifera which a r e taxonomically valid, widely distributed, and occupy well-defined, rather short stratigraphic intervals. Text-figure 4 illustrates and presents notes on the recognized foraminifera1 zones as well as evolutionary lineages recognized by us in the Devonian-early Mississippian. Smaller Foraminifera And Time-Stratigraphic Boundaries Early Mississippian smaller Foraminif e r a are best known of any in the Paleozoic and a number of species have quite restricted ranges, and can be used for age determination, correlation, and boundary placement. Using agglutinated Foraminifera (Text-figure 5) as the biologic parameter and paracontinuities as the physical parameter, we have succeeded in precise placement of t h e long-sought DevonianMississippian boundary, as well as the Kinderhookian-Osagean boundary, in east-central United States (Conkin and Conkin, 1973b and 1975).
-
I
MI S S I S SI PPIAN
c
Oxinoxts Iigula,
Tolypommina bulbosa,
Tolypammina gersterensia,
Tolypommlna cyclops,
Tolypommina jacobschapelensis,
Hyperammino rockfordensis Ammodiscus longexssrtus,
Ammodiscus semiconstrictus,
Hyperommino coaleri Hyperommino compocto, Hyperommino yrocilenta,
I
Rcophnx lochryrnosus
I
Tolypommino bronsoni, Tolyparnminn friizelli, Ammoboculites lepfns,
Hyperommino conslricta,
Thurornmino ? iriradiota, Trepeilopsis glornospiroides, Ammovertella ltsoe,
Reophox calothus
Ammovertella p i k e i , Arnrnobaculites qutschicki, Trepeilopsis recurvidens hnmoboculites beveridgei, rrochommino mehli,
O r i n o r i s swollowi
Hyperammino nitido,
Sorosphoero ?cooperensis,
Trepeilopsis prodigalis,
Saccammino howei, Reophaa Reophax nor t hviewensis, Pseudostrorhizo
buccino,
Lituotu ba semiplana
conic0 , Pseudastrorhiza delicato
, Amrno-
boculites choppelensis, xmsphaero papilla, Thummmina congesta Pseudostrorhiro digitata, rhiza bocculo,
Pseudastrorhiza lonceola, Pseudmiro-
Tolypammina loocoon, Tolypomrnina botanuncus
Tolypammina r o t u l a Reophax %.A,
Thurornmino orenacorna
TEXT-FIGURE 4. FORAMINIFERAL ZONATION IN THE l.OWER PALEOZOIC AND EVOLUTIONARY LINEAGES IN THE DEVONIAN-LOWER MISSISSIPPIAN.
AND MIDOLE NORTH AMERICA.
References Cited Conkin, J. E., 1961. Mississippian smaller Foraminifera of Kentucky, southern Indiana, northern Tennessee, and south-central Ohio: Bull. Amer. Paleont., v. 43, no. 196, p. 129-368.
, and Conkin, B. M., 1967. Arenaceous foraminifera as a key to Upper Devonian and Lower Mississippian relationships in the type Mississippian area: Essays in Paleontology and Stratigraphy. R. C. Moore Commemorative Volume, Spec. Pap. Univ. Kansas, Dept. of Geol., no. 2, p. 85-101. , 1970. North American Kinderhookian (Lower , and Mississippian)naceous For am inif era: Com pte Rendu 6e Congress Intern. Strat. Geol. Carbonif., Sheffield, 1967, v. 2, p. 575-584. and , 1973a. Pre-Pennsylvanian Foraminifera of North America: XXII Int'l. Geol. Congress, India, 1964, p. 319335. Y
, 1973b. The paracontinuity and the determination of the Devonian-Mississippian boundary in the type Lower Mississippian area of North America: Univ. Louisville Studies in Paleont. and Strat. No. 1, Univ. Lou. Reprod. Serv., 36 pp. , and , 1975. The Dovonian-Mississippian and Kinderhookian-Osagean boundaries in the east-central United States are paracontinuities: Univ. of Louisville Studies in Paleont. and Strat. No. 4, Univ. of Lou. Reprod. Serv., 54 pp. Loeblich, A. R., Jr., and Tappan, H., 1964. Sarcodina, chiefly "Thecamoebians" and Foraminifera. In: Moore, R. C., Ed., Treatise on Invertebrate Paleontology. New York: Geol. SOC. Amer., and Univ. Kans. Press, pt. C, v. 1, p. i-xxxi, Cl-C510a.
59
Discussion Dr. R. A. Olsson: Have you done any work on the facies relationships of these fauna? Dr. J . E . Conkin: Well, we d i d n ' t have space in the paper t o go into facies. Agglutinate foraminiferans can, and d o , occur i n great abundance in some Paleozoic limestones, such as the Upper Devonian Louisiana Limestone of northeastern Missouri and western I l l i n o i s (sublithographic limestone bearing much fine-grained s i l t ) which carries a magnificent agglutinate foraminifera1 fauna. Here's a paper [Conkin and C o n k i n , 1964, Devonian Foraminifera, Part 1 - The Louisiana of Missouri and I l l i n o i s : Bull. Amer. Pal., v . 47, no. 213, pp. 53-105, pls. 12-15, text-figs. 1-5, charts 1-31 on i t . I d o n ' t say t h e r e ' s no ecological control for there i s a l ways some. For example, in sparry bioclastic limestone there are few agglutinate foraminiferans f o r often n o t enough det r i t a l grains are available for building t h e i r t e s t s ; the Lower Mississippian (Osagean) Burl ington Limestone of Missour i and I l l i n o i s bears no agglutinate foraminiferans for t h i s reason. I n the essential equivalent of the Burlington Limestone, the New Providence Shale of northwestern Kentucky and southern I n d i a n a , agglutinate foraminiferans are f o u n d . I n some shales and s i l t s t o n e s units in the Osagean Brodhead Formation o f Kentucky and Indiana ( i n which very few and/or poorly preserved arenaceous foraminiferans are f o u n d ) , there are occasional intercalated limestone or ironstone nodules and lenses which produce, upon acidization, well preserved agglutinate foraminiferans. Some agglutinate species have quite restricted stratigraphic ranges such as Hyperammina kentuckyensis which i s a definitive Osagean marker occurring in b o t h limestones and shales. I n general, we may say t h a t in Mississippian limestone facies, the tournayellids and endothyrids are dominant while i n the shale facies the agglutinate foraminiferans dominate.
This Page Intentionally Left Blank
61 LATE PALEOZOIC OSTRACODES OF WESTERN EUROPE AND NORTH AMERICA: A REVIEW L u i s C. Sanchez D e Posada
U n i v e r s i d a d d e O v i e d o , O v i e d o , Espana
ABSTRACT
it is perhaps due t o t h e f a c t t h a t ostracodes have proved to be succ e s s f u l i n so many d i f f e r e n t environments adapting themselves t o every vari a t i o n i n :emperature, s a l i n i t y , water energy, s u b s t r a t m , depth, t h a t t h e y o f t e n sei? l e s s u s e f u l f o r b i o s t r a t i g r a p h i c a l a p p l i c a t i o n s , a t l e a s t a s f a r a s 1ong-d:siince c o r r e l a t i o n s a r e concerned. On t h e one hand we observe several !ong-ranging s p e c i e s , which p e r s i s t i n diachronous f a c i e s and on t h e other &and t h e r e exist many groups which are a b l e to s u r v i v e small f a c i e s c i i a y e s w i t h small changes in t h e i r morphology. The b i o s t r a t i g r a p h i c use o f Paleozoic ostracodes is f u r t h e r hampered b3 t h e f a c t that up to now we have only t h e r a t h e r i s o l a t e d r e s u l t s of work ostracodes from d i f f e r e n L areas and environments a t our d i s p o s a l . T h i s s i t u a t i o n may improve w i t h i n t h e forthcoming y e a r s as has been shown by t h e d'scovery o f so-called s i l i c i f i e d Devonian faunas and Devonian-CarbonlferHowever, at she p r e s e n t time t h e s e a r e o n l y ou8 entomozoacean faunas. k n c m from northwestern Europe, Spain, Portugal, and ilorth America. A t t h e s m e time attempss are being made t o come t o long-distance c o r r e l a t i o n s o f :'?per Devonicr and 9inan:ian assemblages betveen t h e USSR and western Surope by ~?ussiano s t r a c o d o l o g i s t s . In Euurope, she a t t e n t i o n o f Devonian t o Permian ostracode workers has up to nod T a i n l y been concentrated on d e s c r i p t i v e palaeontology and i n t h e l a s t decade 0% p l a e o e c o l o g y . RfSUMi C ' e s t un f a < t que l e s ostracodes o n t rgussi & s ' a d a p t e r 'a beaucoup d ' e n u i r o n n e ~ e n t sdi;;r^e'rerts e t h beaucoup de v a r i a t i o n s de l a tempgrature, de l a sa:initi, de l ' E w r g i e d'eau, de ;a s o u s t r a t e e t de l a profondeur. ~ ' e a :peut-&re h cause de c e c i q?ie l e e ostracodes semblent Ptre moins util e s quans a m a p p l i c a t i o n s b i o s t r a t i g r a p h i q u e f , du moins en ce q u i touche ! e s c o r r e l a t i o n s de longue-portei. D 'un c d t e , on remarque p l u s i e u r s e s pbces h grsnde ,&is;ribu$ion, qui p e r s i s z e n t dans des f a c i e s diachroneuz. De 1 ' a u t r e A t e , il y 2 beaucoup de groupes qui a r r i v e n t h surmonter l e s changenents de p e z i t s f a c i e s avec des changements vinimes dans l e w morphologie. 3e p l u s , l ' u t i l i z a t i o n b i o s t r a t i g r a p h i q u e des ostracodes palakozoiques e a t entrovge par l e f a i t que j u a q u ' i c i , oi: n ' a v a i t eu que des r 6 s u l t a t s isole's sur des ostracodes provenant des rdgions e t des environnements d i p & e n t s . C e t t e s i t u a t i o n p o u r r a i t s'mnZliorer dans 1 ' a v e n i r avec l e de'couvert des soi-disant faunes de'voniennes e t des faunes entomozoacdennea du Divonie n - C a r b o n i f b e . Cependent, pour l e moment, e l l e s ne sont connues qre pour l a p a r t i e cordouest de 1 'Europe, l 'Espagne, l e Portugal e t l 'Amirique du !lord. En rnzrnmo temps. des o s t r a c o d o l o g i s t e s r u s s e s f o n t des e f f o r t s p o w arr i v e r a m c o r r e l a $ i o n s de longue-portke e n t r e 1 'U.R.S.S. e t 1 'Europe de 1 ' Guest pour l e s assemblages du Divonien SupGrieur e t du Dinantien. En Europe, j u s q u 'ici i 'inte'r&?t des recherche6 sur l e s ostracodes du D6vonien jusqu'au P e n i e n a e'te' place' 6w l a p a l e b n t o l o g i e d e s c r i p t i v e e t d i n s l e s diz dernidres anne'es, sur l a palebe'cologie.
INTRODUCTION T h e r e e x i s t s a n e x t e n s i v e l i t e r a t u r e on l a t e P a l e o z o i c ost r a c o d e s from b o t h N o r t h America and w e s t e r n E u r o p e .
Amongst
t h e many p a p e r s w e may q u o t e t h e work by JONES AND KIRBY o n Eur o p e a n o s t r a c o d e s i n t h e p a s t c e n t u r y , and t h a t of U L R I C H and BASSLER $t t h e end of t h e 1 9 t h and b e g i n n i n g of t h e 2 0 t h c e n t u r y , who i n v e s t i g a t e d t h e N o r t h American f a u n a s .
Their funda-
m e n t a l s t u d i e s h a v e b e e n c o n t i n u e d by a l a r g e number of o s t r a code w o r k e r s s i n c e t h e F i r s t World War.
62
I n E u r o p e , l a t e P a l e o z o i c o s t r a c o d o l o g i s t s h a v e b e e n esp e c i a l l y a c t i v e i n Germany a n d s u r r o u n d i n g a r e a s .
It was
t h r o u g h t h e i r i n v e s t i g a t i o n s , t h a t a f i r s t a t t e m p t w a s made t o u s e entomozoan and s i l i c i f i e d o s t r a c o d e f a u n a s f o r b i o s t r a t i graphical purposes.
The p a s t t e n y e a r s h a v e moreover shown a
growing i n t e r e s t i n t h e p a l a e o e c o l o g y o f t h e s e o s t r a c o d e s . I n N o r t h America, work on Devonian t o P e r m i a n o s t r a c o d e s h a s b e e n m a i n l y c o n c e n t r a t e d on d e s c r i p t i v e p a l a e o n t o l o g y r a t h e r t h a n on b i o s t r a t i g r a p h y , a l t h o u g h i n t h e l a s t d e c a d e t h e r e i s a t e n d e n c y t o r e v i e w t h e i r v a l u e f o r i n f r a - and i n t e r b a s i n a l
correlations. I n comparison w i t h post-Paleozoic o s t r a c o d e s t h e s t u d y of P a l e o z o i c o s t r a c o d e s h a s b e e n l e s s i n t e n s i v e on t h e w h o l e as shown by t h e p a p e r s r e a d d u r i n g t h e p a s t f i v e o s t r a c o d e sympos i a a t N a p l e s , H u l l , P a u , Newark, and Hamburg.
I t should be
k e p t i n mind of c o u r s e , t h a t 1 2 P a l e o z o i c o s t r a c o d e w o r k e r s m e t s e p a r a t e l y a f t e r t h e Hamburg symposium a t G o t l a n d , w h e r e e i g h t non-published progress r e p o r t s have been discussed.
It is also
c u r i o u s t o n o t i c e how s e v e r a l y o u n g e r d i s c i p l i n e s w i t h i n t h e f i e l d of Paleozoic micropalaeontology,
such
BS
conodont, a c r i -
t a r c h s , c h i t i n o z o a n s , palynolomorphs and f o r a m i n i f e r s have y i e l d e d world-wide a p p l i c a b l e c o r r e l a t i o n s , w h e r e a s t h e b i o s t r a t i g r a p h i c v a l u e of o s t r a c o d e s i s s t i l l i n d i s c u s s i o n . I n t h i s c o n t e x t it appears t o be impossible t o p r e s e n t a t t h i s moment u n i f o r m b i o s t r a t i g r a p h i c o s t r a c o d e r a n g e c h a r t s which c a n b e u s e d on b o t h s i d e s o f t h e A t l a n t i c .
LOWE2 DEVONIAN Canada and A l a s k a Lower Devonian t o M i s s i s s i p p i a n o s t r a c o d e s i n Canada were r e v i e w e d by COPELAND (1972). G e d i n n i a n a s s e m b l a g e s , c o n t a i n i n g s e v e r a l t h l i p s u r i d and b e y r i c h i i d genera and P o l o n i e l l a s p e c i e s have been d e s c r i b e d from Quebec (BERDAN (COPELAND, 1 9 6 2 ) .
&
COPELAND
&I
BURK,
1 9 6 5 ) a n d N e w Brunswick
Upper Emsian o s t r a c o d e s , i n c l u d i n g many
b e y r i c h i i d s and t h e f i r s t d i v e r s i f i e d hollinomorphs have been d i s c o v e r e d i n A l a s k a a n d Yukon T e r r i t o r y (BERDAN 1973).
&
COPELAND,
The l a t t e r f a u n a a l s o c o n t a i n s s e v e r a l g e n e r a which a r e
known from t h e Lower Devonian o f E u r o p e , s u c h a s A c a n t h o s c a p h a ,
63
Tricornina, Beecherelia, Berounella, Praepilatina and Poloniella. U.S.A. Gedinnian faunas have been described by among others LUNDIN (1968) from Oklahoma and Tennessee (cf. also WILSON, 1935) with many thlipsurid species and Poloniella. Emsian thlipsurid faunas have been described from Tennessee (BASSLER, 1941) and Siegenian thlipsurid faunas have been described from West Virginia (ULRICH and BASSLER, 1913) and Pennsylvania (SWART2, 1932) Germany Czechoslovakia When JORDAN (1970) reviewed the Paleozoic ostracodes of Central Europe, he listed several species of Tricornina, Acanthoscapha and hollin3morphs from the Siegenian-Emsian of the German Democratic Republic, including an unidentified species of Polyzygia from the Emsian. BECKER & BLESS 11974) considered several species of Zygobeyrichia, Kozlowskiella, Polon+ella, Carinokloedenia and Bassleratia to have some biostratigraphical value for a regional subdivision of Emsian strata in the Rhenisch Mountains. Kozlowskiella also occurs the Upper Emsian of Czechoslovakia (PRIBYL, 1962). _ I
.
Acantoacopha Beccherella
62
Plo9ionephrodss
Bcrounellidr
0
Polonleila
Bcyrlchlocco ather than
@3
PoIyZygIo
treposelllmd*
a
Corboniia group
sI
Prmiopsocean Processobaiidm
Entomomids
,
i i
Fig. 1.
a
Kirkbyoccon
p~
Kloedenellacem
?
4
Rectonar (doe
a
Thlipsurids
w
Traporellinids
@
Tricirninidae
W
Hollinomorphs
0
Heaidlacean
J \
Mouryello
Schematic distribution of some ostracod groups in Lower Devonian strata
64
France The b e y r i c h i i d o s t r a c o d e s which have b e e n d e s c r i b e d from t h e S i e g e n i a n of Normandy (WEYANT, 1 9 6 5 , 1 9 6 6 ) t o g e t h e r w i t h a t h l i p s u r i d s p e c i e s and a d i v e r s i f i e d a s s e m b l a g e of P o l j z y g i a show l i t t l e i n common w i t h e i t h e r N o r t h America ( e x c e p t f o r a l o c a l s p e c i e s of O c t o n a r i a ) o r German f a u n a s . Carnic A l p s BANDEL & BECKER,
1 9 7 5 , s t u d i e d s i l i c i f i e d o s t r a c o d e s from
t h e Upper S i l u r i a n t o Lower C a r b o n i f e r o u s of t h e C e n t r a l C a r n i c Alps.
The f a u n a s a r e c h a r a c t e r i z e d by t h e predominance of
d i s t i n c t , r e l a t i v e l y few and l o n g - r a n g i n g s y s t e m a t i c u n i t s . Spain MICHEL ( 1 9 7 2 ) d e s c r i b e d f i v e s p e c i e s of P o l y z y g i a from
S i e g e n i a n and Emsian o f n o r t h w e s t e r n S p a i n . Conclusions B e y r i c h i i d o s t r a c o d s s . 1. and P o l o n i e l l a s . 1. a r e known from b o t h s i d e s o f t h e A t l a n t i c , a l t h o u g h no c l e a r r e l a t i o n s h i p s even a t g e n e r i c l e v e l c a n b e o b s e r v e d .
Thlipsurid ostra-
c o d e s e x c e p t P o l y z y g i a a p p e a r t o be w i d e s p r e a d i n t h e Lower Devonian o f N o r t h America b u t t h e y t e n d t o become r a r e i n t h e Emsian. T h i s g r o u p of o s t r a c o d e s i s p r a c t i c a l l y a b s e n t i n w e s t e r n Europe, where o n l y one s p e c i e s h a s b e e n d e s c r i b e d from Normandy. I n w e s t e r n Europe t h e t h l i p s u r i d genus P o l y z y g i a i s a b u n d a n t i n Normandy and S p a i n w i t h one s p e c i e s o c c u r r i n g i n
German Democratic R e p u b l i c .
-'e r e l l a
Genera l i k e A c a n t h o s c a p h a , Beech-
T r i c o r n i n a , and P r a e p i l a t i n a , which seem t o be r e s t r i c t -
ed t o r a t h e r q u i e t , low e n e r g y f a c i e s ( c f . J O R D A N , 1 9 7 0 ) o c c u r a t s e v e r a l p l a c e s i n Lower Devonian s t r a t a where t h e e n v i r o n m e n t a l c o n d i t i o n s a p p e a r t o have b e e n f a v o r a b l e f o r s u c h assemblages.
They c o n t a i n few marker s p e c i e s i n g e n e r a l and s e v e r a l
of t h e g e n e r a r a n g e i n t o Upper Devonian and D i n a n t i a n .
MIDDLE DEVONIAN Canada The E i f e l i a n of Eilesmere I s l a n d h a s y i e l d e d s e v e r a l s p e c i e s of P o l o n i e l l a
(WEYANT,
1968).
BRAUN,
1966 described
G i v e t i a n a s s e m b l a g e s from t h e N o r t h w e s t T e r r i t o r i e s w i t h p r i m i t s i o p s i d and q u a s i l l i t i d forms and P o l o n i e l l a .
McGILL
(1967)
d e s c r i b e d p o o r l y d i a g n o s t i c o s t r a c o d f a u n a s from t h e G i v e t i a n of t h e Northwest T e r r i t o r i e s , which h e compared w i t h f a u n a s of
65
similar age in Germany. McGILL 21966) discovered primitiopsid ostracodes from the Givetian of Alberta, U.S .A. Rich and diversified hollinomorph assemblages have been described from the Lower Eifelian of Ohio (KESLING & PETERSON, 1958), the Givetian of Michigan (KESLING & TABOR, 1953; KESLING, 1952; KESLING & McMILLAK, 1952) and the Givetian of Ontarie (KESLING, 1952). The Givetian of Ontario has also yielded quasillitid ostracodes and several species of Poloniella (cf. STUMM & WRIGHT, 1958) , Treposellinid (Hibbardia), quasillitid and thlipsurid ostracodes have been found in the Givetian of New York (SWARTZ & ORIEL, 1948). Thlipsurid and hollinomorph ostracodes from the Eifelian of Pennsylvania have been described by SWARTZ & SWAIN (1941). KESLING & KILGORE (1952) and KESLING & WEIS (1953), discovered thlipsurid and quasillitids from the Givetian of Michigan. KESLING (1954) described Poloniella from the Upper Eifelian of Ohio. ULRICH (1891) and KESLING (1955) described treposellinid species from the Eifelian of Kentucky and the Givetian of New York, respectively, Germany- & Poland Endemic black shale faunas of Upper Eifelian age occur in Thuringia with high percentages of Tricornina, Hlubocepina and healdiid ostracodes (K. ZAGORA, 1967). Significant markers are the genera Jenningsina and Poloniella. Silicified ostracode assemblages of presumably low-energy environments with Acanthoscapha, Beecherella, Berounella, Tricornina, Jenningsina and several hollinomorphs occur in the Eifelian of the Harz (BLUMENSTENGEL, 1969) and Vogtland (JORDAN, 1965). Treposellinid (Kozlowskiella, Parakozlowskiella) , hollinomorphs, quasillitid,primitiopsid and Polyzygia appear to be important regional guides in the Eifelian of Poland (ADAMCZAK, 1968) and the Eifelian-Givetian boundary of the Rhenish Mountains (BECKER, 1964, 1965, 1969, 1970, GROOS, 1969, KROMMELBEIN, 1950, 1953, 1955, BECKER & BLESS, 1974). Czechoslovakia POKORNY (1951) described some ostracodes occurring in Middle Devonian of Czechoslovakia, among others fuesillitids, hollinomorphs and primitiopsids.
-
66 (1
MIDDLE DEVONIAN
BULTIYNCK (1967) cited Polyzygia from the Eifelian of the Dinant Basin, Spain The genus Polyzygia is widely distributed in Eifelian and Givetian strata of NW Spain (MICHEL, 1972). Primitsiopsid, hollinomorphs and quasillitid ostracodes together with the genera Acanthoscapha, Tricornina, Berounella, Poloniella and Praepilatina occur in the Eifelian of NW Spain (BECKER & SANCHEZ DE POSADA, and BECKER et al., in Press). Conclusions Treposellinid ostracodes are distinctive elements from the Middle Devonian in both North America and Europe, As in the case of the Lower Devonian thlipsurid genera, treposellinid genera appear to exclude each other on both sides of the Atlantic. Hibbardia and Treposella being restricted to North America, whereas Kozlowskiella and Parakozlowskiella occur in Europe (Germany and Poland) and Australia (PRIBYL, 1962). It should be noted, however, that it is not clear to me, whether Hibbardia appears to possess a more distinct dolonoid scar than
67
Parakozlowskiella. Whether this phenomenom should be considered to be a specific or generic character remains open for discussion. Poloniella s.1. (including Dizyglopeura and Framella as subgenera) and quasillitids are characteristic of many Middle Devonian strata, However species are usually restricted to a single basin and do not permit long distance correlations. The same is true of hollinomorphs, which according to BECKER & BLESS (1971) appear to have migrated in time and space from North America to Europe during the Devonian and Carboniferous, the different genera showing a clear diachronism. The low energy environments containing Tricornina, Acanthoscapha, Berounella, Beecherella, have not been described in North America. The similarity at the specific level between certain German and Spanish Lower Fifelian faunas may be due to similar facies conditions hence their stratigraphical use remains uncertain.
UPPER DEVONIAN Canada Large Frasnian ostracode faunas from Alberta and Northwest Territories containing amongst others several species of gionephrodes, and quasillitids (Eriella, Quasillites! seem to have some similarities with Russian Platform faunas of similar age. LORANGER, 1954, 1963, 1965; McGILL, 1963; BRAUN, 1963, 1968; LORANGER, 1971, describes a Frasnian fauna containing amongst others several entomozoan ostracods which may permit comparison with entomozoan faunas from NW Europe. The apparent absence of hollinomorphs is remarkable. U.S.A. GIBSON (1955) described a fauna from the Cerro Gordo Formation of Iowa containing primitiopsid ostracodes. STEWART & HENDRIX (1945) described rather heterogeneous ostracode assemblages containing some entomozoan species from the Frasnian of Ohio. KESLING & PLOCH, (1960) studied a probably endemic cypridinacean species from the Upper Devonian Black Limestone (Frasnian/Famennian). The absence of hollinomorphs is again remarkable. Germany and Poland BLUMENSTENGEL (1965) described a rich Frasnian/Famennian
e-
68
silicified fauna from lhuringia, with hollinomorphs, Acanthoscapha, Tricornina, Berounella, Processobairdia among others. This fauna has shown to be useful for regional zonation. The Frasnian fauna from the Harz described by BLUMENSTENGEL (1969) also contains Acanthoscapha, Berounella, Tricornina and hollinomorphs but there is very little similarity between the two faunas at specific level. However GRUNDEL (1962) BLUMENSTENGEL (1954) described a rich entomozoan fauna from Thuringia which can be directly compared with the Frasnian/Famennian entomozoan faunas from the Rhenisch Mountains (RABIEN, 1954). KRCMMELBEIN (1954); BECKER (1967, 1968 , 1970) GROOS, (1969), have described rich and diversified ostracod assemblages from the Frasnian of the Rhenisch Mountains which contained several species of Polyzygia, Plagionephrodes, Primitiopsacea, Poloniella, and hollinomorphs. Entomozoan ostracodes also appear in the Upper Devonian of Poland (cf. JORDAN, 1970). Papers by RABIEN, (especially RABIEN, 1954) represent an important attempt to obtain a zonation with entomozoan ostracods and this zonation with minor modifications seems to be mainly accepted in Europe. Fig. 7 shows the Upper Devonian and lowermost Carboniferous entomozoan zonation after BECKER & BLESS (1974). The Richterina ( g . ) aff. latior Zone is proposed by BECKER & BLESS (op. cit.) and characterized by the presence of ostracodes described as 5. ( 5 . ) aff. latior by GRUNDEL (1961, 1963) JORDAN & BLESS (1970) and SANCHEZ DE POSADA & BLESS (1974). This zone is supposed to be equivalent to the Middle-Upper Tournaisian and part of the Visean. France and Belgium -LETHIERS (1970, 1972, 1974) described ostracode faunas ranging from Frasnian to Tournaisian age containing, amongst others, Polyzygia,- Quasillitidae, and several entomozoan species. BECKER & BLESS (1974) noticed that the stratigraphical range of some of the species described by LETHIERS does not coincide with that observed by themselves suggesting that Upper Famennian benthonic ostracodes of the Dinant Basin had been clearly facies controlled in their lateral and vertical distribution.
69
LETHIERS (1974b) proposed a preliminary zonation for ostracodes from Frasnian to lowermost Tournaisian in the Dinant Basin. However, as ye this zonation has not been tested outside of the area in which it was established. Sjain Little is known about Upper Devonian ostracodes in Spain. BLESS & MICHEL (1967 studied a silicified ostracod fauna from northwestern Spain containing among others Acratia,-Bohemina, Ceratacratia, Processobairdia, Ectoplacera and Tricornina. This fauna has been compared with the fauna described by BLUMENSTENGEL (1965) from the Upper Devonian in Thuringia, the age established from the ostracodes is in agreement with that obtained from conodonts and cephalopods. In addition MICHEL (1972) has described 2 species of Polyzygia in Frasnian strata of northwestern Spain. Conclusions Upper Devonian entomozoan ostracode faunas seems to be very useful for stratigraphical purposes, at least in Europe, and entomozoan zonation may be correlated with the orthochronological scale. Benthonic ostracodes are occasionally useful in regional correlations. LOWER C A R B O N I F E R O U S E U RO PE A N FA U N A S AND M I S S I S S I P P I A N AMERICAN FAUNAS
Recent progress in correlation techniques in Carboniferous strata of Europe and North America coupled with the difficulty of not knowing the exact stratigraphical level from which some ostracode faunas (described many years ago) have been obtained, has necessitated that Chester ostracodes from North America should be treated together with Lower Carboniferous European faunas
.
Canada GREEN (1963) described rich and diversified ostracodes assemblages from the Banff Formation of Alberta containing among others hollinomorphs, Kirkbyacea, Beyrichiopsidae, Kloedenellacea, Healdiidae, Paraparchitacea, rich quasillitids, some Berounellidae, Beecherellidae, Bythocytheridae and Entomozoidae. LORANGER (1958) divided the Rundle Formation into a
70
Lower Criboconcha zone and an upper Paraparchites carbonarius zone. Other Mississippian ostracodes in Canada were described by COPELAND (1957), BELL (1960) and BLESS & JORDAN (1971). U.S.A. Carboniferous and Permian ostracodes described in U.S.A. wete compiled by ECHOLS & CREATH, 1959. Most of the known Carboniferous occurrences are in the interior region outside the area of this paper.
Great Britain JONES ( 1 8 8 4 ) and JONES & KIRKBY k 8 7 4 ) described in many publications a great number of ostracodes in the British Isles covering the whole Carboniferous at the end of last century. Most of these species need revision now. As far as we know only a few of the lower Carboniferous species have been restudied. ANDERSON (1970) dealt with species of the genus Carbonita.
DEVONIAN
Fig. 3 Schematic distribution of some ostracode groups in Upper Devonian strata.
71
ROBINSON (1959) described an ostracod fauna from the Cowdor Quarry in Derbyshire containing among others Tetrasac-culus 1 Kirkbyacea, Discoidella, Paraparchitacea, Monoceratina and Healdicea. Germany KUMMEROW 1939) described 80 species from Lower Carboniferous strata of Germany and Belgium containing among others Hollinellidae Kirkbyacea, Bairdiacea, Berounellidae, Glyptopleuridae and Entomozoidae. GRUNDEL (1961) described more than 60 species from the Gattendorfia-stufe. The faunas are mainly composed of Tricorninidae, Kirkbyacea, Healdiidae, Rectonariidae, Bairdiidae, Cypridinadae, Monoceratina, Triceratina and Entomozoidae. Ostracod faunas described by GRUNDEL (1962) from the CU 11 from the southern border of the Ruhr Carboniferous B/Y area contained mainly Rectonariidae, Tricornina, Glyptopleuridae, Healdiopsis and Entomozoidae. Two species, Glyptopleura elapa and Richterina (R.) aff. latior a r e related to species described in 1974 by SANCHEZ DE POSADA & BLESS from Aprath (Federal Republic of Germany). BECKER & BLESS (1974) reviewed the stratigraphical range of the species described from the Lower Carboniferous of Dinant Basin by DE KONINCK. (1840, 1844) ,JONES & KIRKBY (1874), TSCHIGOVA (1970), ROME (1971) and ROME & GOREUX (1960). Some of the species named under open nomenclature are figured in the above papers; some of them are compared with previously described species, and their stratigraphical range has been proposed. Nine ostracod zones ranging from Fa2a to V3 have been proposed. Belaium ROME & GOREUX (1960) described nine species of Cryptophyllus from Strunian strata. ROME (1971) described several species of Kegelites, Beyrichiopsis, Sansabella, Bairdia, Silenites, Microcheilinella, and Paraparchites. Some of the species of Bairdia described by ROME are considered to belong to a single species (LETHIERS, 1975). BECKER & BLESS (1974) studied ostracodes in the Dinant Basin of Germany and Belgium (see above). BECKER & BLESS (1974) figured and described some species occurring in Fr2 strata in the Belgian part of the Dinant Basin. Special attention is
12
p a i d t o B e y r i c h i o p s i s g l y p t o p l e u r o i d e s GREEN and t o a g r o u p of s i m i l a r ( i f n o t i d e n t i c a l ) s p e c i e s composed o f " B e r n i x " v e n u l o s a KUMMEROW
( 1 9 3 9 ) , P s e u d o l e p e r d i t i a p o o l e i SOHN ( 1 9 6 9 ) and
p.
t u b e r c u l i f e r a SCHNEIDER (1956) a l l of which a r e c o n s i d e r e d t o p r o b a b l y have some v a l u e i n l o n g - d i s t a n c e c o r r e l a t i o n s .
g.
g l y p t o p l e u r o i d e s i s known from t h e uppermost Famennian and lower most T o u r n a i s i a n s t r a t a i n t h e USSR, Europe and Canada. The g r o u p of P,. p o o l e i ,
g. t u b e r c u l i f e r a and " B e r n i x " v e n u l o s a
i s known from D i n a n t i a n s t r a t a i n U S A , t h e F e d e r a l R e p u b l i c of Germany and USSR. BLESS & THOREZ (& BECKER e t a l . , 1 9 7 4 ) d i s t i n g u i s h e d f o u r o s t r a c o d e s a s s e m b l a g e s , two o f which o c c u r i n l i m e s t o n e b e l i e v e d t o r e p r e s e n t a r e l a t i v e l y d e e p s u b t i d a l environment, t h e o t h e r occurred i n limestones b e l i e v e d t o have been d e p o s i t e d i n a n i n t e r t i d a l and a s u p r a t i d a l e n v i r o n m e n t . Poland BLASZYK & NATUSIEWICZ ( 1 9 7 4 ) d e s c r i b e d 3 4 o s t r a c o d e s p e c i e s from D i n a n t i a n and Namurian s t r a t a i n n o r t h w e s t e r n P o l a n d .
Only one s p e c i e s i s r e l a t e d t o p r e v i o u s l y d e s c r i b e d s p e c i e s from Europe, f o u r a r e i d e n t i c a l and two are r e l a t e d t o American species. Spain JORDAN & BLESS ( 1 9 7 0 ) d e s c r i b e d two s p e c i e s of entomozoid
O s t r a c o d a o c c u r r i n g i n T o u r n a i s i a n s t r a t a of n o r t h w e s t e r n S p a i n and f i g u r e d s e v e r a l s p e c i e s of K i r k b y a , A c r a t i a and T r i p l a c e r a . Conclusions Entomozoan O s t r a c o d a seem t o be u s e f u l f o r s t r a t i g r a p h i c a l p u r p o s e s i n some Lower C a r b o n i f e r o u s s t r a t a from Europe b u t a l o t of work h a s s t i l l t o be done c o n c e r n i n g t h i s s u b j e c t . BECKER & BLESS
(1974)
( i n BECKER
g . )cited
several
s p e c i e s of B e r n i x , B e y r i c h i o p s i s and P s e u d o l e p e r d i t i a a s p o s s i b l e index f o s s i l s f o r long d i s t a n c e c o r r e l a t i o n s . A most i n t e r e s t i n g a s p e c t o f t h e o s t r a c o & f a u n a on b o t h
s i d e s of t h e A t l a n t i c i s t h e a b s e n c e i n N o r t h America of T r i c o r n i n i d a e and R e c t o n a r i i d a e .
On t h e o t h e r hand t h e most
s t u d i e d American Lower C a r b o n i f e r o u s f a u n a s p o s s e s s a g r e a t e r v a r i e t y of s p e c i e s of h o l l i n o m o r p h s , K i r k b y a c e a , B a i r d i a c e a and H e a l d i a c e a .
73 UPPER CARBONIFEROUS
U.S.A. Most of t h e p a p e r s o n P e n n s y l v a n i a n o s t r a c o d e s d e a l w i t h faunas r e s t r i c t e d e i t h e r s t r a t i g r a p h i c a l l y o r g e o g r a p h i c a l l y . COOPER ( 1 9 4 6 ) , BRADFIELD ( 1 9 3 5 ) , and KELLETT
(1933, 1934, 19351,
however, d e s c r i b e l a r g e and d i v e r s i f i e d f a u n a s from t h r o u g h o u t t h e P e n n s y l v a n i a n of I l l i n o i s and a d j a c e n t a r e a s o f I n d i a n a and Kentucky, t h e Ardmore B a s i n i n Oklahoma and t h e Upper P e n n s y l vanian i n t o Permian of Kansas, a l l of which are o u t s i d e t h e a r e a of t h i s s t u d y .
T h e s e f a u n a s a r e m a i n l y composed o f H o l l i -
n e l l a , Kloedenellacea, Kirkbyacea, Paraparchitacea, Bairdiacea, and H e a l d i a c e a .
L i t t l e can be s a i d about t h e u s e f u l n e s s of t h e
d e s c r i b e d s p e c i e s f o r s t r a t i g r a p h i c a l p u r p o s e s as m o s t o f them
seem t o h a v e a r a t h e r l o n g s t r a t i g r a p h i c a l r a n g e and o n l y a c o m p a r a t i v e l y s m a l l number h a v e a r e s t r i c t e d r a n g e ( c f . C O O P E R , 1 9 4 6 , p . 18; KELLETT, 1 9 3 3 , p . 6 1 ) . SHAVER,
( i n Thomson e t a l . , 1 9 5 9 ) d e s c r i b e d a n o s t r a c o d e
f a u n a from K e n t u c k y , B a i r d i a c e a and K i r k b y a c e a b e i n g t h e p r e d o m i n a n t e l e m e n t s t h e r e i n . SHAVER & SMITH ( 1 9 7 4 ) d e s c r i b e d 1 5 s p e c i e s o f K i r k b y a c e a from Lower and M i d d l e P e n n s y l v a n i a n s t r a t a of I n d i a n a and K e n t u c k y .
The A_. r o t h i f a u n a i s f o u n d i n
rocks belonging t o t h e P r o f u s u l i n e l l a zone whereas t h e
5.
c e n t r o n o t u s f a u n a i s f o u n d i n r o c k s of t h e z o n e s o f F u s u l i n e l l a and F u s u l i n a . Lower P e n n s y l v a n i a n non-marine o s t r a c o d e s from t h e S . A p p a l a c h i a n s were d e s c r i b e d by SCOTT
&
SUMMERSON ( 1 9 4 4 ) .
They
included a s p e c i e s , Cypridina r a d i a t a ( t y p e s p e c i e s of Radiic y p r i d i n a BLESS, 1 9 7 3 ) , w h i c h i s common i n t h e Lower Westp h a l i a n of Europe.
74
MISSISSIPPIAN
ii
LOWER CARBONIFEROUS
Y
Fig. 4 Schematic distribution of some ostracode groups 4.n Lower Carboniferous and Mississippian strata.
'ENNSYLVANIAN
UPPER
CARBONIFEROUS
/,
/
"
/
I' \
.i
,
Fiq. 5 Schematic distribution of some ostracode groups in Uppe; Carboniferous and Pennsylvanian strata.
75 U P P E R C A R B O N I F E R O U S OF E U R O P E
Great Britain JONES and JONES & KIRKBY described in the last century some ostracodes occurring in the Upper Carboniferous of Great Britain, most of these are species of Geisina and Carbonita. As stated before species of Carbonita were reviewed by ANDERSON (1970). POLLARD (1966, 1969) described and noted the occurrence of several Geisina and Carbonita species in non-marine bands from the Coal Measures of Durham and Northumberland. BLESS & POLLARD (1973) studied the palaeocology and content of two ostracodeassemblages, one from Upper Westphalian A strata in The Netherlands, the other one from Lower Westphalian B in Great Britain. Only a small number of marine ostracodespecies are known from the Upper Carboniferous of Great Britain. RAMSBOTTOM, 1952 described some kirkbyacean and Roundyella species from the Similis pulchra zone. BLESS (1974) described the occurrence of several species of Hollinellidae (Hollinella and Jordanites) , Kirkbyacea, Roundyella, Moorites, Cornigella, Pseudoparaparchites, Healdia and Asturiella from the Croft's End Marine Band. Eleven of these species are conspecific or closely related to species of Pennsylvanian age in North America. Germany KUMMEROW (1953) described Upper Carboniferous ostracodes from Germany containing, among others, Paraparchitacea, Kellettina, and Kloedenellacea. Now the fauna needs to be restudied. KREMP & GREBE (1956), VANGEROW (1958, 1970), described nonmarine ostracodes mainly Kloedenellacea and Carbonita species. The Netherlands VAN DER HEIDE (1951) describes species of Carbonita, Geisina, and Qpridina in the Upper Carboniferous of The Netherlands. BLESS, JORDAN & MICHEL (1969) described a rather diversified ostracode fauna from the base of Westphalian C. This included: Hollinellidae, Kirkbyacea, Cornigella, Roundyella, Morrites, Paraparchitacea and Healdiacea. Twelve of these species are conspecific with species described by BLESS (1974) from the same horizon in Great Britain. BLESS (1974) described the new genus Cypridelliforma from the Lower Westphalian A of the Netherlands.
-
76
Spain BLESS (1965, 1967, 1968, 1970 - in VAN AMEROM g.), SANCHEZ DE POSADA & BLESS (1971), BECKER & BLESS (in BECKER g a1.,1375 ) described more than 40 species of Upper Carboniferous ostracodes in northwestern Spain (mostly in Upper Westphalian strata with only a small number from Namurian and Stephanian rocks). Most of the species which had been previously described from elsewhere are identical or closely related to Upper Pennsylvanian species from North America. The faunas are mainly composed of Hollinellidae (Hollinella and Jordanites) , Kirkbyacea, Roundyella, Bairdiacea and Healdiacea. However two genera, Jordanites and Asturiella do not occur in North America. Unpublished data of several localities in northwestern Spain show that some species which occur at the base of Westphalian C in The Netherlands and England are now known from the Westphalian A-B of northwestern Spain: Healdia sp. D (BLESS, JORDAN & MICHEL 1969) , Asturiella limburgensis BLESS, Pseudobythocypris pediformis (KNIGHT) and Rouridyella simplicissima (KNIGHT) Cavellina sp. cf. cumingsi described by BLESS AND by SANCHEZ DE POSADA & BLESS (1971) previously described from Upper Westphalian strata only in Spain has now also been found in Lower Westphalian rocks. A most interesting aspect of the Spanish Upper Carboniferous ostracodefauna is the presence of some ostracodes previously not known from so young strata as Upper Carboniferous, mainly Tricornidae (Tricornina (Sricorninq) and (Bohemina) and Rectonariidae (at least one probable species of Rectoplacera occurs), and Berounella. The occurrence of such ostracodes is believed to have a clear environmental control. All these o s tracods were found in limestones with silicified fauna or in shales (as internal and external molds) in sections without coal bearing strata. The same holds true for a silicified fauna composed of Myodocopida in limestones of Westphalian D age and for the entomozoan ostracode Truyolsina truyolsi described by BECKER & BLESS (in BECKER et a1.1974 ) from rocks of Namurian A age. Czechoslovakia PRIBYL (1958, 1962) described 4 3 ostracode species from the Namurian A of Czechoslovakia, containing amonc; others
.:
77
Carbonita, Hollinella, Healdiacea, Kloedenellidae, and Kirkbyacea. These bear a close relationship to American faunas from Chester and Morrow strata (cf. PRIBYL, 1962, p. 6). Conclusions Possibly due to the greater development of marine sequences in North America, Upper Carboniferous ostracodes in USA are more varied than in western Europe where only a relatively small number of marine faunas has been studied. The marine ostracodes of Europe have frequently been considered to have relationships with American ones (cf. PRIBYL, 1962, BLESS, 1974). Both American and European marine bands are mainly composed of Hollinellidae, Kirkbyacea, Kloedenellacea, Paraparchitacea, Bairdiacea, and Healdiacea. Some of the species are identical or closely related on both sides of the Atlantic. In Europe work was mainly done with non-marine ostracodes. Carbonita and Geisina are the more frequent genera. The presence in Upper Carboniferous strata of northwestern Spain of some ostracodes belonging to the Families Fricorninidae and Rectonariidae is remarkable.
-
PERMIAN
U.S.A.
HARRIS & LALICKER (1932) described Wolfcampian ostracodes from Oklahoma and Arkansas containing Paraparchitacea, Healdiacea and Kloedenellacea. Diversified assemblages of Hollinellidae, Kirkbyacea, Kloedenellacea and Bairdiacea were described from Wolfcampian strata of Kansas by KELLETT (1933, 1934, 1935) Wolfcampian and Lower Leonardian from Nebraska by UPSON (1935). Other Wolfcampian faunas were described by DELO (1930) and HARRIS & WORELL (1936). All of these are outside the present study area. HOLLAND (1934) and SCOTT (1942) studied unique non-marine species from West Virginia,
Fig. 6
Schematic distribution of some ostracode groups in
Permian strata.
79 England Permian o s t r a c o d e s from England were d e s c r i b e d i n t h e l a s t c e n t u r y by K I R K B Y . Germany
&
K i r k b y a c e a a r e t h e most r e m a r k a b l e e l e m e n t s .
Poland
Lower Z e c h s t e i n o s t r a c o d e s from Pommern h a v e b e e n des c r i b e d by KRb'MMELBEIN ( 1 9 5 8 ) .
The f a u n a c o n t a i n s s e v e r a l s p e -
c i e s of M o n o c e r a t i n a , R o u n d y e l l a , H e a l d i a c e a and B a i r d i a c e a , most of which had n o t b e e n p r e v i o u s l y d e s c r i b e d .
RICHTER and
REUSS d e s c r i b e d i n t h e 1 9 t h c e n t u r y o s t r a c o d f a u n a s from t h e
Z e c h s t e i n of Germany and i n 1968 J O R D A N r e v i e w e d t h e P e r m i a n o s t r a c o d e s d e s c r i b e d from t h e Z e c h s t e i n i n c e n t r a l E u r o p e .
REFERENCES Due t o r e a s o n of s p a c e t h e r e a d e r i s r e f e r r e d t o BECKER & BLESS,
1 9 7 4 and ECHOLS & CREATH,
1959 f o r m o s t r e f e r e n c e s d e a l -
ing w i t h o s t r a c o d e s d e s c r i b e d from Devonian and D i n a n t i a n s t r a -
t a of C e n t r a l and W e s t e r n Europe a s w e l l a s M i s s i s s i p p i a n , P e n n s y l v a n i a n and Permian s t r a t a of U.S.A. Amerom, H . W . J . v a n , B l e s s , M . J. M . , and W i n k l e r , P r i n s C . F . , 1 9 7 0 . Some p a l e o n t o l o g i c a l and s t r a t i g r a p h i c a l a s p e c t s of t h e Upper C a r b o n i f e r o u s Sama F o r m a t i o n ( A s t u r i a s , S p a i n ) . Med. R i j k s G e o l . D i e n s t , N.S., n . 2 1 , p p . 9 - 7 9 , p l s . 1 - 1 0 . Anderson, F . r 1 9 f 0 . a r b o n i f e r o u s O s t r a c o d a . The g e n u s C a r b o n i t a S t r a n d . B u l l . Geol. S u r v . G r e a t B r i t a i n , n . 32, DD. 69-121. ~ l s X . II-XIX. B a i i e l , D . , an& B e c k e r , G . , 1975. O s t r a c o d e n a u s p a l s o z o i s c h e n p e l a g i s c h e n K a l k e n d e r K a r n i s c h e n Alpen ( S i l u r i u m b i s U n t e r k a r b o n ) . Senck. l e t h . , Band 5 6 , p p . 1 - 8 3 , p l s . 1 - 8 . 1 9 4 1 . O s t r a c o d a from t h e Devonian (Onondaga) Bassler, R. S. c h e r t of West V i r g i n i a . Washington Acad. S c . J o u r . , v . 3 1 , pp. 2 1 - 2 7 . Becker, F . , B l e s s , M . J. M. , S t r e e l , M. , and T h o r e z , J . , 1 9 7 4 . P a l y n o l o g y and o s t r a c o d e d i s t r i b u t i o n i n t h e Upper Devonian and b a s a l D i n a n t i a n of Belgium and t h e i r d e p e n d e n c e on s e d i +. R i j k s G e o l . D i e n s t , N . S . , n . 25, p p . 9mentary f a c i e s . M 98, 30 p l s . Ostracode s t r a t i g r a p h y Becker, G . , and B l e s s , M. J . M . , 1 9 7 4 . of t h e Ardenno-Rhenisch Devonian and D i n a n t i a n . I n t e r . B e l g i a n Microp. l i m i t s , Namur, 1 9 7 4 , P u b l i c a t i o n n . 1, 52 p p . , 50 p l s . Becker, F . , B l e s s , M. J . M . , and Kullmann, J . , 1975. O b e r k a r b o n i s c h e E n t o m o z o e n - S c h i e f e r i m K a n t a b r i s c h e n G e b i r g e (NordP a l a o n t . Abh. 1 5 0 , p p . 9 2 - 1 1 0 . s p a n i e n ) . 5. Becker, G . , Mendez B e d i a , I . , and Sanchez d e P o s a d a , L . C . , ( i n p r e s s ) ; Una f a u n a d e o s t r i c o d o s d e l a f o r m a c i o n M o n i e l l o (Devonico, A s t u r i a s , NW d e E s p a n a ) . Nota p r e l i m i n a r . T r a b . Geol., n . 7 . Becker G . , and S a n c h e z d e P o s a d a , L . C . ( i n p r e s s ) . Ostracoda aus d e r M o n i e l l o F o r m a t i o n A s t u r i e n s (Devon; NW S p a i n ) Palaeontographica.
,
w.
Jb.
-
e.
,
80
B e l l , W . A . , 1 9 6 0 . M i s s i s s i p p i a n H o r t o n Group o f t y p e WindsorH o r t o n d i s t r i c t , Nova S c o t i a . G e o l . S u r v . C a n a d a , M Z . 3 1 4 , 58 p p . , 2 4 p l s . B e r d a n , J . M . , and C o p e l a n d , M . J . , 1 9 7 3 . O s t r a c o d e s from Lower Devonian f o r m a t i o n s i n A l a s k a and Yukon T e r r i t o r y . % G e o l . S u r v . P r o f . , P a p e r 8 2 5 , 47 p p . , 1 4 p l s . B l e s s , M. J . M . , 1965. On two new s p e c i e s o f m a r i n e o s t r a c o d e s i n t h e C a r b o n i f e r o u s o f A s t u r i a s , S p a i n . L e i d . Geol. E . , V. 3 3 , p p . 177-182. i ' 9 6 6 . C a r b o n i t a a g n e s (JONES) f r o m t h e C o a l Mine "Sab e r o " , (Le6n ( S p a i n ) . N o t s . Corns. I n s t . G e o l . Min. E s p a n a , n . 9 0 , pp. 93-98. , 1 9 6 7 . On t h e m a r i n e b e d s o f some c y c l o t h e m s i n t h e C e n t r a l C a r b o n i f e r o u s B a s i n of A s t u r i a s w i t h s p e c i e s r e f e r ence t o t h e i r o s t r a c o d e fauna. n . 99-100, p p . 91-134. , 1 9 6 8 . On two H o l l i n i d O s t r a c o d e g e n e r a from t h e Upper v . 43, C a r b o n i f e r o u s of N o r t h w e s t e r n S p a i n . L e i d . G e o l . p p . 157-212, p l s . 1 - 1 0 . , 1 9 7 4 . On a new g e n u s and s p e c i e s o f C y p r i d i n a c e a ( O s t r a c o d a ) f r o m t h e Upper C a r b o n i f e r o u s of The N e t h e r l a n d s . Med. R i j k s G e o l . D i e n s t , N . S . , n . 2 2 , p p . 21-23, p l s . I . , 1 9 7 4 . O s t r a c o d s f r o m C r o f t ' s End M a r i n e Band ( b a s e of W e s t p h a l i a n C ) o f t h e B r i s t o l D i s t r i c t . B u l l . Geol. S u r v . Great B r i t a i n , n . 4 7 , p p . 39-53, p l s . 4-6. B l e s s , M. J . M . , and J o r d a n , H . , 1 9 7 1 . The new g e n u s C o p e l a n d e l l a from t h e Lower C a r b o n i f e r o u s t h e y o u n g e s t known beys i a c e a n o s t r a c o d e . L e t h a i a , v . 4 , pp. 185-190.
,
s.,
e.,
-
, 1 9 7 2 . O s t r a c o d e s of t h e f a m i l y H o l l i n e l l i d a e . E . R i j k s G e o l . D i e n s t , s e r . C , v . 3 , 83 p p . , 35 p l s . B l e s s , M . J . M . , J o r d a n , H . , and M i c h e l , M. P h . , 1 9 6 9 . Ostrac o d s f r o m t h e A e g i r M a r i n e Band ( b a s i s W e s t p h a l i a n C ) o f Id. n . 2 0 , pp. 1 9 - 4 9 , p l s . S o u t h Limburg (The N e t h e r l a n d s ) . 1-7. B l e s s , M. J . M . , a n d M i c h e l , M . P h . , 1 9 6 7 . An o s t r a c o d e f a u n a from t h e Upper Devonian o f t h e G i l d a r - M o n t o R e g i o n (NW S p a i n ) . L e i d . Geol:-Med. v . 3 9 , p p . 269-271. Bless, M X M T a n d P o l l a r d , J . , 1973. P a l e o e c o l o g y and O s t r a c o d e f a u n a s o f W e s t p h a l i a n O s t r a c o d e Bands from Limburg, The N e t h e r l a n d s and L a n c a s h i r e , Great B r i t a i n . M S . R i j k s Geol. D i e n s t , N.S., n. 2 4 , pp. 1-33, p l s . 1-5. S t r a t i g r a p h y a n d m i c r o f a u n a o f M i d d l e and Braun, W . K . , 1966. Upper Devonian F o r m a t i o n s , Norman Wells a r e a , N o r t h w e s t T e r r i t o r i e s . N . J b . G e o l . P a l a o n t . , Abh. 1 2 5 , pp. 2 4 7 - 2 6 4 . , 1 9 6 8 . Upper Devonian O s t r a c o d f a u n a s o f Great S l a v e Lake a n d N o r t h e a s t e r n A l b e r t a , Canada. I n t e r n a t . n i a n S y s t e m , Calgary, 1 9 6 7 , A l b e r t a A s s o c . P e t r o . G e o l . , v . I .. D D . 617-652. . 9 DX & B u r k , C . F . , 1 9 6 5 . S i l u r i a n s t r a t i g r a p h y o f Gasp; P e n i n s u l a . A Reply. A m . Assoc. P e t r . , v . 4 9 , pp. 2305-2316. C o p e l a n d , M . J . ,e.7591 Arthropod f a u n a i n t h e Legger C a r b o n i f e r o u s r o c k s of t h e M a r i t i m e P r o v i n c e s . G e o l . S u r v . Cana&, 286, 1 1 0 p p . , 2 1 p l s . , 1 9 6 2 . O s t r a c o d a f r o m t h e Lower Devonian D a l h o u s i e b e d s , n o r t h e r n New B r u n s w i c k . G e o l . S u r v . Canada, B u l l . 9 1 , p p . 18-51. , 1 9 7 4 . B i o s t r a t i g r a p h i c z o n a t i o n o f Devonian a n d M i s s i s s i p p i a n O s t r a c o d a from Canada. A summary a c c o u n t . I n t e r -
m.w-
-
L L
s.
@.
w.
81
n a t . &. Belgian n. 7 , 9 p p .
M i c r o p . l i m i t s , Namur, 1 9 7 4 .
Publication
E c h o l s , D . J . , and C r e a t h , W . B . , 1 9 5 9 . S u r v e y o f M i s s i s s i p p i a n , P e n n s y l v a n i a n and Permian o s t r a c o d a r e c o r d e d i n t h e U n i t e d S t a t e s . M i c r o p a l e o n t . , v . 5 , p p . 389-414. G i b s o n , L . B . , 1 9 5 5 . Upper Devonian O s t r a c o d a f r o m t h e C e r r o Gordo F o r m a t i o n o f Iowa. Bull. A m . P a l e o n t . , v . 3 5 , p p . 5-36, p l s . 1-2. G r e e n , R . , 1 9 6 3 . Lower M i s s i s s i p p i a n o s t r a c o d e s from t h e B a n f f F o r m a t i o n , A l b e r t a . R e s e a r c h C o u n c i l A l b e r t a , B u l l . 11, 273 pp., 1 7 pls. Heide, S . , van d e r , 1951. L e s A r t h r o p o d e s du t e r r a i n H o u i l l i e r du Limburg M g r i d i o n a l ( e x c e p t ; l e s s c o r p i o n s e t l e s i n s e c t e s ) S t i c h t i n g , s e r . C-IV-3, n . 5 , 84 p p . , 1 0 p l s . J o n e s , T . R . , and K i r k b y , J . W . , 1 8 7 4 . A monograph o f t h e B r i t i s h f o s s i l Entomostraca from t h e C a r b o n i f e r o u s F o r m a t i o n s . P a r t I . The C y p r i d i n i d a e and a l l i e d g r o u p s . P a l e o n t . Monographs, p p . 1 - 5 6 , p l s . 1-5. , 1 8 8 7 . A l i s t of t h e g e n e r a and s p e c i e s o f b i v a l v e d E n t o m o s t r a c a f o u n d i n t h e C a r b o n i f e r o u s F o r m a t i o n s of G r e a t B r i t a i n and I r e l a n d , w i t h n o t e s o n t h e g e n e r a and t h e i r d i s t r i b u t i o n . E.A s s . v . 9 , p p . 495-515. J o r d a n , H . , 1 9 6 8 . Neue t a x i n o m i s c h e und b i o s t r a t i g r a p h i s c h e E r g e b n i s s e m i k r o p a l z o n t o l o g i s c h e r U n t e r s u c h u n g e n i m germanischen Zechsteinbacken u n t e r besonderer Beriicksichtigung d e r O s t r a c o d e n . G e o l . und P a l a o n t . , Band 1 3 , p p . 1 9 9 - 2 1 3 ; . J o r d a n , H . $ and B l e s s , M . J . M ; , 1 9 7 0 . Nota p r e l i m i n a r s o b r e 10s o s t r a c o d o s d e l a f o r m a c i o n Vegamiin. Geol. A s t u r i c a , A N ~ O XIV, p p . 37-44. K e z n g , R. V . , 1952. O s t r a c o d e s o f t h e F a m i l i e s L e p e r d i t e l l i d a e , P r i m i t i i d a e , D r e p a n e l l i d a e , Aechminidae and K i r k b y i d a e from t h e M i d d l e Devonian B e l l S h a l e o f M i c h i g a n . P a l e o n t . C o n t r . M i c h i g a n Univ. Museum, v . 1 0 , p p . 2 1 - 4 4 , 5 p l s .
.
s. s.
e.
=.,
s.
-
, 1953. O s t r a c o d s of t h e f a m i l y H o l l i n i d a e from t h e v . 1 0 , p p . 203-219, p l s . 1 - 4 . Arkona S h a l e o f O n t a r i o . g., , 1 9 5 4 . O s t r a c o d s from t h e M i d d l e Devonian Dundee L i m e s t o n e i n N o r t h w e s t e r n Ohio. v . 11, pp. 167-186, 3 p l s . , 1 9 5 5 . Two new s p e c i e s o f o s t r a c o d s from t h e C e n t e r v . 1 2 , p p . 273-284, f i e l d L i m e s t o n e of W e s t e r n N e w York. 3 pls. K e s l i n g , R . V . , and K i l g o r e , J . E . , 1952. O s t r a c o d s of t h e families Leperditellidae, Drepanellidae, Glyptopleuridae, K l o e d e n e l l i d a e , B a i r d i i d a e , B a r y c h i l i n i d a e and T h l i p s u r i d a e from t h e Genshaw f o r m a t i o n o f M i c h i g a n . Id., v . 1 0 , p p . 1 - 1 9 , pls. 104. K e s l i n g , R . V . , and McMilland, G . V . , 1 9 5 1 . O s t r a c o d e s o f t h e Family H o l l i n i d a e from t h e B e l l S h a l e o f M i c h i g a n . v. 9 , pp. 45-81, p l s . 1 - 7 . K e s l i n g , R . V . , and P e t e r s o n , R . M . , 1 9 5 8 . M i d d l e Devonian o s t r a c o d s from t h e F a l l s o f Ohio. M i c r o p a l e o n t . , v . 4 , pp. 129-148, p l s . 1, 2 . K e s l i n g , R . V . , and P l o c h , R . A . , 1 9 6 0 . N e w Upper Devonian c y p r i d i n a c e a n o s t r a c o d from S o u t h e r n I n d i a n a . P a l e o n t . C o n t r . M i c h i g a n Univ. Museum, v . 1 5 , pp. 281-292, 3 p l s . K e s l i n g , R . V . , a n d T a b o r , N . L . , 1 9 5 2 . Two new s p e c i e s of o s t r a c o d s from t h e Genshaw F o r m a t i o n ( M i d d l e D e v o n i a n ) of MichP a l e o n t . , v . 2 6 , p p . 761-763, p l . 111. igan.
s.,
=.,
=.,
Jour.
82
K e s l i n a , R. V . , and Weiss. M . . 1 9 5 3 . O s t r a c o d a from t h e Norwav P o i n t ' F o r m a t i o n o f Michigan: P a l e o n t . C o n t r . Michigan Museum, v . 11, p p . 33-76, 5 p l s . Kremv. G . . a n d G r e b e , H . . 1 9 5 6 . B e s c h r e i b u n a und s t r a t i a r a D h i s c h e r Wert e i n i q e r O s t r a c o d e n f ormen a u s dGm Rhurkarbo;. G e o l . J a h r b . , Band 7 1 , p p . 145-170, p l . 1 6 . KrBmmelbein, K . , 1 9 5 8 . O s t r a c o d e n a u s a u s dem U n t e r e n ZechBand 7 5 , p p . 115-134, s t e i n d e r ' B o h i u n g Leba i n Ponunern. d., p l s . 1-3. Kummerow, E . , 1 9 5 3 . Ueber O b e r k a r b o n i s c h e und D e v o n i s c h e O s t r a c o d e n i n D e u t s c h l a n d und i n d e r B o l k r e p u b l i k P o l e n . Geolog i e B e i h . z u r Z e i t c h B e i h . 7, 7 5 p p . , 7 p l s . Latham,, m 2 m ; t i s h Carboniferous 0stracoda. E d i n b u r g h T r a n s . , v . 5 7 , p p . 351-395. L e t h i e r s , F . , 1 9 7 2 . O s t r a c o d e s f a m e n n i e n s d a n s l ' 0 u e s t du G e o l . Nord. , t . X C I I , B a s s i n du D i n a n t (Ardenne) A s . p p . 155-169, p l s . 23-25. , 1 9 7 4 . O s t r a c o d e s du p a s s a g e F r a s n i e n - F a m e n n i e n d e S e n z e l l l e s ( A r d e n n e ) Palaeontograph. , A . Band 1 4 7 , p p . 366 9 ,. v- l s . 7-9. , 1974. O s t r a c o d e s d e l a l i m i t e D 6 v o n i e n - C a r b o n i f i r e d a n s i ' A s v e n o i s . 2. E . AS..%. P a r i s , s e r . D , t . 2 7 8 , pp. 1015-1017. , 1 9 7 4 . B i o s t r a t i g r a p h i e d e s O s t r a c o d e s d a n s l e D&on i e n s u p 6 r i e u r du Nord d e l a F r a n c e e t B e l g i q u e . N e w l s . S t r a t i g r a p h . , v . 3 , p p . 73-79. , 1 9 7 5 . L e s e n t o m o z o i d 6 s ( O s t r a c o d e s ) d u F a c i e s Matagne dans l e F r a s n i e n S u d - O c c i d e n t a l d e 1'Ardenne. Geobios, n. 8 , pp. 135-138. L o r a n g e r , D. M . , 1 9 5 4 . I r e t o n m i c r o f o s s i l s z o n e s o f c e n t r a l a n d n o r t h e a s t e r n A l b e r t a ; i n C l a r k , L . M. ( e d . ) : W e s t e r n Canada Sedimentary b a s i n . Petr. Ralph Leslie R u t h e r f o r d M e m . v o l . , pp: 1 8 2 - 2 0 3 . 1 9 6 3 7 D e v o n i a n Microfauna of n o r t h e a s t e r n A l b e r t a . P t . I . O r d e r s L e p e r d i t o c o p i d a a n d , P a l a e o c o p i d a . P u b l i s h e d by t h e a u t h o r 55 p p . , 4 p l s . P t . 11. O r d e r P o d o c o p i d a . Id., 53 PP. I 3 P l S * , 1 9 6 5 . Devonian P a l e o e c o l o g y of n o r t h e a s t e r n A l b e r t a . P e t r . , v . 3 5 , p p . 818-837. , 1 9 7 1 3 s t r a c o d s . t r a c e e l e m e n t s and F r a s n i a n r e e f s i n S t u r g e o n Lake a r e a . B u l l . C e n t r e Rech. SNPA, C o l l o q u e P a l e o e c . o s t r a c . , P a u l F r a n c e , v . 5 , s u p l . , p p . 769-768. L u n d i n , R . , 1 9 6 8 . H a r a g a n O s t r a c o d e s . Oklahoma G e o l . S u r v . B u l l . 116, 1 2 1 pp., 22 p l s . M c m . P . . 1963. Uvver and M i d d l e Devonian o s t r a c o d e s f r o m t h e B e a v e r h i l l Lake- F o r m a t i o n , A l b e r t a , Canada. g .Can. P e t r . G e o l . , v . 11, p p . 1-26. , 1966. O s t r a c o d s o f p r o b a b l e L a t e G i v e t i a n a g e from S l a v e P o i n t F o r m a t i o n , A l b e r t a . I d . , v . 1 4 : p p . 104-133. , 1 9 6 7 . Comparison o f a M i a l e G i v e t i a n o s t r a c o d e f a u n a from C a r c a j o u R i d g e , N o r t h w e s t T e r r i t o r i e s , C a n a d a , w i t h s i m i l a r f a u n a s f r o m E u r o p e . I n t e r n a t . _Symp. Devonian S s t e m C a l g a r y , 1967. A l b e r t a E.P e t r . G e o l . , v . 11, p p h l 1085, 4 p i s . McGuire, 0; S . , 1 9 6 6 . P o p u l ' a t i o n s t u d i e s i n t h e o s t r a c o d e gen u s p o l y t y l i t e s f r o m t h e C h e s t e r S e r i e s . Jour. P a l e o n t . , v . 4 0 , p p . 883-910, p l s . 103-104.
w. A
--
a.e.
.
e.
.
%.Ass.
,
-
our.
s.
--
=.,
83 M i c h e l , M. P h . , 1972. P o l y z y g i a G u r i c h ( O s t r a c o d a ) i n t h e D e v o n i a n of A s t u r i a s and Lebn ( S p a i n ) . L e i d . G e o l . M A . , v . 48, pp. 207-273, p l s . I - X V . Oswald, D . H . ( e d . ) , 1968. I n t e r n a t i o n a l Symposium o n t h e D e vonian S y s t e m Y A l b e r t a Assoc. P e t r . G e o l . , v . I , 1055 p p . , v . 11, 1377 p p . P o l l a r d , J . , 1 9 6 6 . A non-marine o s t r a c o d f a u n a from t h e C o a l Measures of Durhan and N o r t h u m b e r l a n d . P a l a e o n t o l o g y , v . 9 , DP. 6 6 7 - 6 9 7 . , 1 9 6 9 . T h r e e O s t r a c o d - M u s s e l Bands i n t h e Coal-Measu r e s ( W e s t p h a l i a n ) of N o r t h u m b e r l a n d and Durhan. P r o c . Yorks h i r e G e o l . S O C . , v . 3 7 , p p . 239-276, p l . 8 . P r i b y l , A . , 1 9 5 8 . The o s t r a c o d e s t h e Upper C a r b o n i f e r o u s (Nam. A ) of C z e c h o s l o v a k i a ( P o r u b a b e d s ) and i t s i m p o r t a n c e f o r t h e O s t r a v a - K a r a n i v a c o a l d i s t r i c t ( i n Czech w i t h E n g l i s h summary). S b o r n i k U s t r e d U s t a v , G e o l o g i e , v . 2 4 ( 1 9 5 7 ) , odd. P a l e o n t . , p p . :29-7 , 1 9 6 1 . B i o s t r a t i g r a p h i c a l s i g n i f i c a n c e o f t h e Carboni f e r o u s O s t r a c o d a and t h e i r d i s t r i b u t i o n i n t h e O s t r a v a K a r v i n 6 . C o a l D i s t r i c t (Upper S i l e s i a n B a s i n ) of C z e c h o s l o v a k i a . Compt. Rend. Q u a t r i e m Congr. p o u r l ' a v a n c e m e n t etud. S t r a t i g r . e t Ggolog. & C a r b o n i f . , t . 11, p p . 553557. , 1 9 6 1 . Upper C a r b o n i f e r o u s o s t 5 a c o d e s of t h e Hrusov and P e t r k o v i c e b e d s o f O s t r a v a - K a r v i n a C o a l D i s t r i c t (Czecho s l o v a k i a ) . Rozbravy A X . v s . r . 7 1 , pp. 1-54, 9 p l s . Ramsbottom, W . H . C . , 1952. The f a u n a o f t h e C e f n Coed M a r i n e Band i n t h e C o a l Measures a t A b e r b a i d e n n e a r Tondu, Glamorg a n . B u l l . G e o l . S u r v . G r e a t B r i t a i n , v . 4 , p p . 8-30, p l s .
__
-- -
OF
-
9
-
=.
11, 111.
Robinson, J . E . , 1 9 5 9 . The o s t r a c o d e f a u n a o f t h e s h a l e f a c i e s of t h e Cowdor L i m e s t o n e s , n o r y h end o f Cawdor Q u a r r y . Q u a r t . J o u r . Geol. London, v . 1 1 4 , p p . 435-448. Sanchez d e o s a d a , L . C . , and B l e s s , M. J . M . , 1 9 7 1 . Una m i crofauna d e l Westphaliense C de Asturias. d e Microp a l e o n t . , v . 111, pp: 193-204, p l s . I , 11. , 1 9 7 4 . P r e l i m i n a r y n o t e on t h e Lower C a r b o n i f e r o u s O s t r a c o d s from A p r a t h ( F e d e r a l R e p u b l i c o f Germany). I n t e r n a t . 0" B e l g i a n M i c r o p a l e o n t . l i m i t s , Namur, 1 9 7 4 . P u b l i c a t i o n n. 2 , 5 p p . , 2 p l s . Some P e n n s y l v a n i a n k i r k b y S h a v e r , R . H . , and S m i t h , S . , 1 9 7 4 . a c e a n o s t r a c o d e s o f I n d i a n a and M i d - c o n t i n e n t S e r i e s t e r m i n o logy. P r o g r e s s , n . 3 1 , 59 p p . , 3 p l s . Sohn, I . G . , 1 9 6 0 . P a l e o z o i c sDecies of B a i r d i a and r e l a t e d g e n e r a . U.S. G e o l . S u r v . P r o f e s s . ,Paper 330A, 1 1 5 p p . , 6 p l s . 1 9 6 1 . A e c h m i n e l l a , Amphissltes, K i r k b y e l l a and rel- a.t _ e d g e n e-r a _. _ U . S . G e o l . S u r v . P r o f e s s . P a p e r 330B, p p . 107160, p l s . 7 - 1 2 . , ( i n Gordon e t a l . ) , 1 9 6 9 . R e v i s i o n of some G i r t y ' s i n v e r t e b r a t e f o s s i E =om t h e F a y e t t v i l l e S h a l e ( M i s s i s s i p p i a n ) of A r k a n s a s and Oklahoma. O s t r a c o d e s . Id. 606-F, pp. 41-55, p l s . 6-8. Sohn, I . G . , 1 9 6 9 . P s e u d o l e p e r d i t i a S c h n e i d e r , 1956 ( O s t r a c o d a , C r u s t a c e a ) a n e a r l y M i s s i s s i p p i a n g e n u s from S o u t h w e s t e r n Nevada. Id., 643C, p p . 1 - 6 , 1 p l . , 1 9 7 1 . N e w l a t e M i s s i s s i p p i a n O s t r a c o d e g e n e r a and s p e c i e s from N o r t h e r n A l a s k a . g., 7 1 1 A , pp. 1 - 2 4 , p l s . 1-9.
e.
Q~J.
=.of
.
=.=.
, 1 9 7 2 . L a t e P a l e o z o i c O s t r a c o d e s p e c i e s from t h e Con7 1 1 B , p p . 1-15, p l s . 1, 2 . terminous United S t a t e s . , 1975. M i s s i s s i p p i a n O s t r a c o d a of t h e Amsdem F o r m a t i o n ( M i s s i s s i p p i a n and P e n n s y l v a n i a n ) o f Wyoming. Id. 848G, p p . 1 - 2 2 , p l s . 1-3. S t e w a r t , G . A . , and H e n d r i x , W . E . , 1945. O s t r a c o d a of t h e Plum Broo S h a l e , E r i e C o u n t y , O h i o . Jour. P a l e o n t . , v . 1 9 , pp. 87-95, p l . 1 0 . Stumm, E . C . , and W r i g h t , J . D . , 1958. Check l i s t of f o s s i l i n v e r t e b r a t e s d e s c r i b e d from t h e M i d d l e Devonian r o c k s o f t h e Thedford-Arkona R e s i o n of S o u t h e a s t e r n O h i o . P a l e o n t . C o n t r . Michigan Univ. Museum, v . 1 4 , p p . 81-132. Swartz, F . , m 2 . R e v i s i o n of t h e o s t r a c o d e f a m i l y T h l i p s u r i d a e , w i t h d e s c r i p t i o n of a new s p e c i e s from t h e Lower Devoni a n of P e n n s y l v a n i a . P a l e o n t . v . 6 , p p . 36-58, p l s . 1 0 , 11. S w a r t z , F . , and O r i e l , S . S . . 1 9 4 8 . O s t r a c o d a from t h e M i d d l e Devonian’Windom Beds i n W e s t e r n N e w York. Penns l v a n i a C o l l . T e c h . , P a p e r 1 4 2 , p p . 541-566, p l s . S w a r t z , F . , and S w a i n , F . M . . 1 9 4 1 . O s t r a c o d e s o f t h e M i d d l e Devonian’ Onondaga Beds of C e n t r a l P e n n s y l v a n i a . G e o l . SOC. America v . 5 2 , p p . 381-458, p l s . 1 - 8 . S w e e t , W . C . , and B e r a s t r o m , S . M . . 1 9 7 0 . SvmDosium on Conoa . don; B i o s t r a t i g r a p h ; . =.’America 1 2 7 , 499 pp. Thompson, M. J . , and S h a v e r , R . H . , 1 9 6 4 . E a r l y P e n n s y l v a n i a n M i c r o f a u n a s of t h e I l l i n o i s B a s i n . T r a n s . I l l i n o i s S t a t e Acad. S c . , v . 5 7 , p p . 4-23, 1 p l . T h e o n T M . J . , S h a v e r , R . H . , and R i g g s , E . A . , 1959. E a r l y P e n n s y l v a n i a n f u s u l i n i d s and o s t r a c o d s of t h e I l l i n o i s B a s i n . J o u r . P a l e o n t . , v . 33, pp. 770-792, p l s . 104-107. U l r i c h , E . O . , and B a s s l e r , R . S . , 1 9 1 3 . S y s t e m a t i c P a l e o n t o l o g y Lower Devonian O s t r a c o d a Maryland Geol. S u r v . Lower Devonian volume, p p . 513-542, p l s . 95-98. Vanqerow, E. F . , 1 9 5 7 . M i k r o p a l d o n t o l o q i s c h e U n t e r s u c h u n s e n i n d e n K o h l s c h e i d e r S c h i c h t e n i m Wurmrevier b e i Aachen. J b . , Band 7 3 , p p . 457-506, p l s . 20-23. , 1 9 7 0 . D i e f a u n a d e s W e s t d e u t s c h e n O b e r k a r b o n s . pal a e o n t o g r . Band 1 3 4 , p p . 133-152, p l . 1 3 . Weyant, M . , 1 9 7 1 . Recherches m i c r o p a l 6 o n t o l o g i q u e s s u r l e P a l g o z o i c i n f k r i e u r e t moyen d e l ’ h r c h i p e l A r c t i q u e C a n a d i e n . U n p u b l i s h . Ph.D. t h e s e s Univ. Caen. W i l s o n , Ch. W . , 1 9 3 5 . The o s t r a c o d e f a u n a of t h e B i r d s o n g S h a l e , H e l d e r b e g , o f W e s t e r n T e n n e s s e e . Jour. P a l e o n t . , v . 9 , p p . 6 2 9 - 6 4 6 , p l s . 77-78.
s.,
Jour.
s.
--
x.,
&.
s.
,
&.
I
85 EARLY PALEOZOIC CONODONT BIOSTRATIGRAPfIY IS THE ATLANTIC BORDERLAHE S t i g M. B e r g s t r B m Department of Geology and Mineralogy, Columbus,
The O h i o S t a t e U n i v e r s i t y
O h i o 43210
ABSTRACT
Cambrian c o n o d o n t s ,
s t i l l r e l a t i v e l y p o o r l y known b u t a p p a r e i i t l y
w i d e s p r e a d i n t h e A t l a n t i c B o r d e r l a n d s , i n c l u d e s o m e 15 m u l t i e l e m e n t genera.
D e s c r i b e d f a u n a s a r e m a i n l y frsm S c a n d i n a v i a , P o l a n d , a n d
Germany ( e r r a t i c s ) w h e r e a s l i t t l e i n f o r m a t i o n i s a v a i l a b l e
from, f o r
i n s t a n c e , e a s t e r n N o r t h A m e r i c a a n d t h e B r i t i s h I s l e s . A l t h o u g h no s u c c e s s i o n of f o r n a l l y d e f i n e d c o n o d o n t z o n e s h a s a s y e t b e e n e s t a b l i s h e d t h r o u g h o u t t h e s y s t e m , Cambrian c o n o d o n t s have c o n s i d e r a b l e p o t e n t i a l as g u i d e f o s s i l s , p a r t i c u l a r l y i n t h e U p p e r C a m b r i a n , w h e r e many forms show l i m i t e d v e r t i c a l r a n g e s a n d v e r y w i d e h o r i z o n t a l distributions. O r d o v i c i a n c o n o c b n t s , w h i c h i n c l u d e a b o u t 80 m u l t i e l e m e n t g e n e r a , a r e much b e t t e r known t h a n t h e C a m b r i a n o n e s , a n d show s t r i k i n g provincial d i f f e r e n t i a t i o n throughout
the period.
I n the North
A m e r i c a n M i d c o n t i n e n t P r o v i n c e , some 1 7 b i o s t r a t i g r a p h i c u n i t s of zonal t y p e have been recognized and i n t h e North A t l a n t i c Province, some 1 5 z o n e s ,
a n d more t h a n 10 s u b z o n e s , h a v e b e e n f o r m a l l y d e f i n e d .
Ordovician conodonts have proved very u s e f u l biDstratigrdphically,
in
many i n s t a n c e s p r o v i d i n g a s t r a t i g r a p h i c r e s o l u t i o n s u p e r i o r t o t h a t of any o t h e r f o s s i l group. S i l u r i a n c o n o d o n t f a u n a s e x h i b i t f a r less t a x o n o m i c d i v e r s i t y ( a b o u t 15 m u l t i e l e m e n t g e n e r a ) a n d p r o v i n c i a l d i f f e r e n t i a t i o n t h a n Ordovician ones.
T h e S i l u r i a n c o n o d o n t s u c c e s s i o n , b e s t known i n t h e
A t l a n t i c B o r d e r l a n d s from Great B r i t a i n , S c a n d i n a v i a , V i r g i n i a , a n d t h e e a s t e r n part
o f t h e North American M i d c o n t i n e n t , form t h e b a s i s
f o r a b o u t 1 2 z o n e s . Many of t h e s e z o n e s h a v e b e e n w i d e l y r e c o g n i z e d n o t o n l y i n Europe and North A m e r i c a
but also i n Australia,
Asia,
and n o r t h e r n Africa. A l t h o u g h p u b l i s h e d r e p o r t s on E a r l y P a l e o z o i c c o n o d o n t s of A f r i c a a n d S o u t h America i n c l u d e o n l y a f e w p a p e r s , t h e a v a i l a b l e
d a t a s u g g e s t t h a t t h e conodonts f a u n a s from t h o s e c o n t i n e n t s are, by and l a r g e ,
v e r y s i m i l a r t o t h o s e known f r o m t h e n o r t h e r n h e m i s p h e r e .
C o - o c c u r r e n c e of s t r a t i g r a p h i c a l l y d i a g n o s t i c c o n o d o n t s a n d
g r a p t o l i t e s h a s made i t p o s s i b l e t o t i e t o g e t h e r c o n o d o n t and g r a p t 3 l i t e z o n a l u n i t s i n t h e O rd o v i cian and S i l u r i a n a t a r e l a t i v e l y l a r g e
number o f s t r a t i g r a p h i c l e v e l s . G e o g r a p h i c d i s t a n c e a p p a r e n t l y was o f r e l a t i v e l y minor i m p o r t a n c e i n comparison w i t h e c o l o g i c f a c t o r s f o r t h e e s t a b l i s h m e n t of p a t t e r n s i n t h e E a r l y P a l e o z o i c conodont biogeography.
This conclu-
s i o n , a l o n g w i t h t h e f a c t t h a t v e r y similar c o n o d o n t f a u n a s were p r e s e n t on b o t h s i d e s o f t h e P r o t o - A t l a n t i c ,
make c o n o d o n t s o f l i t t l e
u s e f o r e v a l u a t i o n s o f t h e s i z e a n d d e v e l o p m e n t of c l i e P r o t o - A t l a n t i c Ocean, a t l e a s t a t t h e p r e s e n t t i m e .
------------INTRODUCTION
C o n o d o n t s a r e common m i c r o f o s s i l s i n many t y p e s o f Lower P a l e o z o i c m a r i n e r o c k s . Due t o t h e i r r a p i d e v o l u t i o n , t h e v e r y w i d e h o r i z o n t a l d i s t r i b u t i o n o f many t a x a , a n d t h e f a c t t h a t numerous s p e c i e s a p p a r e n t l y were n o t s t r o n g l y f a c i e s - c o n t r o l l e d ,
conodonts
now r a n k among t h e most u s e f u l f o s s i l s b i o s t r a t i g r a p h i c a l l y i n t h e
Lower P a l e o z o i c .
The g r o u p h a s b e e n known f o r 1 2 0 y e a r s b u t t h e
p e r i o d of modern a n d i n t e n s e s t u d y o f t h e s e f o s s i l s b e g a n a b o u t 1950 when t h e i r g r e a t b i o s t r a t i g r a p h i c p o t e n t i a l b e g a n t o b e a p p a r e n t . Since
then,
t h e c o n o d o n t l i t e r a t u r e h a s grown v e r y r a p i d l y a n d c l o s e
t o 100 p a p e r s ,
i n which
c o n o d o n t s a r e d e a l t w i t h i n o n e form o r a n -
o t h e r , a r e now p u b l i s h e d a n n u a l l y .
Most p a p e r s a r e of b i o s t r a t i -
g r a p h i c o r taxonomic n a t u r e b u t d u r i n g t h e l a s t f e w y e a r s ,
the paleo-
e c o l o g y and m i c r o m o r p h o l o g y o f t h e s e f o s s i l s h a v e a t t r a c t e d c o n s i d e r a b l e i n t e r e s t , and i t i s l i k e l y t h a t t h e s e areas of r e s e a r c h w i l l be increasingly important i n the future. Since t h e mid-l960's,
c o n o d o n t taxonomy h a s gone t h r o u g h a
p e r i o d of f u n d a m e n t a l change. d i f f e r e n t types
On t h e b a s i s o f t h e f a c t t h a t s e v e r a l
of c o n o d o n t e l e m e n t s e v i d e n t l y o c c u r r e d i n t h e
a p p a r a t u s of a s i n g l e i n d i v i d u a l ,
t h e taxonomy h a s
developed rapid-
l y f r o m a p u r e f o r m taxonomy b a s e d o n e x t e r n a l s h a p e o f s i n g l e e l e m e n t s , t o a z o o l o g i c a l l y more s o u n d m u l t i e l e m e n t taxonomy b a s e d on r e c o n s t r u c t e d a s s e m b l a g e s o f e l e m e n t s . A g r e a t many c o n o d o n t t a x a h a v e now b e e n r e - e v a l u a t e d
on a m u l t i e l e m e n t b a s i s b u t i t w i l l t a k e
a l o n g t i m e t o c o m p l e t e t h i s r a d i c a l t a x o n o m i c r e v i s i o n b e c a u s e many
h u n d r e d s p e c i e s and more t h a n 300 g e n e r a were o r i g i n a l l y p r o p o s e d
87 a s form t a x a . The p u r p o s e o f t h e p r e s e n t c o n t r i b u t i o n i s t o s u m m a r i z e b r i e f l y t h e conodont b i o s t r a t i s r a p h y s o f a r e s t a b l i s h e d f o r Cambrian t h r o u g h S i l u r i a n rocks i n t h e areas b o r d e r i n g t h e p r e s e n t A t l a n t i c . of geographic e x t e n t ,
I n terms
t h e area d e a l t w i t h i n c l u d e s S o r t h America e a s t
o f 90°w L o n g i t u d e , w e s t e r n and c e n t r a l E u r o p e , and t h e B a l t o s c a n d i c region.
B e c a u s e l i t t l e i s known a b o u t Lower P a l e o z o i c c o n o d o n t s i n
South A m e r i c a (see, f o r i n s t a n c e , S e r p a g l i ,
for i n s t a n c e , E t h i n g t o n a n d F u r n i s h ,
1 9 7 4 ) and A f r i c a (see,
1962), t h e review w i l l i n c l u d e
d a t a m a i n l y f r o m N o r t h A m e r i c a and E u r o p e . A l t h o u g h p u b l i s h e d s e v e r a l y e a r s a g o , and t h e r e f o r e o u t d a t e d i n some r e s p e c t s , a summary volume
e n t i t l e d "Symposium on Conodont B i o s t r a t i g r a p h y " eds.,
(Sweet and Bergstrbm,
1 9 7 1 ) s t i l l p r o v i d e s a u s e f u l summary o f Lower P a l e o z o i c c o n o -
dont b i o s t r a t i g r a p h y ,
a n d t h e r e a d e r is r e f e r r e d t o t h a t volume f o r
a more d e t a i l e d t r e a t m e n t t h a n i s p o s s i b l e w i t h i n t h e s c o p e o f t h e present contribution.
P a p e r s i n t h a t volume a l s o g i v e i l l u s t r a t i o n s
of many of t h e s t r a t i g r a p h i c a l l y i m p o r t a n t c o n o d o n t s m e n t i o n e d b e l o w .
CAMBRIAR
The o l d e s t c o n o d o n t s known a r e f r o m s t r a t a i n S i b e r i a c u r r e n t l y c l a s s i f i e d as l a t e Precambrian (Missarzhevsky, Missarzhevsky,
1 9 7 3 ; Matthews a n d
1 9 7 5 ) b u t t h e g r o u p , w h i c h i s r e p r e s e n t e d by s p e c i e s
of t h e o r d e r P a r a c o n o d o n t i d a ,
exhibited l i t t l e d i v e r s i t y before Late
Cambrian time. The p a r a c o n o d o n t s a r e w e a k l y p h o s p h a t i z e d a n d d i f f e r a l s o i n o t h e r r e s p e c t s from t h e co n o d o n t i f or m c o n o d o n t s , which i n c l u d e p r a c t i c a l l y a l l post-Cambrian forms. A l t h o u g h known f r o m t h e B a l t o s c a n d i c area ( M C l l e r ,
1959; Poul-
s e n , 1 9 6 6 ; B e n g t s s o n , 1 9 7 6 ) , Germany (Mfiller, 1 9 5 9 , e r r a t i c b o u l d e r s ) , Poland ( S z a n i a w s k i , 1 9 7 1 ) , t h e B r i t i s h Isles (Miller and Rushton, 1 9 7 3 ) , a n d N e w York S t a t e ( L a n d i n g , 1 9 7 4 b , 1 9 7 6 ) , C a m b r i a n c o n o d o n t s are s t i l l v e r y l i t t l e s t u d i e d i n t h e A t l a n t i c B o r d e r l a n d s ( F i g . 1 ) . T h e i r p o t e n t i a l as b i o s t r a t i g r a p h i c t o o l s was s u g g e s t e d i n t h e p i o n e e r work by M g l l e r
( 1 9 5 9 ) , b u t s t u d i e s on Cambrian c o n o d o n t
b i o s t r a t i g r a p h y have s o f a r b een co n cerned o n l y w i t h m ai nl y L a t e Cambrian s u c c e s s i o n s i n a r e a s s u c h a s w e s t e r n N o r t h America ( M i l l e r , 1 9 7 5 ) , I r a n (Mcller, 1 9 7 3 ) , Q u e e n s l a n d ( D r u c e a n d J o n e s ,
1 9 7 1 ) , and
C h i n a (Nogami, 1 9 6 6 , 1 9 6 7 ) . Mcller ( 1 9 7 3 ) a n d Miller ( 1 9 7 5 ) i n t r o -
88
2oi
W Spitsbergen
w
10
D
=
I
N Scotland
3
PRO TO-ATLANTIC OCEAN
20 E Spltsbergen
F i g . 1 . Cambrian conodont o c c u r r e n c e s i n N o r t h A m e r i c a and n o r t h w e s t e r n Europe p l o t t e d on a s k e t c h - m a p s h o w i n g i n f e r r e d c o n t i n e n t p o s i t i o n s d u r i n g M i d d l e Cambrian t i m e . A s i n F i g . 2 a n d 4, e a c h d o t d e n o t e s one o r s e v e r a l conodont l o c a l i t i e s which have p r o d u c e d c o l l e c t i o n s d e s c r i b e d i n t h e l i t e r a t u r e o r s e e n by t h e w r i t e r . L a t i t u d i n a l p o s i t i o n of n o r t h w e s t e r n E u r o p e b a s e d on K o l t i m i e r and B e r g s t r h ( i n p r e p a r a t i o n ) .
d u c e d z o n a l u n i t s b a s e d on Late Cambrian species a n d t h e r e a r e i n d i c a t i o n s t h a t a t l e a s t some of t h e s e z o n e s a r e r e c o g n i z e a b l e i n t h e Appalachians
( L a n d i n g , 1 9 7 6 ) . N o a t t e m p t h a s b e e n made s o f a r t o
e s t a b l i s h f o r m a l c o n o d o n t b i o s t r a t i g r a p h i c u n i t s i n t h e Lower a n d M i d d l e C a m b r i a n a n d a p a r t f r o m a f e w p a p e r s ( L a n d i n g , 197413; P o u l s e n , 1 9 6 6 ; B e n g t s s o n , 1976), c o n o d o n t s o f t h a t a g e r e m a i n v i r t u a l l y un-
s t u d i e d i n t h e A t l a n t i c area.
The known Cambrian c o n o d o n t f a u n a s i n -
c l u d e some 1 5 g e n e r a of s i m p l e f o r m s s u c h a s F u r n i s h i n a ,
",
Prosagittodontus,
Muellerina, Hertzina,
Proconodon-
Prooneotodus,
P r o s c a n d o d u s a n d h a v e a somewhat monotonous c h a r a c t e r .
and
However,
there
a r e c l e a r i n d i c a t i o n s t h a t some s p e c i e s h a v e a w i d e g e o g r a p h i c d i s t r i b u t i o n combined w i t h a r e a s o n a b l y s h o r t v e r t i c a l r a n g e ; accor di n g l y , a p r o m i s i n g and c h a l l e n g i n g t a s k would b e t o e x p l o r e t h e i r
89 W Spitsbergen
N . Ireland
0 0 0
PROTO-ATLANTIC OCEAN ? E\
N e w Brunswick
Nova Scotia \
Fig.2. Ordovician conodont occurrences i n North America and northwestern Europe plotted on a sketch-map showing inferred positions of continents in Early Ordovician (Arenigian) time. Latitudinal position of Europe according to Noltimier and Bergstram (1976; in preparation), that of North America based on McElhinny and Opdyke (1973). Dark dots denote North Atlantic Province faunas, open circles Midcontinent Province faunas. Note the presence of North Atlantic Province conodont faunas in the eastern Appalachians and in California (Klamath Mountains). utility as regionally useful index fossils in the Atlantic area.
ORDOVICIAN
The Early and Middle Ordovician was a time of rapid conodont evolution that led to a diversification at the generic and specific
90 l e v e l t h a t i s a p p a r e n t l y g r e a t e r than t h a t of any other comparable t i m e i n t e r v a l i n t h e s t r a t i g r a p h i c record o f conodonts.
This is i l l u s -
t r a t e d by t h e f a c t t h a t some 150 f o r m g e n e r a o f c o n o d o n t s h a v e b e e n p r o p o s e d on t h e b a s i s of O r d o v i c i a n c o l l e c t i o n s a n d more t h a n h a l f of t h e s e a r e d i s t i n c t as m u l t i e l e m e n t g e n e r a .
In a d d i t i o n , t h e r e a r e
q u i t e a f e w c h a r a c t e r i s t i c , b u t s t i l l unnamed,
g e n e r a , and t h e t o t a l
number o f O r d o v i c i a n m u l t i e l e m e n t g e n e r a may u l t i m a t e l y p r o v e t o b e
w e l l i n e x c e s s o f 100. T h i s f i g u r e i s more t h a n t w i c e as h i g h a s t h a t f o r any o t h e r g e o l o g i c p e r i o d . O r d o v i c i a n conodonts h a v e b e e n f a r more i n t e n s e l y s t u d i e d t h a n t h e Cambrian o n e s i n t h e A t l a n t i c B o r d e r l a n d s b u t t h e r e a r e , n e v e r t h e -
l e s s , l a r g e g a p s i n o u r k n o w l e d g e a b o u t f a u n a s of t h a t a g e .
Best
known a r e f a u n a s f r o m t h e B a l t o s c a n d i c a r e a a n d t h e N o r t h A m e r i c a n Midcontinent b u t t h o s e from s e v e r a l o t h e r major r e g i o n s ,
f o r instance,
S o u t h America, n o r t h e r n A f r i c a , and G r e e n l a n d , r e m a i n v i r t u a l l y unexplored.
Fig.2 g i v e s t h e g e n e r a l geographic l o c a t i o n of important
Ordovician conodont o c c u r r e n c e s i n areas b o r d e r i n g t h e p r e s e n t North Atlantic. Taken as a w h o l e , O r d o v i c i a n c o n o d o n t f a u n a s p r o b a b l y e x h i b i t a s t r o n g e r biogeographic d i f f e r e n t i a t i o n than t h o s e of any o t h e r system (Bergstram,
1 9 7 3 a ; B a r n e s e t a l . , 1 9 7 3 ; Sweet and B e r g s t r a m , 1 9 7 4 ) .
This f a u n a l provincialism,
which c a n b e t r a c e d b a c k t o t h e Tremadoc-
i a n , p r e v a i l e d to t h e end of t h e p e r i o d a l t h o u g h modified as t o a r e a l e x t e n t and d i s t i n c t i v e n e s s . recognized,
The two main f a u n a l p r o v i n c e s g e n e r a l l y
t h e North A m e r i c a n M i d c o n t i n e n t P r o v i n c e and t h e N o r t h
A t l a n t i c P r o v i n c e , a r e c h a r a c t e r i z e d b y two c o n o d o n t f a u n a s s o d i f f e r e n t t h a t s e p a r a t e z o n a l schemes have been e s t a b l i s h e d f o r e a c h o f them ( S w e e t e t a l . ,
1971; E t h i n g t o n and C l a r k ,
1971; Bergstram,
1 9 7 1 a , 1 9 7 1 b ; L i n d s t r B m , 1 9 7 1 ) and t h e r e i s s t i l l c o n s i d e r a b l e unc e r t a i n t y regarding t h e p r e c i s e r e l a t i o n s between s e v e r a l u n i t s i n t h e s e schemes. A p r o v i s i o n a l c o r r e l a t i o n between t h e s e p r o v i n c i a l schemes i s g i v e n i n F ig . 3 .
North American M i d c o n t i n e n t P r o v i n c e
Lower O r d o v i c i a n ( C a n a d i a n ) c o n o d o n t f a u n a s i n t h i s p r o v i n c e , w h i c h o c c u p i e s t h e c e n t r a l p o r t i o n o f t h e N o r t h A m e r i c a n c o n t i n e n t as w e l l a s p a r t s of S i b e r i a a n d A u s t r a l i a , are c h a r a c t e r i z e d by " s i m p l e -
cone" g e n e r a s u c h as Acanthodus, Scolopodus, U l r i c h o d i n a , Oneotodus,
91 IIDCONTINENT I A M G BRITAIN EALTOSCANDIE
‘ z
ASHGlLLlAN
5 -z
a I
LLANDEILIAN
4
3
1-I
HARJUAN
10
CARADOCIAN
Z
UJNAS
NORTH ATLANTIC CONODONT
ZONES
I
SUEZONES
Amorphognathus ordovicfcus
NOT YET DISTINGUISHED Amorphognathus superbus
Arnorphognathus tvaerensfs
1
Prfoniodus alobarus Prioniodus gerdae
Pygodus ansennus
VIRUAN
’
1
?
i
Eoplacognarhus robusrus
Pygodus serrus
I
Eoplacognarhus reclfnarus
I
Eoplacognarhus fohaceus
0
- ?
b
LLANVIRNIAN
UNNAMED
Eoolacoonarhus suecfcus
Eoplacognarhus variabilis 4
3
Mfcrozarkodina parva Paroistodus originalis Prioniodus navis
7-
ARENIGIAN
z 9
OELANDIAN
NOT YET DISTINGUISHED
?
2
5
TREMADOCIAN
Fig.3. C o r r e l a t i o n between S o r t h American and European Ordovician standard series, Midcontinent Province conodont f a u n a s (Sweet e t a l . , 1971; Ethington and C l a r k , 1 9 7 1 ) , and S o r t h A t l a n t i c Province conodont zones and subzones (Lindstrcm, 1971; B e r g s t r h , 1 9 7 1 a , 1 9 7 1 b ) . A s shown in t h e d i a g r a m , t h e s t r a t i g r a p h i c s c o p e of t h e t h r e e m a i n s u b d i v i s i o n s o f t h e O r d o v i c i a n i s , by t r a d i t i o n , n o t t h e same i n E u r o p e a n d N o r t h America; i n t h e t e x t b e l o w , t h e terms L o w e r , M i d d l e , a n d Upper O r d o v i c i a n a r e u s e d i n t h e i r l o c a l s e n s e , t h a t i s , t o d e n o t e t h e Oelandian, Viruan, and H a r j u a n i n Europe, and t h e Canadian, Champlainian, and C i n c i n n a t i a n i n N o r t h A m e r i c a . Top o f C a n a d i a n i s t a k e n a s t h e b a s e o f t h e W h i t e r o c k i a n S t a g e . A s t o t h e un-named i n t e r v a l s b e t w e e n t h e T r e m a d o c i a n / A r e n i g i a n a n d t h e L l a n v i r n i a d L l a n d e i l i a n , see Bergstrom e t a l . (1973, 1 9 7 4 ) . “ P a l t o d u s f ‘ , a n d o t h e r s . Compound-element g e n e r a are less w e l l repres e n t e d b u t i n c l u d e Cordylodus, Loxodus, Chosonodina,
and Oepikodus.
A l t h o u g h e a r l i e s t O r d o v i c i a n f a u n a s b e g i n t o b e w e l l known
1969; D r u c e a n d J o n e s ,
(Miller,
1 9 7 1 ) much r e m a i n s t o b e l e a r n e d a b o u t y o u n g e r
E a r l y O r d o v i c i a n c o n o d o n t f a u n a s of t h i s p r o v i n c e .
Fortunately,
this
s e r i o u s gap i n o u r k n o w l e d g e may b e a t l e a s t p a r t i a l l y f i l l e d b y new d a t a f r o m s t i l l l a r g e l y u n p u b l i s h e d s t u d i e s i n h’ew S l e x i c o - T e x a s ( R e p e t s k i , 1974) a n d Oklahoma
(Mound, 1 9 6 8 ; B r a n d ,
1 9 7 6 ; P o t t e r , 1975).
92 C o r r e s p o n d i n g work i n A s i a i s b e i n g c a r r i e d o u t b y , among o t h e r s , Lee ( 1970, 1975 )
.
D e s c r i p t i o n s o f e a r l i e s t Middle O r d o v i c i a n ( e a r l i e s t Champlaini a n ) Midcontinent Province conodont f a u n a s remain l a r g e l y unpublished i n N o r t h America b u t t h e a v a i l a b l e p u b l i s h e d a n d u n p u b l i s h e d d a t a
( s e e , f o r i n s t a n c e , Mound, 1 9 6 5 ; Bradshaw, 1 9 6 9 ) show t h a t M i d c o n t i n e n t f a u n a s of t h i s a g e i n c l u d e , a p a r t f r o n a n a r r a y o f " s i m p l e - c o n e " g e n e r a s u c h as D r e p a n o i s t o d u s , P a n d e r o d u s , O i s t o d u s , a n d S c o l o p o d u s , a d i s t i n c t i v e s u i t e o f compound-element
genera, f o r instance, Multi-
oistodus, Leptochirognathus, Histiodella, d u s , and T r i c l a d i o d u s . -
Bergstroemognathus
,
Erismo-
Typical p l a t f o r m elements are rare o r missing
i n most f a u n a s of t h i s t y p e , b u t o c c u r ( P o l y p l a c o g n a t h u s , S c y p h i o d u s ) i n s l i g h t l y y o u n g e r M i d d l e O r d o v i c i a n f a u n a s a l o n g w i t h a b u n d a n t rep r e s e n t a t i v e s o f Phragmodus,
Plectodina,
and f i b r o u s c o n o d o n t g e n e r a
s u c h a s C u r t o g n a t h u s , C h i r o g n a t h u s , a n d C o l e o d u s ( B e r g s t r h a n d Sweet, 1966 ; S c h o p f , 1966 ; W e b e r s ,
1966 ).
U n f o r t u n a t e l y , nowhere i n t h e
North American M i d c o n t i n e n t i s t h e r e a r e a s o n a b l y c o m p l e t e Middle O r d o v i c i a n ( C h a m p l a i n i a n ) s u c c e s s i o n and t h e s e q u e n c e of f a u n a s p r o p o s e d by S w e e t e t a l .
( 1 9 7 1 ) is p i e c e d t o g e t h e r f r o m s e c t i o n s i n
s e v e r a l d i f f e r e n t areas. Although t h e p r e c i s e c o r r e l a t i o n between some u n i t s i n t h e s e a r e a s is s t i l l u n c e r t a i n , t h e g e n e r a l s u c c e s s i o n o f u n i t s i n t h i s scheme h a s p r o v e d t o b e v e r y u s e f u l r e g i o n a l l y o v e r t h e N o r t h A m e r i c a n c r a t o n ( S w e e t and B e r g s t r a m ,
i n press).
A t p r e s e n t t h e b e s t known L a t e O r d o v i c i a n ( C i n c i n n a t i a n ) c o n o -
dorit f a u n a l s u c c e s s i o n i n N o r t h A m e r i c a i s t h a t o f t h e C i n c i n n a t i a r c h a r e a i n O h i o a n d a d j a c e n t s t a t e s ( S w e e t , i n p r e s s ) . The L a t e O r d o v i c i a n f a u n a s t h e r e i n c l u d e r e p r e s e n t a t i v e s o f , among o t h e r s , A p h e l o g n a t h u s , B e l o d i n a , O u l o d u s , Phragmodus, dognathus.
I n some i n t e r v a l s ,
t a t i v e s of Amorphognathus,
Plectodina,
and R h i p i -
t h e r e are a l s o o c c u r r e n c e s o f r e p r e s e n -
I c r i o d e l l a , Periodon,
Protopanderodus,and
Rhodesognathus, which are g e n e r a l l y tak e n t o b e b a s i c a l l y Nor t h A t l a n t i c P r o v i n c e f o r m s . Some o f t h e s e are i m p o r t a n t s t r a t i g r a p h i c a l l y and s e r v e as c o r r e l a t i v e l i n k s w i t h t h e North A t l a n t i c conodont
z o n a l s u c c e s s i o n , w h e r e t h e y a r e f a r more w i d e l y d i s t r i b u t e d .
By a n d
l a r g e , t h e i n d i g e n o u s C i n c i n n a t i a n c o n o d o n t f a u n a s a r e of a r a t h e r monotonous a n d c o n s e r v a t i v e t y p e and t h e y are n o t n o t e d f o r s h o w i n g conspicuous e v o l u t i o n a r y changes through t h e sequence.
In t h e C i n c i n n a t i r e g i o n , as w e l l a s e l s e w h e r e i n e a s t e r n USA, t h e i n t e r v a l w i t h i n which t h e O r d o v i c i a n / S i l u r i a n boundary f a l l s
is
93 marked b y a s t r i k i n g c h a n g e i n t h e c o m p o s i t i o n o f t h e c o n o d o n t faunas.
A m a j o r i t y of t h e t y p i c a l M i d c o n t i n e n t p r o v i n c e g e n e r a a p p a r -
e n t l y became e x t i n c t i n l a t e s t O r d o v i c i a n t i m e a n d o n l y P a n d e r o d u s and a f e w o t h e r " s i m p l e - c o n e " Amorphognathus,
Icriodella,
g e n e r a , as w e l l a s r e p r e s e n t a t i v e s o f
O u l o d u s , and a f e w o t h e r c o m p o u n d - e l e m e n t
genera, survived i n t o S i l u r i a n t i m e .
Accordingly, S i l u r i a n conodont
f a u n a s d i f f e r a good d e a l f r o m O r d o v i c i a n o n e s a n d t h e g e n e r a l e x t i n c t i o n o f c o n o d o n t g e n e r a i n Late O r d o v i c i a n time i s one o f t h e most s e v e r e d u r i n g t h e e n t i r e t i m e o f e x i s t e n c e o f t h e g r o u p .
The N o r t h A t l a n t i c P r o v i n c e
Early Ordovician
(Oelandian) conodont f a u n a s of t h e North Atlan-
t i c P r o v i n c e a r e c u r r e n t l y b e s t known f r o m t h e B a l t o s c a n d i c a r e a ( L i n d s t r c m , 1955, 1960, 1971; S e r g e e v a , 1964; V i i r a , Kohut,
1972; van W a m e l ,
d e s c r i b e d f r o m S c o t l a n d (Lamont a n d L i n d s t r g m ,
America ( F $ h r a e u s ,
1967, 1975;
1 9 7 4 ) a l t h o u g h a few c o l l e c t i o n s h a v e b e e n
1970; Bergstram e t a l . ,
1957), e a s t e r n Korth
1972; Landing,
1974a),
and A r g e n t i n a ( S e r p a g l i , 1 9 7 4 ) . E a r l y O r d o v i c i a n f a u n a s o f t h i s p r o vince are c h a r a c t e r i z e d by a wide v a r i e t y of such as Drepanodus, A co n ti o d u s ,
Paroistodus,
Protopanderodus,
"simple-cone"
Drepanoistodus,
Oistodus,
genera Paltodus,
S c a n d o d u s , and S t o l o d u s a s w e l l a s
compound-element g e n e r a s u c h a s C o r d y l o d u s , M i c r o z a r k o d i n a , and P e r i o d o n .
Oepikodus,
It is of i n t e r e s t t o n o t e t h a t t r u e platform conodonts
a p p e a r e d as e a r l y a s i n e a r l i e s t O r d o v i c i a n ( T r e m a d o c i a n ) t i m e i n t h e B a l t o s c a n d i c area ( L i n d s t r B m , 1 9 5 5 ) b u t s u c h c o n o d o n t s d i d n o t c o n s t i t u t e a s u b s t a n t i a l p a r t of the conodont f a u n a s u n t i l Middle Ordovician (Viruan) t i m e .
S u c c e s s i o n s o f c o n o d o n t z o n e s p r o p o s e d by
S e r g e e v a ( 1 9 6 4 ) , L i n d s t r b ( 1 9 7 1 ) , a n d V i i r a ( 1 9 7 5 ) , among o t h e r s , d i f f e r r e l a t i v e l y l i t t l e f r o m e a c h o t h e r a n d t h e one i n t r o d u c e d by Lindstrbm (1971) has proved t o be widely a p p l i c a b l e .
Van Wamel ( 1 9 7 4
p r o p o s e d a f a r more d e t a i l e d z o n a l s u b d i v i s i o n o f T r e m a d o c i a n a n d E a r l y A r e n i g i a n s t r a t a i n e a s t e r n Sweden b u t some o f h i s 2 0 assemb l a g e z o n e s , w h i c h were b a s e d on s i n g l e s a m p l e s , may b e o n l y o f l o c a b i o s t r a t i g r a p h i c s i g n i f i c a n c e , i f any. Lindstram's
On t h e o t h e r h a n d , u n i t s of
( 1 9 7 1 ) z o n a l s u c c e s s i o n have been r e c o g n i z e d i n areas as
f a r a p a r t a s N e w f o u n d l a n d , Texas, Nevada, and A r g e n t i n a , a n d i t i s c l e a r t h a t conodonts r i v a l g r a p t o l i t e s as t h e b i o s t r a t i g r a p h i c a l l y most u s e f u l f o s s i l g r o u p i n t h e Lower O r d o v i c i a n o f t h i s p r o v i n c e .
94 In t h e North A t l a n t i c P r o v i n c e , Middle Ordovician ( V i r u a n ) c o n o d o n t s a r e b e s t known f r o m t h e B a l t o s c a n d i c a r e a ( B e r g s t r B m , 1971a, 1971b; H a m a r ,
1964, 1966; Lindstram,
1960; V i i r a ,
1962,
1975) and
e a s t e r n S o r t h America ( B e r g s t r 6 m , 1 9 7 1 a , 1 9 7 3 c ; B e r g s t r b e t a l . , 1974; Bergstrijm and C a r n e s ,
i n p r e s s ; Fahraeus, 1970, 1973; Sweet
a n d I i e r g s t r B m , 1 9 6 2 ) b u t some u n i t s i n Great B r i t a i n (Lamont a n d L i n d s t r h , 1957; Lindstrom,
1 9 5 9 ; B e r g s t r a m , 1 9 6 4 , 1 9 7 1 a ; Rhodes,
1951) and France (Lindstrhm e t a l . ,
1974) have a l s o y i e l d e d conodonts
of t h i s a g e . Z o n a l u n i t s h a v e b e e n p r o p o s e d by B e r g s t r z m ( 1 9 7 1 a ) a n d V i i r a ( 1 9 7 5 ) a n d many o f t h e u n i t s i n t h e f o r m e r ' s z o n a l s u c c e s s i o n
have been r e c o g n i z e d r e g i o n a l l y i n t h e North A t l a n t i c P r o v i n c e . Rliddle O r d o v i c i a n c o n o d o n t f a u n a s o f t h i s a r e a a r e c h a r a c t e r i z e d by a s u c c e s s i o n of s t r a t i g r a p h i c a l l y i m p o r t a n t s p e c i e s of Amorphognathus, E o p l a c o g n a t h u s , P r i o n i o d u s ( B a l t o n i o d u s ) , a n d P y g o d u s as w e l l a s by t h e w i d e s p r e a d p r e s e n c e of P e r i o d o n a n d " s i m p l e - c o n e " Protopanderodus.
genera such as
R e p r e s e n t a t i v e s of I c r i o d e l l a , a l t h o u g h r a t h e r
common i n B r i t i s h f a u n a s , a r e r a r e i n B a l t o s c a n d i a b u t known a l s o f r o m t h e A p p a l a c h i a n s ; i t s h o u l d be n o t e d , h o w e v e r , t h a t t y p i c a l Korth A t l a n t i c P r o v i n c e f a u n a s o f l a t e Middle O r d o v i c i a n a g e are c u r r e n t l y n o t known f r o m t h e A p p a l a c h i a n s . L a t e Ordovician ( H a r j u a n ) f a u n a s of North A t l a n t i c Province
ts'pe a r e as y e t n o t d o c u m e n t e d i n e a s t e r n N o r t h America, b u t a r e known f r o m s e v e r a l a r e a s i n E u r o p e , strbm,
1971a; V i i r a ,
including Ealtoscandia (Berg-
1 9 7 5 ) , Great B r i t a i n ( R h o d e s , 1 9 5 5 ; B e r g s t r a m ,
1964, 1 9 7 1 a ) , t h e C a r n i c Alps i n I t a l y and A u s t r i a ( S e r p a g l i , 1967; SchBnlaub, 1 9 7 1 ) , T h u r i n g i a (KnGpfer, 1 9 6 7 ) , and S p a i n ( F u g a n t i and S e r p a g l i , 1 9 6 8 ) . The b i o s t r a t i g r a p h i c z o n a t i o n o f t h e Upper O r d o v i c i a n b a s e d on c o n o d o n t s i s s t i l l p r e l i m i n a r y ( B e r g s t r a m , 1 9 7 1 a ) a n d some r e f i n e m e n t w i l l n o d o u b t b e p o s s i b l e when t h e p e r t i n e n t f a u n a s a r e b e t t e r known t h a n a t t h e p r e s e n t t i m e .
Sorth Atlantic Province
f a u n a s of t h i s a g e a r e less v a r i e d t h a n f a u n a s from o l d e r p a r t s o f t h e s y s t e m ; comnon a n d w i d e s p r e a d e l e m e n t s i n c l u d e A m o r p h o g n a t h u s , D a p s i l o d d s , Hamarodus, P e r i o d o n , G t o p a n d e r o d u s , P r i o n i o d u s , a n d Strachanoynathus wnereas forms o f I c r i o d e l l a , D i c h o d e l l a , and Nordiod u s h a v e a more l i m i t e d d i s t r i b u t i o n . A s i s a l s o t h e c a s e i n t h e
_-
Nidcontinent Province, the i n t e r v a l near the Ordovician/Silurian b o u n d a r y i s marked b y a d r a s t i c e x t i n c t i o n of common a n d w i d e s p r e a d Uorth A t l a n t i c P r o v i n c e t a x a ; o n l y Amorphognathus, H i n d e o d e l l a ( O z a r k o d i n a ) , and I c r i o d e l l a , a l o n g w i t h sosne " s i m p l e - c o n e " g e n e r a ,
95 survived i n t o t h e S i l u r i a n .
I t s h o u l d be n o t e d , however, t h a t e a r l i -
est S i l u r i a n ( e a r l y L l a n d o v e r i a n ) conodonts are v i r t u a l l y
unknown i n
t h e a r e a u n d e r d i s c u s s i o n a n d t h i s may make t h e e x t i n c t i o n seem more a b r u p t t h a n i s r e a l l y t h e case.
Interestingly, the extinction appears
t o have a f f e c t e d t r o p i c a l - s u b t r o p i c a l
( M i d c o n t i n e n t ) f a u n a s as w e l l
a s temperate (North A t l a n t i c ) faunas e q u a l l y severely. Accordingly, t h e c a u s e may n o t be s o l e l y a s h a r p d r o p i n water t e m p e r a t u r e a s s o c i a t e d with t h e Late Ordovician g l a c i a t i o n . I t
may b e s i g n i f i c a n t t h a t
t h e g e n e r a s u r v i v i n g i n t o t h e S i l u r i a n were a l l among t h e more w i d e l y d i s t r i b u t e d , a n d t h o s e n o t r e s t r i c t e d t o one p r o v i n c e , d u r i n g t h e O r d o v i c i a n w h i c h s u g g e s t s t h a t t h e s e c o n o d o n t s had t h e c a p a c i t y t o adapt t o a r e l a t i v e l y wide range of environmental c o n d i t i o n s .
SILURIAN
In comparison w i t h t h o s e from t h e Ordovician, S i l u r i a n conod o n t s a r e l e s s w e l l known r e g i o n a l l y i n t h e A t l a n t i c B o r d e r l a n d s , A l s o , p r a c t i c a l l y n o t h i n g h a s b e e n p u b l i s h e d on c o n o d o n t f a u n a s f r o m t h e v e r y l o w e r m o s t p a r t o f t h e s y s t e m . However, j u d g i n g f r o m d a t a c u r r e n t l y a v a i l a b l e , t h e M i d d l e ( W e n l o c k i a n ) a n d Upper ( L u d l o v i a n and P r i d o l i a n ) S i l u r i a n c o n o d o n t s a r e now r e l a t i v e l y w e l l known a s f a r as t h e i r g e n e r a l taxonomy a n d s t r a t i g r a p h i c r a n g e s a r e c o n c e r n e d . A c o n o d o n t - b a s e d z o n a l s c h e m e f o r most o f t h e S i l u r i a n S y s t e m
was i n t r o d u c e d by W a l l i s e r ( 1 9 6 4 ) . I t i s b a s e d l a r g e l y on t h e C e l l o n s e c t i o n i n t h e C a r n i c A l p s o f A u s t r i a . With some m o d i f i c a t i o n s , e s p e c i a l l y i n t h e L l a n d o v e r i a n , Walliser's z o n a l system has proved a p p l i c a b l e world-wide.
The z o n a l s c h e m e s c u r r e n t l y i n u s e i n E u r o p e
and N o r t h A m e r i c a i n c l u d e a b o u t 1 2 a s s e m b l a g e z o n e s ; a c c o r d i n g l y , t h e y a r e l e s s d e t a i l e d t h a n t h a t b a s e d on g r a p t o l i t e s , w h i c h i n c l u d e s a b o u t 32 z o n e s ( B u l m a n , 1 9 7 0 ) .
I n t h e a r e a u n d e r d i s c u s s i o n , S i l u r i a n c o n o d o n t s a r e b e s t known from G r e a t B r i t a i n ( A l d r i d g e , 1 9 7 2 , 1 9 7 5 ) , s o u t h e r n a n d e a s t e r n Sweden ( J e p p s s o n , 1 9 7 5 ) , c e n t r a l E u r o p e (Walliser, 1 9 6 4 ; S c h d n l a u b , 1 9 7 1 ; Walmsley e t a l . ,
1974), Virginia
( H e l f r i c h , 1 9 7 5 ) , and t h e
e a s t e r n M i d c o n t i n e n t o f N o r t h America ( C o o p e r , 1 9 7 5 ; N i c o l l a n d Rexroad, 1968; P o l l o c k and Rexroad, 1973; Rexroad, 1967; Rexroad and Nicoll,
1 9 7 1 ) . S i l u r i a n c o n o d o n s a r e a l S o known f r o m N o r t h A f r i c a
( E t h i n g t o n and F u r n i s h , 1 9 6 2 ) , e a s t e r n Canada ( L e g a u l t , 1 9 6 8 ) , t h e
30 -
-0
J
I
20-
30i
SILURIAN
L
I
F i g . 4 . S i l u r i a n conodont o c c u r r e n c e s i n N o r t h America and n o r t h w e s t e r n Europe p l o t t e d on a s k e t c h - m a p s h o w i n g i n f e r r e d p o s i t i o n o f c o n t i n e n t s i n E a r l y S i l u r i a n ( L l a n d o v e r i a n ) t i m e . Note t h a t p r a c t i c a l l y a l l t h e s e conodont f a u n a s f a l l w i t h i n a b e l t between 2 0 d e g r e e s S o u t h and North l a t i t u d e . I b e r i a n P e n i n s u l a ( K o c k e l , 1958; v a n d e n B o o g a a r d ,
1965), France
( F e i s t a n d S c h B n l a u b , 1 9 7 4 ) and s c a t t e r e d l o c a l i t i e s i n a f e w o t h e r
areas i n t h e A t l a n t i c B o r d e r l a n d s . A r e v i e w of S i l u r i a n c o n o d o n t o c c u r r e n c e s i n t h e A t l a n t i c B o r d e r l a n d s i n N o r t h America a n d n o r t h e r n Europe i s g i v e n i n F i g . 4 . S i l u r i a n conodont f a u n a s ,
a s now known, i n c l u d e 15-20 r n u l t i e l e -
ment g e n e r a and a r e f a r l e s s v a r i e d a t b o t h t h e g e n e r i c a n d s p e c i f i c l e v e l t h a n t h e O r d o v i c i a n o n e s . F u r t h e r , t h e y d o n o t show a n y s t r i k i n g b i o g e o g r a p h l c d i f f e r e n t i a t i o n a l t h o u g h t h e r e a r e some regional d i f f e r e n c e s i n t h e s p e c i f i c composition of t h e faunas
(Jeppsson, 1 9 7 5 ) . Zarliest S i l u r i a n ( e a r l y L l a n d o v e r i a n ) conodonts are as y e t v i r t u a l l y unknown b u t y o u n g e r L l a n d o v e r i a n f a u n a s a r e c h a r a c t e r i z e d by A p s i d o g n a t h u s ,
Carniodus, Distomodus, I c r i o d e l l a , Llandovery-
g n a t h u s , H i n d e o d e l l a ( = O z a r k o d i n a ) , I c r i o d i n a , and Hadrognathus.
The
c o n s i d e r a b l e number o f p l a t f o r m g e n e r a i n t h e L l a n d o v e r i a n f a u n a s i s a n o t a b l e f e a t u r e ; i n younger S i l u r i a n f a u n a s , p l a t f o r m conodont s a r e uncommon a n d l i t t l e v a r i e d , L l a n d o v e r i a n s t r a t a , a s w e l l a s younger S i l u r i a n b e d s , c o n t a i n s e v e r a l wi despr ead
"simple-cone"
97 CONODONT ZONES
SERIES
CARNIC ALPS
PRlDOLlAN
H s
GREAT BRITAIN
H
eosieinhornensis
S
eosreinhornensis
NORTH AMEPICA
H s eosreinhornensis
H crispa / laria/aius
H crispa I laiialaius
LUDLOVIAN
P sr1uricus
i
NOT YET IDENTIFIED
P si/uricus
H sagitia
H sagiiia
NOT YET IDENTIFIED P amorphognarhoides
p amorphognathoides
A ploeckensis H crassa
H segiiia WENLOCKIAN I
K peiule P amorphognathoides
~
P celloni
LLANDOVERIAN
7
/ inconsfans H sieurognarhoides
?
/ discrete
Bereich 1
I deflecia
NOT YET IDENTIFIED
P celloni / irregularis P simplex
?
F i g . 5 . S i l u r i a n c o n o d o n t z o n e s i n E u r o p e a n d K o r t h America. N o t e t h a t t h e p r e c i s e c o r r e l a t i o n b e t w e e n some u n i t s i s s o m e w h a t u n c e r t a i n , p a r t i c u l a r l y i n t h e Lower a n d M i d d l e L l a n d o v e r i a n . g e n e r a b u t t h e s e a r e of m i n o r s i g n i f i c a n c e b i o s t r a t i g r a p h i c a l l y ,
at
l e a s t as f a r a s our p r e s e n t k n o w l e d g e g o e s , Wenlockian and younger S i l u r i a n f a u n a s are dominated by numerous species of Hindeodella
(=Ozarkodina) and Ligonodina b u t s t r a t i g r a p h i -
c a l l y i m p o r t a n t forms o f t h e p l a t f o r m g e n e r a K o c k e l e l l a ,
Pelekys-
g n a t h u s , a n d P o l y g n a t h o i d e s a r e p r e s e n t i n s o m e i n t e r v a l s . A compreh e n s i v e a n d a u t h o r a t i v e r e v i e w e s p e c i a l l y of E u r o p e a n L a t e S i l u r i a n faunas has been given r e c e n t l y by Jeppsson
(1975) a n d a n e x c e l l e n t
summary of t h e known 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 o f B r i t i s h S i l u r i a n c o n o d o n t s h a s b e e n p r e s e n t e d by A l d r i d g e ( 1 9 7 5 ) . T h e i r work c l e a r l y s u g g e s t s t h a t n o t o n l y are c o n o d o n t s v e r y u s e f u l g u i d e fossils i n t h e E u r o p e a n S i l u r i a n , b u t a l s o t h a t t h e p r e s e n t l y u s e d s y s t e m of a b o u t 12 zones c a n b e c o n s i d e r a b l y r e f i n e d after a t h o r o u g h taxonomic r e v i s i o n of t h e f a u n a s .
The commonly u s e d S i l u r i a n c o n o d o n t z o n e s
i n E u r o p e a n d N o r t h America a r e l i s t e d a n d c o r r e l a t e d i n F i g . 5 w h i c h a l s o i l l u s t r a t e s t h e r e l a t i o n s between conodont zones and European s t a n d a r d series.
RELATIONS BETWEEN COUODOST AND GRAPTOLITE ZOSAL UNITS
For a hundred years or more, graptolites have served as the principal guide fossils in regional correlation of Ordovician and Silurian rocks in many parts of the world. Although they are still unsurpassed as biostratigraphic tools in shaly facies, the very scattered occurrence or absence of these fossils in many important carbonate units in America and Eurasia has led to problems in applying the graptolite biostratigraphy to such sequences, especially to those so widely developed on the continental shields. On the
other hand, conodonts are present in a wider r m g e of facies types than are graptolites and they may be used for correlations across facies boundaries. The fact that conodonts are most common, and most easily studied, in shelly rocks makes them particularly useful to compliment graptolites in biostratigraphic work. Considerable efforts have been made in recent years to tie the z o n a l system based on conodonts into that based on graptolites. Much
of this work has been carried out in the Atlantic Borderlands where, at relatively numerous localities, stratigraphically diagnostic conodonts and graptolites have been found together or in such a position that they provide useful information regarding the mutual relations between zonal units based on each of these fossil groups. These relations will be briefly reviewed below.
Cambrian No standard graptolite and conodont zone systems have been established for the Cambrian and the biostratigraphic significance of the dendroid graptolites present in rocks of that system remains unclear.
Ordovician
The striking provincialism exhibited by both conodonts and graptolites in large parts of the Ordovician has greatly complicated the work aimed at establishing as precisely as possible the relations between conodont and graptolite zones. However, as shown by recent summaries (Bergstrsm, 1971b, 1973b), there is now a large number of ties between North Atlantic Province conodont zones and the standard
99 SERIES
NORTH ATLANTIC C O N O D O N T ZCNES
BRITISH GRAPTOLITE
AND SUBZONES ~
ZONES
~~
Dicellogr anceps
ASHGlLLlAN
Amorphognarhus ordovicicus
Dicellogr
complanatus
Pleurogr
linearis
~
Amorohoonathus suDerbus
Dicranoqr clingani
CARADOCIAN
13
L0
3
LLANVIRNIAN
Lower
9
-
ARENlGlAN
?
n z
I
P ygodus ansertnus
LLANDEILIAN
z
I
Eopl reclinatus ~ o p lfoliaceus UNNAMED Eopl suecicus Eoplacognaihus variabilrs Microzarkodine parva Parotstodus originalis Prioniodus navis Prioniodus triangularis Oeoikodus evae Prioniodus elegans , Paroistodus oroteus
Didymogr murchisoni Didymogr Didymogr
1
Didymogr
.
extensus
I
I
(Tetragr approximetus)
4
Anisograptidae
Paltodus deltifer
TREMADOCIAN
btfrdus' hirundo
Clonogr tenellus
Cordylodus angulatus
Dicfyonema flabelliforme
B r i t i s h - B a l t i c g r a p t o l i t e zones. A review of t h e r e l a t i o n s between t h e s e z o n a l s y s t e n s , a s now known,
is given
Irl
i'ig.6.
Some p r o g r e s s
h a s a l s o b e e n made i n e l u c i d a t i n g t h e r e l a t i o n s b e t w e e n N o r t h A t l a n t i c P r o v i n c e conodont zo n es and North Am er i can- P aci f i c
Province
g r a p t o l i t e zon es ( B e r g s t r h , 1971b, 1974, 1976b: Landi ng, 1974a) b u t much more work a l o n g t h o s e l i n e s i s n e e d e d .
This applies a l s o t o
t h e p r o b l e n o f t h e p r e c i s e r e l a t i o n s b e t w e e n Midcontineri-1 P r o v i n c e c o n o d o n t u n i t s a n d g r a p t o l i t e z o n e s , f o r whic!i r e l a t i v e l y l i t t l e d i r e c t e v i d e n c e i s now a v a i l a b l e .
The s c a r c i t y o f z o n a l g r a p t o l i t e s
i n r o c k s w i t h h o r t h American M i d c o n t i n e n t P r o v i n c e conodont f a u n a s w i l l d o u b t l e s s make t h i s work b o t h d i f f i c u l t a n d t i m e - c o n s u m i n g .
Silurian
The many g r a p t o l i t e z o n e s r e c o g n i z e d i n t h i s s y s t e m w i t h i n t h e a r e a d e a l t w i t h h e r e i n a r e , by a n d l a r g e ,
somewhat l e s s s a t i s f a c t o r i -
l y t i e d d i r e c t l y i n t o t h e c o n o d o n t z o n a l s u c c e s s i o n t h a n i s t h e case
100
F i g . 7 . C o r r e l a t i o n between European S i l u r i a n s t a n d a r d s e r i e s , conodont z o n e s , and g r a p t o l i t e zo nes. i n t h e Ordovician.
I n h i s s t a n d a r d work, Walliser ( 1 9 6 4 ) p r e s e n t e d a
p r e l i m i n a r y c o r r e l a t i o n between h i s conodont z o n e s and s t a n d a r d g r a 7 t o l i t e z o n e s t h a t i s s t i l l v a l i d in most r e s p e c t s a l t h o u g h s u b s e q u e n t r e s e a r c h h a s made i t p o s s i b l e t o r e f i n e i t t o a c e r t a i n extent.
However,
t h e r e i s s t i l l some u n c e r t a i n t y r e g a r d i n g t h e
p r e c i s e r e l a t i o n b e t w e e n some c o n o d o n t a n d g r a p t o l i t e z o n e s , e s p e c i a l l y i n t h e L l a n d o v e r i a n , a n d more d e t a i l e d c o n o d o n t work i s c l e a r l y needed i n g r a p t o l i t e - b e a r i n g
s e c t i o n s . Fig.7 is an attempt t o
summarize t h e c u r r e n t l y known r e l a t i o n s b e t w e a
S i l u r i a n conodont
and g r a p t o l i t e zo n es .
In t h e c o m b i n e d g r a p t o l i t e - c o n o d o n t
z o n a l systems w e have a
powerful t o o l t o r e c o g n i z e b i o s t r a t i g r a p h i c u n i t s a c r o s s facies and p r o v i n c i a l b o u n d a r i e s , and t o c o r r e l a t e with unusual precision.
Indeed,
,dinits
3ver l a r g e distaric'-s
t h e r e a r e now i n t e r v a l s w i t h i n w h i c h
t r a n s - A t l a n t i c c o r r e l a t i o n s can be achieved w i t h a s t r a t i g r a p h i c r e s o l u t i o n o f a few f e e t , a n d i t i s t o b e e x p e c t e d t h a t f u r - t h r ? - : r e f i n e m e n t w i l l b e p o s s i b l e i n many i n t e r v a l s when a d d i t i o n a l d a t a
become a v a i l a b l e from c r i t i c a l areas a n d k e y s e c t i o n s . f u t u r e g o a l may b e t o t r y t o e s t a b l i s h
one d e t a i l e d
Obvi ousl y, one
z o n a l framework
b a s e d on a l l s t r a t i g r a p h i c a l l y i m p o r t a n t f o s s i l g r o u p s b u t much remains t o be l e a r n e d about t h e mutual r e l a t i o n s between t h e ranges
of r e p r e s e n t a t i v e s of t h e s e g r o u p s b e f o r e a m e a n i n g f u l a t t e m p t a t s u c h a s y n t h e s i s c a n b e made.
CONOUOKTS AND THE EVOLUTION OF THE PROTO-ATLANTIC
F o s s i l s played a s i g n i f i c a n t r o l e i n t h e e a r l y development of the concept of a Proto-Atlantic
Ocean ( W i l s o n , 1 9 6 6 ) a n d i n d i c a t i o n s
f r o m e s p e c i a l l y b e n t h i c m e g a f o s s i l s s u c h as b r a c h i o p o d s a n d t r i l o b i t e s have a l s o been w i d e l y used i n s u b s e q u e n t d i s c u s s i o n s r e g a r d i n g P a l e o z o i c p l a t e t e c t o n i c s a n d t h e e v o l u t i o n of t h e P r o t o - A t l a n t i c (Williams, 1 9 7 3 ; W h i t t i n g t o n a n d Hughes, 1 9 7 2 , 1 9 7 3 ; B u r r e t t ,
1973).
The d i s t r i b u t i o n o f p r e d o m i n a n t l y p l a n k t i c o r e p i p l a n k t i c o r g a n i s m s s u c h as g r a p t o l i t e s a n d c o n o d o n t s h a s a t t r a c t e d somewhat l e s s i n t e r -
est i n t h e s e d i s c u s s i o n s (Bergstram, 1971a, 1973a, 1976a; Skevington, 1976; B e r g s t r h e t a l . ,
1972, 1 9 7 4 ) . T h i s might b e r e l a t e d t o t h e
f a c t t h a t t h e v e r y w i d e h o r i z o n t a l d i s t r i b u t i o n o f many t a x a o f t h e s e g r o u p s c e r t a i n l y s u g g e s t s t h a t s u c h f o r m s were c a p a b l e o f c r o s s i n g w a t e r b o d i e s t h a t were l a r g e e n o u g h t o s e r v e a s m i g r a t i o n a l b a r r i e r s t o n o s t benthic o r g a n i s m s a n d , a c c o r d i n g l y ,
i t might have been assumed
t h a t t h e s e g r o u p s would b e l e s s l i k e l y t o p r o v i d e u s e f u l d a t a r e g a r ding t h e mutual p o s i t i o n s of t h e c o n t i n e n t a l p l a t e s , T h i s i d e a i s r e e n f o r c e d by t h e f a c t t h a t , a s h a s b e e n n o t e d by Cook a n d T a y l o r ( 1 9 7 5 ) , t h e d e g r e e o f f a u n a l r e s e m b l a n c e b e t w e e n two areas i s n o t n e c e s s a r i l y d i r e c t l y r e l a t e d t o t h e g e o g r a p h i c d i s t a n c e b e t w e e n them, and p r e s u m a b l y ,
t h i s i s l i k e l y t o a p p l y t o planlctic and e p i p l a n k t i c
f o r m s e v e n more t h a n t o b e n a h i c o r g a n i s m s .
However, t h i s s t u d y i s
concerned w i t h a time i n t e r v a l and a g e o g r a p h i c r e g i o n c h a r a c t e r i z e d by v e r y a c t i v e s e a f l o o r s p r e a d i n g and i t i s t h e r e f o r e a p p r o p r i a t e t o c o n s i d e r b r i e f l y t h e r e l a t i o n s between t h e c o n o d o n t f a u n a s i n Europe and N o r t h America and t h e g e n e r a l model o f t h e d e v e l o p m e n t of t h e Prot o-A t l a n t i c
.
Even a c a s u a l l o o k a t t h e t r a n s - A t l a n t i c r e l a t i o n s during Cambro-Silurian
conodont f a u n a l
t i m e shows t h a t t h e r e i s a p p a r e n t l y
no s i m p l e r e l a t i o n b e t w e e n v a r y i n g f a u n a l r e s e m b l a n c e a n d g e o g r a p h i c
proximity of t h e
NQrth-European and North American p l a t e s ,
t h e p e r i o d o f maximum e x p a n s i o n of t h e P r o t o - A t l a n t i c
During
i n t h e Cambrian,
c o n o d o n t f a u n a s show a h i g h d e g r e e o f s i m i l a r i t y r e g i o n a l l y w i t h In Early Ordovician t i m e ,
numerous c o s m o p o l i t a n t a x a .
a striking
p r o v i n c i a l d i f f e r e n t i a t i o n developed, which l a s t e d through v i r t u a l l y the e n t i r e period.
This r e s u l t e d i n t h e establishment of Midcontinent
Province f a u n a s o v e r t h e North American P l a t f o r m a n d t h e s t r i k i n g l y d i f f e r e n t N o r t h A t l a n t i c P r o v i n c e f a u n a s i n n o r t h w e s t e r n Europe a n d , i n t e r e s t i n g l y , i n r e g i o n s a l o n g t h e e a s t e r n and w e s t e r n m a r g i n s o f t h e Iiorth American c r a t o n .
Due t o t h e f a c t t h a t most o f t h e p e r t i n e n t
f a u n a s have n o t y e t been r e v i s e d on m u l t i e l e m e n t b a s i s ,
i t is current-
l y i m p o s s i b l e t o g i v e m e a n i n g f u l n u m e r i c a l v a l u e s of f a u n a l resein-
b l a n c e b e t w e e n t h e two p r o v i n c e s d u r i n g d i f f e r e n t t i m e i n t e r v a l s i n t h e O r d o v i c i a n ; however,
i t a p p e a r s t h a t t h e d i f f e r e n c e s were
e s p e c i a l l y c o n s p i c u o u s d u r i n g Middle O r d o v i c i a n
(Champlainian) t i m e .
T h i s p e r i o d o f maximum c o n o d o n t p r o v i n c i a l i s m i s n o t i n p h a s e w i t h commonly p r o p o s e d m o d e l s f o r t h e e v o l u t i o n o f t h e P r o t o - A t l a n t i c , w h i c h s u g g e s t maximum c o n t i n e n t a l s e p a r a t i o n i n t h e L a t e C a m b r i a n (when c o n o d o n t f a u n a s a r e l a r g e l y c o s m o p o l i t a n ) a n d s u c c e s s i v e c l o s i n g d u r i n g t h e O r d o v i c i a n , t h a t i s , d u r i n g t h e p e r i o d o f most pronounced conodont p r o v i n c i a l i s m .
I t should be n o t e d ,
however,
that
t h e c o s m o p o l i t a n f a u n a s i n t h e S i l u r i a n a r e i n a c c o r d w i t h a model of a t h e n l a r g e l y s u b d u c t e d P r o t o - A t l a n t i c
(Fig.4).
The i m p o r t a n t f a c t t h a t N o r t h A t l a n t i c P r o v i n c e c o n o d o n t f a u n a s a r e p r e s e n t n o t o n l y i n E u r o p e b u t a l s o on t h e A m e r i c a n s i d e o f t h e Proto-Atlantic
i n r e g i o n s c l e a r l y b e l o n g i n g t o t h e N o r t h American
p l a t e ( F i g . 2 ) i n d i c a t e s t h a t t h e Ordovician P r o t o - A t l a n t i c w a s an i n e f f i c i e n t migration b a r r i e r t o t h e s e conodonts. Also, i t is clear t h a t t h e prime f a c t o r s t h a t served t o prevent migration of t h e s e faunas i n t o t h e Midcontinent
P r o v i n c e a r e a f u r t h e r on t o t h e c r a t o n
evidently did not include geographic distance. l y t h a t t h e y were o f e c o l o g i c c h a r a c t e r .
Rather,
c o n t r o l has been d e a l t w i t h r e p e a t e d l y i n r e c e n t y e a r s 1973; Barnes and F s h r a e u s , and Carnes,
i t seems l i k e -
The n a t u r e o f t h i s e c o l o g i c a l
1975; B e r g s t r a m ,
i n p r e s s ; Sweet and BergstrBm,
(Barnes et al.,
1971a, 1973a; Bergstram 1974) and a d i s c u s s i o n of
i t is b e y o n d t h e s c o p e o f t h e p r e s e n t p a p e r .
Y e t i t is o f i n t e r e s t
t o n o t e t h a t t h e s e e c o l o g i c a l f a c t o r s (among w h i c h water t e m p e r a t u r e no d o u b t was i m p o r t a n t ) a p p a r e n t l y a l s o s t r o n g l y i n f l u e n c e d t h e d i s t r i b u t i o n a l patterns of several megafossil groups, including
103 g r a p t o l i t e s and brachiopods. I n v i e w o f t h e a p p a r e n t l y i n s i g n i f i c a n t r o l e p l a y e d by g e o g r a p h ic distance alone vis-a-vis
e c o l o g i c a l f a c t o r s f o r t h e changing
c o n o d o n t d i s t r i b u t i o n a l p a t t e r n s p a r t i c u l a r l y i n t h e Cambro-Ordovici a n , a l o n g w i t h t h e f a c t t h a t v e r y s i m i l a r c o n o d o n t f a u n a s were p r e s e n t on b o t h s i d e s o f t h e P r o t o - A t l a n t i c ,
i t a p p e a r s t h a t cono-
donts are n o t l i k e l y t o p r o v i d e c r i t i c a l d a t a toward a n i n t e r p r e t a t i o n of t h e r e l a t i v e w i d t h o f t h e P r o t o - A t l a n t i c d u r i n g d i f f e r e n t time i n t e r v a l s .
A c c o r d i n g l y , one h as t o b e c o n t e n t w i t h t h e f a c t
t h a t although conodonts are s u p e r i o r t o o l s f o r t r a n s - A t l a n t i c correlations, a t least at the present t i m e they give l i t t l e evidence r e g a r d i n g tk s i z e a n d d e v e l o p m e n t o f t h e P r o t o - A t l a n t i c Ocean.
ACKNOWLEDGEMENTS
I a m indebted t o Dr.
Walter C.
Sweet f o r r e a d i n g t h e m a n u s c r i p t
and o f f e r i n g v a l u a b l e comments a n d t o Mrs. H e l e n J o n e s a n d Mrs. K a r e n Tayler f o r t e c h n i c a l a s s i s t a n c e .
REF EKENCES
1 9 7 2 . L l a n d o v e r y c o n o d o n t s f r o m t h e Welsh B o r d e r l a n d : Aldridge, K.J., B r i t . Mus. ( N a t . H i s t . ) B u l l . G e o l . , 22:2, 125-131. Aldridge, R . J . , 1 9 7 5 . The 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 of c o n o d o n t s i n t h e B r i t i s h S i l u r i a n : Geol. S O C . London J o u r . , 131, 6 0 7 - 6 1 8 . B a r n e s , C . K . , a n d F % h r a e u s , L.F., 1 9 7 5 . P r o v i n c e s , c o m m u n i t i e s , a n d t h e p r o p o s e d n e c t o b e n t h i c h a b i t of O r d o v i c i a n c o n o d o n t o p h o r i d s : L e t h a i a , 8 , 133-149. R e x r o a d , C.B., a n d M i l l e r , J.F., 1 9 7 3 . Lower P a l e o z o i c Barnes, C.R., c o n o d o n t p r o v i n c i a l i s m : G e o l . SOC. A m e r i c a S p e c . P a p e r 141, 157190. B e n g t s s o n , S . , 1 9 7 6 . The s t r u c t u r e of some M i d d l e C a m b r i a n c o n o d o n t s , a n d t h e e a r l y e v o l u t i o n of c o n o d o n t s t r u c t u r e a n d f u n c t i o n : L e t h a i a , 9 , 185-206. B e r g s t r s m , S.M., 1 9 6 2 . C o n o d o n t s f r o m t h e L u d i b u n d u s L i m e s t o n e ( M i d d l e O r d o v i c i a n ) o f t h e T v a r e n area (S.E. S w e d e n ) : Ark. f . M i n e r a l o g i 0 . G e o l . , 3:1, 1 - 6 1 . B e r g s t r b m , S.M., 1 9 6 4 . Remarks o n some O r d o v i c i a n c o n o d o n t f a u n a s f r o m W a l e s : A c t a Univ. L u n d e n s i s , I I : 3 , 1 - 6 6 . B e r g s t r d m , S.M., 1 9 7 1 a . Conodont b i o s t r a t i g r a p h y o f t h e M i d d l e and Upper O r d o v i c i a n o f Europe a n d e a s t e r n N o r t h America: Geo soc. America M e m . 1 2 7 , 8 3 - 1 5 7 . B e r g s t r a m , S.M., 1971b. C o r r e l a t i o n o f t h e N o r t h A t l a n t i c Midd e a n d Upper O r d o v i c i a n c o n o d o n t z o n a t i o n w i t h t h e g r a p t o l i t e s u c c e s s i o n : Mgm. Rech. Ge'ol. e t M i n . , 7 3 , 177-187.
.
104 B e r g s t r b ' m , S.M. 1 9 7 3 a . O r d o v i c i a n c o n o d o n t s . I" H a l l a m , A, (Ed. ) $ A t l a s o f P a l a e o b i o g e o g r a p h y , 4 7 - 5 8 . E l s e v i e r P u b l . Company. 1973b. C o r r e l a t i o n o f t h e l a t e L a s n a n g g i a n S t a g e BergstrSm, S . M . , ( M i d d l e O r d o v i c i a n ) w i t h t h e g r a p t o l i t e s u c c e s s i o n : Geol. F a r e n . S t o c k h o l m F b r h a n d l . , 9 5 , 9-18. 1973c. B i o s t r a t i g r a p h y and f a c i e s r e l a t i o n s i n t h e B e r g s t r i i m , S.M., lower Middle O r d o v i c i a n of e a s t e r n m o s t T e n n e s s e e : A m e r . J o u r . S c i e n c e , 273-A, 2 6 1 - 2 9 3 . B e r g s t r G m , S.M., 1 9 7 4 . O r d o v i c i a n c o r r e l a t i o n s b y means o f n o n b e n t h o n i c f o s s i l s : On t h e r e l a t i o n s b e t w e e n S o r t h A m e r i c a n s t a n d a r d g r a p t o l i t e zo n es and North A t l a n t i c P r o v i n c e conodont z o n e s : G e o l . SOC. A m e r i c a , X b s t r . w i t h P r o g r . , 6 : 7 , 652-653. B e r g s t r h , S.M., 1976a. O r d o v i c i a n c o n o d o n t s and t h e " P r o t o - A t l a n t i c " O c e a n : G e o l . S O C . A m e r i c a , A b s t r . w i t h P r o g r . , 8 : 2 , 133. B e r y s t r E m , S.M. 1 9 7 6 b . The M a r a t h o n M i d d l e a n d Upper O r d o v i c i a n s u c c e s s i o n r e c o n s i d e r e d : Conodont a n d g r a p t o l i t e b i o s t r a t i g r a p h y o f t h e Woods Hollow a n d M a r a v i l l a s F o r m a t i o n s : G e o l . S O C . America, A b s t r . w i t h P r o g r . , 8 : 4 , 4 6 3 - 4 6 4 . i n p r e s s . Conodont b i o s t r a t i g r a p h y B e r g s t r g m , S.M., a n d C a r n e s , J . B . , and paleoecology of t h e Holston Formation (Middle Ordovician) a n d a s s o c i a t e d s t r a t a i n e a s t e r n T e n n e s s e e . 2 B a r n e s , C. R . ( E d . ) , Conodont P a l e o e c o l o g y : G e o l . A s s o c . C a n a d a S p e c . P a p . 1 5 . B e r g s t r a m , S.M. E p s t e i n , A.G. and E p s t e i n , J.B. 1972. E a r l y O r d o v i c i a n N o r t h A t l a n t i c p r o v i n c e c o n o d o n t s i n e a s t e r n Penns y l v a n i a : U . S . G e o l . S u r v . P r o f . P a p . 8 0 0 D , D37-D4?. R i v a l J . , a n d Kay, M . , 1 9 7 4 . S i g n i f i c a n c e o f c o n o B e r g s t r z m , S.M., d o n t s , g r a p t o l i t e s , and s h e l l y f a u n a s from t h e O r d o v i c i a n of w e s t e r n a n d n o r t h - c e n t r a l N e w f o u n d l a n d : Can. J o u r . E a r t h S c i . , 11:12, 1 6 2 5 - 1 6 6 0 . 1966. Conodonts f r o m t h e L e x i n g t o n B e r g s t r E m , S.M., a n d S w e e t , W.C., L i m e s t o n e ( M i d d l e O r d o v i c i a n ) of K e n t u c k y a n d i t s l a t e r a l e q u i v a l e n t s i n O h i o a n d I n d i a n a : B u l l . A m e r . P a l . , 5 0 : 2 2 9 , 271441. van den Boogaard, M., 1 9 6 5 . Two c o n o d o n t f a u n a s f r o m t h e P a l e o z o i c o f t h e B e t i c o f M a l a q a n e a r V 6 l e z R u b i o , S.E. S p a i n : K o n i n k l . N e d e r l . Akad. W e t e n s c h . P r o c . S e r . B, 6 8 , 3 3 - 3 7 . B r a d s h a w , L.E., 1969. C o n o d o n t s f r o m t h e F o r t Pexa F o r m a t i o n ( M i d d l e O r d o v i c i a n ) , M a r a t h o n B a s i n , Texas: J o u r . P a l e o n t . , 4 3 , 1 1 3 7 1168. B r a n d , U., 1 9 7 6 . Lower O r d o v i c i a n c o n o d o n t s f r o m t h e K i n d b l a d e F o r m a t i o n , A r b u c k l e M o u n t a i n s , Oklahoma: U n p u b l . M . A . t h e s i s , U n i v . o f M i s s o u r i , C o l u m b i a , 110 p p . B u l n a n , O.M.B., 1970. G r a p t o l i t h i n a w i t h s e c t i o n s on E n t e r e p n e u s t a and P t e r o b r a n c h i a . & T e i c h e r t , C. ( E d . ) , T r e a t i s e o n I n v e r t e b r a t e P a l e o n t o l o g y , V ( 2 n d e d . ) , 1 6 3 pp. B u r r e t t , C . , 1 9 7 3 . O r d o v i c i a n b i o g e o g r a p h y and c o n t i n e n t a l d r i f t : P a l a e o g e o g r . , P a l a e o c l i m . , P a l a e o e c o l . , 13, 1 6 1 - 2 0 1 . Cook, H.E., a n d T a y l o r , M.E., 1 9 7 5 . E a r l y P a l e o z o i c c o n t i n e n t a l margin s e d i m e n t a t i o n , t r i l o b i t e b i o f a c i e s , and t h e t h e r m o c l i n e , w e s t e r n U n i t e d S t a t e s : G e o l o g y , 3:10, 5 5 9 - 5 6 2 . Cooper, B.J., 1975. M u l t i e l e m e n t co n o dont s from t h e B r a s s f i e l d L i m e s t o n e ( S i l u r i a n ) of s o u t h e r n O h i o : J o u r . P a l e o n t . 49, 98 4- 1008. D r u c e , E.C., a n d J o n e s , P . J . , 1 9 7 1 . C a m b r o - O r d o v i c i a n c o n o d o n t s f r o m t h e B u r k e R i v e r s t r u c t u r a l b e l t , Q u e e n s l a n d : Bur. Min. R e s o u r . G e o l . G e o p h y s . B u l l . 110, 118 p ~ . E t h i n g t o n , R.L., a n d C l a r k , D.L., 1 9 7 1 . Lower O r d o v i c i a n c o n o d o n t s i n N o r t h America: G e o l . S O C . America M e m . 1 2 7 , 6 3 - 8 2 . ~
,
,
,
,
,
105 E t h i n q t o n , R . L . , a n d F u r n i s h , li., 1 9 6 2 . S i l u r i a n a n d D e v o n i a n c o n o d o n t s from S p a n i s h S a h a r a : J o u r . P a l e o n t . , 36, 1253-1290. Fzhraeus, L. E . , 1 9 7 0 . C o n o d o n t - b a s e d c o r r e l a t i o n s o f Lower a n d M i d d l e O r d o v i c i a n s t r a t a i n w e s t e r n N e w f o u n d l a n d : G e o l . S O C . America Bdll., 8 1 , 2 0 6 1 - 2 0 7 6 . F z h r n e u s , L.E., 1973. D e p o s i t i o n a l e n v i r o n n e n t s and c o n o d o n t - b a s e d c o r r e l a t i o n s o f t h e Long P o i n t F o r m a t i o n ( M i d d l e O r d o v i c i a n ) w e s t e r n S e w f o u n d l a n d : Can. J o u r . E a r t h S c i . , 10, 1822-18.33. 1 9 7 4 . Zur S i l u r / D e v o n - G r e n z r i n F e i s t , R . , and S c h i j n l a u b , H . - P . , 8 s t l i c h e n Montagne K o i r e S G d - F r a n k r e i c h s : Neues J a h r b . G e o l . P a l a o n t . Monatsh., J a h r g . 1 9 7 4 , 4 , 200-219. F u g a n t i , A., a n d S e r p a g l i , E . , 1 9 6 8 . G e o l o g i c a l r e m a r k s on U r b a n a L i m e s t o n e a n d e v i d e n c e for i t s Upper O r d o v i c i a n a g e b y means o f c o n o d o n t s - e a s t e r n S i e r r a Elorena, S o u t h S p a i n : G e o l . S O C , I t a l . B o l l . , 8 7 , 511-521. H a m a r , G . , 1 9 6 4 . The b l i d d l e O r d o v i c i a n o f t h e Oslo R e g i o n , Norway. 17. Conodonts from t h e lower Middle O r d o v i c i a n o f R i n g e r i k e : Korsk G e o l . T i d s s k r . , 44, 243-292. Ha.nar, G . , 1 9 6 6 . The M i d d l e O r d o v i c i a n o f t h e Oslo R e g i o n , Norway, 22. P r e l i m i n a r y r e p o r t on co n o d o n t s from t h e Oslo-Asker and R i n g e r i k e d i s t r i c t s : Norslc G e o l . T i d s s k r . , 4 6 , 2 7 - 8 3 . S e l f r i c h , C . T . , 1 9 7 5 . S i l u r i a n c o n o d o n t s f r o m \Gills M o u n t a i n a n t i c l i n e , V i r g i n i a , West V i r g i n i a , a n d M a r y l a n d : G e o l . S O C . A m e r . S p e c . P a p e r 1 6 1 , 8 2 pp.+ a p p e n d i x . J e p p s s o n , L . , 1 9 7 5 . A s p e c t s of S i l u r i a n c o n o d o n t s : F o s s i l s and S t r a t a 6 , 54 pp. KnGpfer, J., 1 9 6 7 . Z u r Fauna und B i o s t r a t i g r a p h i e d e s O r d o v i c i u m s ( G r s f e n t h a l e r S c h i c h t e n ) i n ThGringen: F r e i b e r g e r ForschungsH e f t e , C220, 119 p p . K o c k e l , F . , 1 9 5 8 . C o n o d o n t e n a u s dern P a l a o z o i k u m von M a l a g a (SiidS p a n i e n ) : Seues J a h r b . Geol. P a l s o n t . Monatsh., Jahrg.,225-263. K o h u t , J. J . , 1 9 7 2 . Conodont b i o s t r a t i g r a p h y o f t h e Lower O r d o v i c i a n O r t h o c e r a s a n d S t e i n L i m e s t o n e s ( 3 c ) , Norway: N o r s k G e o l . T i d s s k r . , 5 2 , 427-445. Lamont, A . , a n d L i n d s t r B n , b l . , 1 9 5 7 . A r e n i g i a n and L l a n d e i l i a n c h e r t s i d e n t i f i e d i n t h e S o u t h e r n U p l a n d s o f S c o t l a n d b y means o f c o n o d o n t s , e t c . : E d i n b u r g h G e o l . S O C , T r a n s , , 17:1, 6 0 - 7 0 . L a n d i n g , E., 1 9 7 4 a . Lower O r d o v i c i a n c o n o d o n t a n d g r a p t o l i t e b i o s t r a t i g r a p h y o f t h e T a c o n i c P r o v i n c e , e a s t e r n N e w York: Geol. S O C . America, A b s t r . w i t h P r o g r . , 6 : 6 , 5 2 5 - 5 2 6 . L a n d i n g , E . , 1 9 7 4 b . E a r l y a n d M i d d l e C a m b r i a n c o n o d o r i t s from t h e T a c o n i c A l l o c h t h o n , e a s t e r n Sew York : J o u r . P a l e o n t . , 48 12411248. L a n d i n g , E . , 1 9 7 6 . L a t e C a m b r i a n P r o o n e o t o d u s t e n u i s (Miiller ) a p p a r a t u s e s and a s s o c i a t e d conodonts from t h e Taconic Allochthon, e a s t e r n N e w York: G e o l . S O C . A m e r i c a , A b s t r . w i t h P r o g r . , 8 : 4 , 487-488. 1970. Conodonten a u s d e r Choson-Gruppe ( U n t e r e s O r d o v i z Lee, H . - Y . , i u m ) von K o r e a : Neues J a h r b . G e o l . P a l z o n t . A b h . , 1 3 6 : 3 , 303344. L e e , H.-Y., 1 9 7 5 . C o n o d o n t e n aus dem u n t e r e n und n i t t l e r e n O r d o v i z ium von N o r d k o r e a : P a l a e o n t o g r a p h i c a , A b t . A , 1 5 0 , 1 6 1 - 1 8 6 . 1968. Conodonts and f i s h r e m a i n s from t h e S t o n e h o u s e L e g a u l t , J.A., F o r m a t i o n , A r i s a i g , Nova S c o t i a : C a n a d a G e o l . S u r v . B u l l . 1 6 5 , 3 0 ??. L i n d s t r a m , M., 1955. Conodonts from t h e lowermost O r d o v i c i a n s t r a t a o f s o u t h - c e n t r a l Sweden: Geol. F a r e n . S t o c k h o l n F 6 r h a n d l . , 7 6 , 517-614.
,
106 L i n d s t r b n , >I., 1959. Conodonts f r o m t h e Crdg L i m e s t o n e ( O r d o v i c i a n , Wales) : M i c r o p a l e o n t o l o g y , 5 , 4 2 7 - 4 5 2 . L i n d s t r i j n , bl., 1960. A Lower-Middle O r d o v i c i a n s u c c e s s i o n o f c o n o d o n t f a u n a s : I n t e r n a t . Geol. Congr., XXI S e s s . , Repts., 7 , 88-96. L i n d s t r B n , b l . , 1 9 7 1 . Lower O r d o v i c i a n c o n o d o n t s o f E u r o p e : G e o l . S O C . A m e r i c a M e m . 1 2 7 , 21-61. L i n d s t r B m , M., R a c h e b o e f , P. R . , a n d H e n r y , J . - L . , 1 9 7 4 . O r d o v i c i a n conodonts from t h e P o s t o l o n n e c Formation (Crozon P e n i n s u l a , Massif A r m o r i c a i n ) a n d t h e i r s t r a t i g r a p h i c s i g n i f i c a n c e : G e o l . e t P a l a e o n t . , 8 , 15-28. 1975. S m a l l s h e l l y f o s s i l s ?.!atthews, S . C . , a n d M i s s a r z h e v s k y , V . V . , o f l a t e P r e c a m S r i a n and e a r l y Cambrian a g e : a r e v i e w o f r e c e n t work: G e o l . S O C . Londsn J o u r . , 131, 2 8 9 - 3 0 4 . a n d Opdyke, K . D . , 1973. R e m a g n e t i z a t i o n h y p o t h e s i s blcElhinny, M.W., d i s c o u i t e d : a pxleomagnetic s t u d y of t h e Trenton Limestone, N e w York S t a t e : G e o l . S O C . America B u l l . , 8 4 , 3 6 9 7 - 3 7 0 8 . b l i l l e r , J . F . , 1 9 6 9 . Conodont f a u n a of t h e Notch Peak L i m e s t o n e ( C a m b r o - O r d o v i c i a n ) , House R a n g e , U t a h : J o u r . P a l e o n t . , 4 3 , 413-4.39. Miller, J . F . , 1975. Conodont f a u n a s from t h e Cambrian a n d l o w e s t O r d o v i c i a n o f w e s t e r n N o r t h America: G e o l , S O C . America, A b s t r . w i t h P r o g r . , 7 : 7 , 1200-1201. Miller, J.F., a n d R u s h t o n , A . k ' . A . , 1973. K a t u r a l c o n o d o n t a s s e m b l a g e s f r o m t h e Upper C a m b r i a n o f W a r w i c k s h i r e , G r e a t B r i t a i n : G e o l . S O C . America, A b s t r . w i t h P r o g r . , 5 : 4 , 3 3 8 - 3 3 9 . 1973. Conodontiforin o r g a n i s m s from b e d s c l o s e Nissarzhevsky, V.V., t o t h e Precambrian-Cambrian boundary on t h e S i b e r i a n P l a t f o r m (Ed. ) , P a l e o n t o l o g i c a l a n d i n K a z a k h s t a n . _Ir! Z h u r a l e v a , I . T . a n d b i o s t r a t i g r a p h i c p r o b l e m s i n t h e Lower C a m b r i a n o f S i b i r i a and t h e F a r East, 5 7 - 5 9 . "Nauka"Pub1. H o u s e , N o v o s i b i r s k . ( i n Russian). blound, >!., 1 9 6 5 . A c o n o d o n t f a u n a f r o m t h e J o i n s F o r m a t i o n ( O r d o v i c i a i l ) , Oklahoma: T u l a n e S t u d i e s G e o l . , 4 , 1-46. Mound, M., 1 9 6 8 . C o n o d o n t s a n d b i o s t r a t i g r a p h y o f t h e Lower A r b u c k l e Group ( O r d o v i c i a n ) , A r b u c k l e M o u n t a i n s , Oklahoma: M i c r o p a l e o n t . 14:4, 3 9 3 - 4 3 4 . bliiller, K . J . , 1 9 5 9 . K a m b r i s c h e C o n o d o n t e n : Z e i t s c h r . D e u t s c h . G e o l . G e s . , 111, 4 3 5 - 4 8 5 . bliiller, K.J., 1 9 7 3 . L a t e Cambrian and E a r l y O r d o v i c i a n c o n o d o n t s f r o m n o r t h e r n I r a n : G e o l . S u r v . I r a n , Repts., 3 0 , 7 7 p p . S i c o l l , R.S., and Rexroad, C.B., 1968. S t r a t i g r a p h y and conodont p a l e o n t o l o g y o f t h e S a l a m o n i e D o l o m i t e a n d L e e C r e e k Member of t h e B r a s s f i e l d L i m e s t o n e ( S i l u r i a n ) i n s o u t h e a s t e r n I n d i a n a a n d a d j a c e n t K e n t u c k y : I n d i a n a G e o l . S u r v . B u l l . 40, 7 3 p p . Kogami, Y., 1 9 6 6 , K a m b r i s c h e C o n o d o n t e n von C h i n a . T e i l 1. C o n o d o n t e n a u s d e n o b e r k a m b r i s c h e n K u s h a n - S c h i c h t e n : C o l l . S c i . Univ. K y o t o , M e m . , 3 2 , 351-367. Nogami, Y . , 1 9 6 7 . K a m b r i s c h e C o n o d o n t e n v o n C h i n a , T e i l 2 . C o n o d o n t e n a u s den hoch o b e r k a m b r i s c h e n Y e n c h o - S c h i c h t e n : C o l l . S c i . Univ. K y o t o , M e m . , 33, 2 1 1 - 2 1 8 . and B e r g s t r i j m , S.M., 1 9 7 6 . P a l e o m a g n e t i c s t u d i e s O f S o l t i m i e r , H.C., E a r l y and Middle O r d o v i c i a n l i m e s t o n e s f r om t h e B a l t i c S h i e l d : G e o l . S O C . America, A b s t r . w i t h P r o g r . , 8 : 4 , 501. P o l l o c k , C . A . , and Rexroad, C.B., 1973. Conodonts f r o m t h e S a l i n a F o r m a t i o n a n d t h e u p p e r p a r t o f t h e Wabash F o r m a t i o n ( S i l u r i a n ) i n N o r t h - C e n t r a l I n d i a n a : Geol. e t P a l a e o n t . , 7 , 7 7 - 9 2 . 1 9 7 5 . Lower O r d o v i c i a n c o n o d o n t s o f t h e u p p e r West P o t t e r , C.A., S p r i n g C r e e k F o r m a t i o n , A r b u c k l e M o u n t a i n s , Oklahoma: U n p u b l . >l.A. t h e s i s , U n i v . of M i s s o u r i , C o l u m b i a , 133 p p .
107 P o u l s e n , V . , 1966. E a r l y C a m b r i a n d i s t a c o d o n t i d c o n o d o n t s f r o m B o r n h o l m : B i o l . Medd. K o n g l . Dan. V i d e n s k . S e l s k . , 23:15, 1 - 1 2 . Repetski, J.E., 1974. C o n o d o n t s from t h e Lower O r d o v i c i a n E l P a s o G r o u p of l i ' e s t T e x a s : G e o l . S O C . A m e r i c a , A b s t r . w i t h P r o g r a m . ,
6:6, 540. R e x r o a d , C . B . , 1967. S t r a t i g r a p h y a n d c o n o d o n t p a l e o n t o l o g y o f t h e B r a s s f i e l d ( S i l u r i a n ) i n t h e C i n c i n n a t i Arch area: I n d i a n a G e o l . S u r v . B u l l . 3 6 , 6 4 pp. R e x r o a d , C.B., a n d S i c o l l , R . S . , 1971. Summary o f c o n o d o n t b i o s t r a t i g r a p h y of t h e S i l u r i a n S y s t e m of S o r t h A m e r i c a : G e o l . S O C . America M e m . 127, 207-225. Rhodes, F.H.T., 1953. Some B r i t i s h L o w e r P a l e o z o i c c o n o d o n t f a u n a s : Roy. S O C . L o n d o n P h i l . T r a n s . , S e r . B , 647, 237, 261-333. Rhodrs, F.H.T., 1955. T h e c o n o d o n t f a u n a of tile K e i s l e y L i m e s t o n e : G e o l . S O C . L o n d o n , Q u a r t . J o u r . , 11, 117-142. 1971. Z u r P r o b l e m a t i k d e r C o n o d o n t e n - C h r o n o l o g i e S c h z n l a u b , H.-P., an d e r Wende O r d o v i z / S i l u r m i t b e s o n d e r e r B e r G c k s i c h t i g u n g d e r V e r h z l t n i s s e i m L l a n d o v e r y : G e o l . e t P a l e o n t . , 5, 35-57. Schopf, T.J.M., 1966. C o n o d o n t s of t h e T r e n t o n G r o u p ( O r d o v i c i a n ) i n Yew Y o r k , s o u t h e r n O n t a r i o , a n d Q u e b e c : N e w York S t a t e lilus.
Bull., 405, 105 pp. S e r g e e v a , S . P . , 1964. On t h e s i g n i f i c a n c e of t h e Lower O r d o v i c i a n c o n o d o n t s i n t h e L e n i n g r a d r e g i o n : L e n i n g r a d Univ. V e s t n i k , S e r . G e o l o g i y a i G e o g r a f i i , 12:2, 56-60. ( i n R u s s i a n ) . S e r p a g l i , E . , 1967. I c o n o d o n t i d e l l ' O r d o v i c i a n o S u p e r i o r e ( A s h g i l l i a n o ) d e l l e A l p i C a r n i c h e : S O C , P a l e o n t . I t a l . Bo11.,6, 30-111. S e r p a g l i , E . , 1974, L o w e r O r d o v i c i a n c o n o d o n t s f r o m P r e c o r d i l l e r a n > A r g e n t i n a ( P r o v i n c e of S a n J u a n ) : S O C . P a l e o n t . I t a l . Boll., 13, 17-98. S k e v i n g t o n , D . , 1976. P r o t o c h o r d a t e s , p a l a e o l a t i t u d e s a n d t h e P r o t o A t l a n t i c O c e a n : G e o l . S O C . America, A b s t r . w i t h P r o g r . , 8 : 2 ,
269. S w e e t , W.C., i n p r e s s . C o n o d o n t s a n d c o n o d o n t b i o s t r a t i g r a p h y o f p o s t - T y r o n e O r d o v i c i a n r o c k s o f t h e C i n c i n n a t i r e g i o n : U. S . G e o l . Surv. Prof. Paper. S w e e t , W.C., a n d d e r g s t r a m , S.M., 1 9 6 2 . C o n o d o n t s f r o ? t h e P r a t t F e r r y F o r m a t i o n ( M i d d l e O r d o v i c i a n ) of A l a b a m a : J o u r . P a l e o n . i . ,
36, 1214-1252. W.C., a n d B e r g s t r a m , S.M., 1971. Sy:lposium o n c o n o d o n t b i o s t r a t i g r a p h y : G e o l . S O C . America l i l e m . 127, 499 p p . S t v e e t , W.C., a n d B e r g s t r d m , S . N . , 1974. P r o v i n c i a l i s m e x h i b i t e d b y O r d o v i c i a n c o n o d o n t f a u n a s : SOC. Econ. P a l e o n t . Min. S p e c . P u b l . 2 1 , 189-202. S w e e t , W.C., a n d B e r g s t r B m , S.M., i n p r e s s . C o n o d o n t b i o s t r a t i g r a p h y o f t h e M i d d l e a n d Upper O r d o v i c i a n of t h e m i d c o n t i n e n t of t h e U n i t e d S t a t e s : P a l a e o n t o l . Assoc. S p e c . P a p e r . S w e e t , h'.C., E t h i n g t o n , R . L . , a n d B a r n e s , C . R . , 1971. S o r t h A m e r i c a n b l i d d l e a n d U p p e r O r d o v i c i a n c o n o d o n t f a u n a s : G e o l . S O C . America S\veet,
Slem.
127, 163-193.
S z a n i a w s k i , H . , 1971. N e w s p e c i e s o f U p p e r C a m b r i a n c o n o d o n t s f r o m P o l a n d : ‘\eta G e o l , P o l o n i c a , 16, 401-413. V i i r a , V . , 1967. O r d o v i c i a n c o n o d o n t s u c c e s s i o n i n t h e O h e s a a r e c o r e : E e s t i NSV T e a d , A k a d . T o i m e t i s e d , 1 6 , 319-329. L'iira, V . , 1975. O r d o v i c i a n c o n o d o n t s of t h e E a s t B a l t i c : E e s t i S S V T e a d . .Akad. G e o l . I n s t . T a l l i n n , 142 pp. ( i n R u s s i a n ) . K a l l i s e r , O.H., 1 9 6 4 . C o n o d o n t e n d e s S i l u r s : H e s s . L.-Amt B o d e n f o r s c h . A b h . , 41, 106 pp. I i ' a l m s l e y , V.G., A l d r i d g e , R. J . , a n d A u s t i n , R . L . , 1974. B r a c h i o p o d
108 a n d c o n o d o n t f a u n a s f r o m t h e S i l u r i a n a n d Lower D e v o n i a n of Bohemia: G e o l . e t P a l e o n t . , 8 , 39-47. v a n liamel, W.A., 1974. Conodont b i o s t r a t i g r a p h y of,.the Upper Camb r i a n a n d Lower O r d o v i c i a n of n o r t h - w e s t e r n O l a n d , s o u t h e a s t e r n Sweden: U t r e c h t M i c r o p a l e o n t . B u l l . 10, 126 p p . W e b e r s , G.F., 1 9 6 6 . The M i d d l e a n d Upper O r d o v i c i a n c o n o d o n t f a u n a s of Minnesota: Minnesota Geol. S u r v. Spec. Publ . SP-4, 123 pp. W h i t t i n g t o n , H . B . , a n d Hughes, C.P., 1 9 7 2 . O r d o v i c i a n g e o g r a p h y a n d f a u n a l p r o v i n c e s deduced from t r i l o b i t e d i s t r i b u t i o n : Royal S O C . London P h i l . T r a n s . , B263, 235-278. W h i t t i n g t o n , H . B . , a n d Hughes, C . P . , 1 9 7 3 . O r d o v i c i a n t r i l o b i t e d i s t r i b u t i o n and g eo g rap h y : P a l a e o n t o l . Assoc. Spec. Pap., 12, 2 35- 2 40. W i l l i a m s , A . , 1973. D i s t r i b u t i o n o f b r a c h i o p o d a s s e m b l a g e s i n r e l a t i o n t o Ordovician palaeogeography: P a l a e o n t . Assoc. Spec. P a p . , 1 2 , 241-269. Wilson, J . T . , 1 9 6 6 . Did t h e A t l a n t i c c l o s e a n d t h e n r e - o p e n ? N a t u r e , London, 2 1 1 , 6 7 6 - 6 8 1 .
Dr.
I. G.
Sohn:
Discussion Do you c o n s i d e r t h e d i v e r s i t y of c o n o d o n t s t o
be g r e a t e r i n d e e p t h a n i n s h a l l o w w a t e r ? S . M. Bergstrzm: I t i s d i f f i c u l t t o g i v e a d e f i n i t e answer
Dr.
t o t h i s q u e s t i o n b u t a s a r u l e , f a u n a s were more d i v e r s i f i e d i n somewhat d e e p e r t h a n i n v e r y s h a l l o w w a t e r .
In, for in-
s t a n c e , t h e North American Middle O r d o v i c i a n , t h e d i v e r s i t y i n conodont f a u n a s i n c r e a s e s c o n s i d e r a b l y from t h e c e n t r a l p a r t of t h e c o n t i n e n t t o t h e A p p a l a c h i a n s .
It is especially
s t r i k i n g t h a t O r d o v i c i a n p l a t f o r m c o n o d o n t s were much more a b u n d a n t and v a r i e d i n t h e presumably somewhat d e e p e r w a t e r along t h e c p n t i n e n t a l margins t h a n i n t h e s h a l l o w e p i c o n t i n e n t a l s e a s on t h e p l a t f o r m i t s e l f . Dr.
B.
K.
Holdsworth:
1.
What e v i d e n c e i s t h e r e f o r c o n s i d e r -
i n g t h e D a l r a d i a n p a r t of t h e American p l a t e i n Cambrian t i m e , a s t h e o n l y known t r i l o b i t e s d o n o t n e c e s s a r i l y seem t o b e of t h e American p r o v i n c e ? What e v i d e n c e e x i s t s f o r an O r d o v i c i a n ocean e x t e n d i n g n o r t h w a r d s i n t o S.W. E n g l a n d , a l l F r a n c e and w e l l i n t o cen2.
t r a l Europe? Bergstrgm:
A s f a r a s t h e f i r s t q u e s t i o n i s c o n c e r n e d , I am
aware of t h e weaknesses of some of t h e d a t a b e h i n d t h e Cambrian reconstruction.
A s you p c i n t o u t , f a u n a l and g e o l o g i c
e v i d e n c e f o r p l a c i n g p a r t s of t h e B r i t i s h I s l e s , e t c . on one or t h e other s i d e of the Proto-Atlantic i s not very strong and a t t h e p r e s e n t t i m e , i t i s c e r t a i n l y n o t s u f f i c i e n t f o r
109
making any f i r m judgements a s t o t h e f o r m e r g e o g r a p h i c p o s i t i o n s of t h e s e a r e a s .
The p o s i t i o n of t h e B a l t i c S h i e l d i n
my map i s b a s e d on p a l e o m a g n e t i c measurements on M i d d l e Camb r i a n r o c k s t h a t w e have c a r r i e d o u t r e c e n t l y , and I have a d o p t e d t h e c o n v e n t i o n a l view and k e p t p a r t o f t h e B r i t i s h I s l e s -- The E n g l i s h M i d l a n d s , e t c . -- i n a b o u t t h e same geographic p o s i t i o n t o t h e B a l t i c Shield as today.
But I h a v e
no s t r o n g f e e l i n g s r e g a r d i n g where S c o t l a n d i s l i k e l y t o h a v e I b e l i e v e , however, t h a t t h e m a r i n e been i n Cambrian t i m e . Lower P a l e o z o i c s e d i m e n t s o n t h e H e b r i d e a n P l a t f o r m i n n o r t h -
w e s t e r n m o s t S c o t l a n d , f o r i n s t a n c e , t h e Durness L i m e s t o n e , show so c l o s e a f f i n i t i e s w i t h e q u i v a l e n t s e d i m e n t s i n e a s t e r n most N o r t h America, G r e e n l a n d , S p i t s b e r g e n , e t c . t h a t i t i s r e a s o n a b l e t o s u g g e s t t h a t t h e y a l l b e l o n g t o t h e same g e n e r a l province. A s f a r a s t h e s e c o n d q u e s t i o n i s c o n c e r n e d , I have p r e f e r r e d , i n t h e model of t h e E a r l y O r d o v i c i a n P r o t o - A t l a n t i c , t o l e a v e o u t from t h e map C o r n w a l l and t h e o t h e r H e r c y n i a n a r e a s i n t h e S W p a r t of t h e B r i t i s h I s l e s .
I personally believe
t h a t t h e r e w a s a mid-European sea e x t e n d i n g i n t o t h a t area a s s u g g e s t e d by o t h e r a u t h o r s , b u t I do n o t know i f C o r n w a l l
w a s a p a r t of Great B r i t a i n o r n o t d u r i n g E a r l y O r d o v i c i a n
time. Dr.
J. W.
Neale:
I f I c a n f o l l o w up w h a t D r .
Holdsworth s a i d ,
I w a s r a t h e r w o r r i e d a b o u t some of y o u r p l a t e t e c t o n i c r e c o n -
structions.
I n terms of t h e c o n o d o n t s , I n o t i c e d t h i s p a r -
t i c u l a r l y i n y o u r L l a n d e i l i a n r e c o n s t r u c t i o n where you had your American and N o r t h A t l a n t i c f a u n a s s h a r p l y s e p a r a t e d somewhere i n t h e r e g i o n of t h e A p p a l a c h i a n s .
A t t h e same
t i m e , t h e p r o t o - A t l a n t i c Ocean a p p a r e n t l y had no e f f e c t on t h e e a s e o f d i s s e m i n a t i o n of c o n o d o n t s from t h e European a r e a s t o t h e e a s t American a r e a . I n t h i s i n s t a n c e i f y o u r p r o t o - o c e a n i s no b a r r i e r t o y o u r N o r t h A t l a n t i c f a u n a , what c o n t r o l s t h e s h a r p d i v i s i o n between y o u r N o r t h A t l a n t i c f a u n a and y o u r American f a u n a i n t h e A p p a l a c h i a n a r e a ?
In
o t h e r w o r d s , would i t n o t b e b e t t e r t o a t t a c h t h e e a s t Americ a n p a r t of your p l a t e t o t h e European a r e a a t t h a t t i m e ? Bergstram: T h i s i s a v e r y i n t e r e s t i n g q u e s t i o n and a problem on w h i c h , a s you know, I have b e e n working f o r s e v e r a l y e a r s . I t i s
110
s t r i k i n g how s i m i l a r t h e Lower-Middle O r d o v i c i a n c o n o d o n t faunas i n t h e e a s t e r n p a r t of t h e Appalachians are t o t h o s e i n Europe. by s p e c i e s .
I n many cases o n e c a n s i m p l y m a t c h f a u n a s s p e c i e s On t h e o t h e r h a n d , i f o n e g o e s t o t h e w e s t e r n
Appalachians t w e n t y - t h i r t y miles t o t h e w e s t , one encounters M i d c o n t i n e n t - t y p e c o n o d o n t f a u n a s w h i c h a r e v i r t u a l l y completely different.
F o r a w h i l e , I was i n c l i n e d t o b e l i e v e
t h a t maybe, t h e r e w a s a p l a t e b o u n d a r y b e t w e e n t h e s e two conodont f a u n a l provinces.
However, i f o n e s t u d i e s t h e l o c a l
Ordovician geology i n Tennessee, V i r g i n i a , e t c . , one f i n d s
-
t h a t e v e n i f t h e s e r o c k s h a v e b e e n t h r u s t e d some d i s t a n c e
probably a r a t h e r s h o r t d i s t a n c e - t o t h e northwest, one can t r a c e many f o r m a t i o n s from t h e w e s t e r n A p p a l a c h i a n s i n t o a t l e a s t t h e c e n t r a l p a r t of t h e A p p a l a c h i a n V a l l e y ; a c c o r d i n g l y , t h e r e a p p e a r s t o b e no r e a s o n t o r e g a r d t h e c o n o d o n t f a u n a l p r o v i n c i a l boundary as c o i n c i d i n g w i t h a p l a t e boundary.
Ac-
t u a l l y , t h e r e i s a zone o f t r a n s i t i o n t h a t may b e o n e o r t w o t h r u s t b e l t s w i d e i n w h i c h some N o r t h A t l a n t i c p r o v i n c e e l e m e n t s a r e mixed w i t h r e p r e s e n t a t i v e s of t h e M i d c o n t i n e n t p r o v i n c e , and t h i s r a t h e r n a r r o w b e l t h a s b e e n n o t e d l o c a l l y from Alabama a l l t h e way u p t o Maryland i n t h e c e n t r a l p a r t of t h e A p p a l a c h i a n V a l l e y . I t h i n k t h i s f a u n a l d i f f e r e n t i a t i o n i n t h e Appalachian Val-
l e y i s l i k e l y t o h a v e b e e n c a u s e d by e c o l o g i c a l c o n t r o l , n o t by g e o g r a p h i c s e p a r a t i o n .
I believe t h a t basic controlling
f a c t o r s were w a t e r t e m p e r a t u r e and w a t e r d e p t h .
According t o
r e c e n t l y p u b l i s h e d r e c o n s t r u c t i o n s by Walker and o t h e r s of s e d i m e n t a r y e n v i r o n m e n t s i n t h e M i d d l e O r d o v i c i a n of e a s t e r n T e n n e s s e e , t h e c r i t i c a l a r e a was n o t o n t h e c o n t i n e n t a l s l o p e b u t on t h e o u t e r p a r t o f t h e p l a t f o r m .
The N o r t h A t l a n t i c
c o n o d o n t s p r e f e r r e d t h e somewhat d e e p e r w a t e r a l o n g t h e cont i n e n t a l m a r g i n , and w i t h few e x c e p t i o n s , t h e y d i d n o t i n h a b i t t h e s h a l l o w e r water f a r t h e r o n t o t h e p l a t f o r m .
Although
now p a r t l y o b s c u r e d by t h r u s t f a u l t s , t h e t r a n s i t i o n z o n e may h a v e b e e n f i v e m i l e s o r so w i d e w h e r e o n e c a n g o from o n e t y p e of fauna i n t o a n o t h e r .
So I d o n o t t h i n k t h a t t h e p r e -
s e n t p r o v i n c i a l d i f f e r e n t i a t i o n of conodonts i n t h a t a r e a can s e r v e f o r i d e n t i f i c a t i o n of p l a t f o r m b o u n d a r i e s .
111 LATE P A L E O Z O I C AND T R I A S S I C CONODONT BIOSTRATIGRAPHY: CORRELATIONS AROUND THE E X P A N D I N G ATLANTIC OCEAN
David L . C l a r k D e p a r t m e n t o f Geology a n d G e o p h y s i c s U n i v e r s i t y o f W i s c o n s i n , Madison ABSTRACT The b e s t L a t e Devonian c o n f i g u r a t i o n f o r t h e p r e s e n t A t l a n t i c b o r d e r l a n d a r e a i n c l u d e s N o r t h America and n o r t h e r n Europe i n c o n t a c t o r c l o s e p r o x i m i t y and a " p r o t o - A t l a n t i c Ocean" s e p a r a t i n g t h e s e c o n t i n e n t s f r o m A f r i c a and S o u t h America.
D u r i n g t h e C a r b o n i f e r o u s , t h e A f r i c a - S o u t h America b l o c k
approached t h e North America-northern Europe b l o c k , d e s t r o y i n g t h e " p r o t o - A t l a n t i c " and d u r i n g t h e P e r m o - T r i a s s i c , t h e r e w a s no m a j o r s e p a r a t i o n o f a t e n u o u s l y f u s e d S o u t h A m e r i c a , A f r i c a , N o r t h America, E u r o p e c o n t i n e n t .
By L a t e T r i a s s i c , t h e p r e s e n t
N o r t h A t l a n t i c b e g a n foYming a s t h e E u r o p e a n a n d A f r i c a n c o n t i nents began t h e s h i f t t o t h e i r p r e s e n t p o s i t i o n s . The r e l a t i o n s h i p o f t h i s c o n t i n e n t a l movement t o c o n o d o n t b i o s t r a t i g r a p h y on e i t h e r s i d e of t h e p r e s e n t A t l a n t i c Basin c a n b e compared u s i n g t h e s i m i l a r i t y i n d e x , 2w/ a conodont f o r m - s p e c i e s .
+
b, for
Conodont s i m i l a r i t y b e t w e e n E u r o p e and
N o r t h America was g r e a t e s t d u r i n g t h e L a t e D e v o n i a n and P e r mian, w i t h a g r a d u a l decrease i n t h e T r i a s s i c t h a t m i g h t b e r e l a t e d t o formation of t h e p r e s e n t A t l a n t i c Basin. i n d i c e s among o t h e r A t l a n t i c b o r d e r l a n d
Similarity
c o n t i n e n t s a r e some-
what ambiguous, a s f a r a s d o c u m e n t i n g c o n t i n e n t a l p r o x i m i t y . T h i s i s b e c a u s e of l i m i t e d t a x o n o m i c d a t a ,
Also, such t h i n g s
a s low C a r b o n i f e r o u s A t l a n t i c b o r d e r l a n d c o n o d o n t s i m i l a r i t y may b e e x p l a i n e d by o t h e r f a c t o r s s u c h a s t h e e c o l o g i c i d i o s y n c r a s i e s of t h e a n c i e n t P a c i f i c and T e t h y a n w a t e r masses, or different continental elevations during t h e t i m e interval.
Re-
g a r d l e s s , some 100 c o n o d o n t z o n e s ( 4 3 D e v o n i a n , 23 C a r b o n i f e r ous, 1 2 Permian, 22 Triassic) p r o v i d e an important b i o s t r a t i g r a p h i c framework f o r t h e A t l a n t i c b o r d e r l a n d s . A c a l c u l a t e d z o n a l s i m i l a r i t y i n d e x m i m i c s t h a t of
form-species index.
the
The g r e a t e s t z o n a l s i m i l a r i t y i s d u r i n g
t h e L a t e D e v o n i a n , much l e s s d u r i n g t h e C a r b o n i f e r o u s , b u t h i g h again i n t h e Triassic b e f o r e dropping i n t h e L a t e Triassic, perhaps, i n p a r t , r e f l e c t i n g L a t e T r i a s s i c s e p a r a t i o n of North America and E u r o p e .
112
R~SUM~ La meillure configuration du d6vonien rkcent pour la r6giI on de bord actuelle de 1'Atlantique comprend l'Amerique du Nord et 1'Europe du nord, soit en contacte l'un avec l'autre soit en proche proximitk , et une "Oc6an proto-Atlantique" qui skpare ces continents de 1'Afrique et 1'Amkrique du Sud. Pendant le
carbonif&re, le bloc comprenant 1'Am;rique du Sud et 1'Afrique se rapprochait du bloc comprenant l'Am&ique du Nord et 1'Europe du nord. La "proto-Atlantique" ktait dktruite et pendant le permo-trias, il n'y avait aucune division dans le continent tentativement li6 de 1 'Amkrique du Sud , 1 'Afrique, 1 'Amhrique du Nord et 1'Europe. Par le trias rkcent, 1'Atlantique du Nord actuelle avait commenc; h prendre forme pendant que les continents europken et africain prenaient leurs positions actuelles. Le rapport de ce mouvement continental 'a la biostratigraphie conodonte des deux cotes du Bassin Atlantique actuel peut 2tre compar;! en utilisant l'indice de similitude, 2w/a+b , pour le genre-forme conodont. La similitude conodonte entre 1'Europe et 1'Amkrique du Nord ktait la plus grande pendant le dkvonien rkcent et le permien, avec une iente r6duction pendant le trias qui pourrait se rapporter la formation du Bassin AtI lantique actuel. Les indices de similitude parmi d'autres regions de bord de 1'Atlantique sont assez ambiguzs, quant 'a la documentation de la proximit; continentale. Ceci est dii aux I donnees taxonomiques limitkes. A U S S ~ ,des choses c o m e une basse similitude conodonte de la r.&gion de bord de 1'Atlantique carbonifLre peuvent hetre expliqu.&es par d'autres facteurs comme les idiosyncrasies des anciennes masses d'eaux de la Pacifique et Tethyan ou des differentes kl6vations continentales pendant ces p&iodes de temps. NQanmoins, il y a une centaine de zones conodontes (43 devoniennes, 23 carbonifbres, 12 permiennes, 22 trias) qui comprennent un cadre biostratigraphique important pour les r6gions de bord de 1'Atlantique. Une indice de similitude zonale calcul&e copie celle du genre-forme. La plus grande similitude zonale arrive pendant le dkvonien rkcent, encore moins pendant le carbonifkre, mais beaucoup plus encore pendant le trias avant de se rabaisser pendant le trias rAcent, peut-Gtre ainsi r&flLchissant en partie la s;!paration de l'Am&ique du Nord de 1'Europe pendant le trias rkcent.
113
INTRODUCTION
Worldwide c o n o d o n t b i o s t r a t i g r a p h y was summarized i n 1 9 7 1 (Sweet and B e r g s t r o m , 1 9 7 1 ) .
Since t h e n advances i n conodont
r e s e a r c h p r i m a r i l y have been i n a r e a s of b i o l o g y , paleoecology and p a l e o g e o g r a p h y .
Because s i g n i f i c a n t new i d e a s h a v e d e v e l -
oped c o n c e r n i n g c o n t i n e n t a l p l a c e m e n t d u r i n g t h e g e o l o g i c p a s t , i t i s now p o s s i b l e t o r e l a t e c o n o d o n t b i o s t r a t i g r a p h y more
c l o s e l y t o paleogeography.
N e w P h a n e r o z o i c w o r l d maps h a v e
provided a base f o r A t l a n t i c borderland b i o s t r a t i g r a p h y , i n particular
(Smith and o t h e r s , 1 9 7 3 ) .
T h i s p a p e r , w h i l e r e v i e w i n g and i n some i n s t a n c e s u p d a t i n g L a t e P a l e o z o i c and T r i a s s i c c o n o d o n t b i o s t r a t i g r a p h y ,
in-
cludes an a t t e m p t a t r e l a t i n g t h e A t l a n t i c Basin development of t h e t i m e t o c o n o d o n t s i m i l a r i t i e s i n t h e b o r d e r l a n d s . A t l a n t i c borderlands geology i n c l u d e s l i t t l e marine Pennsylvani a n , Permian o r T r i a s s i c .
The p o r t i o n o f t h i s r e v i e w r e l a t e d
t o c o n o d o n t s of t h e s e s y s t e m s i s b a s e d on w e s t e r n N o r t h A m e r i c a n and E u r o p e a n o c c u r r e n c e s t h a C , i n t h e s t r i c t e s t s e n s e , a r e not Atlantic borderlands.
The s i m i l a r i t i e s and d i f f e r e n c e s
among t h e s e c o n o d o n t f a u n a s a r e r e l a t e d d i r e c t l y t o t h e A t l a n t i c B a s i n d e v e l o p m e n t , however, and have a l e g i t i m a t e p l a c e i n any d i s c u s s i o n of A t l a n t i c B a s i n b o r d e r l a n d b i o s t r a t i g r a p h y . DEVONIAN
Introduction Devonian c o n o d o n t s a r e w e l l known i n t h e B r i t i s h I s l e s , Belgium, Germany, F r a n c e , S p a i n , P o r t u g a l and w e s t e r n N o r t h A f r i c a (Morocco and S p a n i s h S a h a r a ) f o r t h e e a s t e r n A t l a n t i c b o r d e r l a n d s , b u t l e s s w e l l known a l o n g t h e A t l a n t i c s e a b o a r d o f N o r t h America a n d r e l a t i v e l y unknown i n S o u t h America.
The
most c o m p r e h e n s i v e b i o s t r a t i g r a p h y i s b a s e d on work i n c e n t r a l Flirope (Germany, A u s t r i a , P o l a n d , B u l q a r i a , and I t a l y ) and w e y t e r n and c e n t r a l N o r t h America ( I o w a , I l l i n o i s , Nevada, Utah, O n t a r i o , A l b e r t a , and t h e A r c t i c I s l a n d s ) ,
The p r e c i s i o n
of Upper Devonian c o r r e l a t i o n a c r o s s t h e p r e s e n t A t l a n t i c B a s i n i s u n e x c e l l e d i n any o t h e r s y s t e m .
D e t a i l s o f L o w e r and M i d d l e
Devonian f a u n a s a r e n o t a s c o m p r e h e n s i v e .
114
Fig. 1. Devonian to Triassic "index" species. A l l figures except 14, adapted from Lindstrom, 1964. Figures, X28, except as indicated. 1
-
2
3
4 - 6
Epigondolella mungoensis (Diebel): 1, lateral: 2 , upper view; Upper Middle Triassic species; genus important in Upper Triassic, as well. Neospathodus cristagalli (Huckriede); Lower Triassic species: genus important in Upper Permian and throughout Triassic. Ellisonia sp.; multielement species important in Upper Permian and Triassic: 4, LC-element; 5, ozarkodinid element; 6 , U-element. (continued on next page)
115
Lower Devonian Gedinnian The m o s t i m p o r t a n t G e d i n n i a n c o n o d o n t i s I c r i o d u s w o s c h m i d t i ( F i g . 1 ) . I t marks t h e b a s e o f t h e Devonian o n b o t h s i d e s of t h e A t l a n t i c and r a n g e s t h r o u g h much o f t h e G e d i n n i a n ,
a t l e a s t a s r e c o g n i z e d i n N e w York Other s p e c i e s of I c r i o d u s
288).
-
v i s i n E u r o p e and
(2
I. latericrescens
Klapper
g . , 1971,
e-woschmidti,
p.
I.pesa-
i n North A m e r i c a ) p l u s
s p e c i e s of S p a t h o g n a t h o d u s c o m p r i s e t h e i m p o r t a n t Upper G e d i n n i a n c o n o d o n t s ( F i g . 2). F i g u r e 1 ic oi:ti nueri) 7 - 8 9 - 10
11 12
-
13
14
Giadiogondolella tethydis (Huckriede); 7 , upper; 8 , lower view; Middle T r i a s s i c s p e c i e s . F u r n i s h i u s t r i s e r r a t u s C l a r k ; 9 , l a t e r a l . ; 10, u p p e r view; Lower Triassic s p e c i e s . N e o g o n d o l e l l a s e r r a t a ( C l a r k and E t h i n g t o n ) ; Upper Permian s p e c i e s ; genus i m p o r t a n t t h r o u g h o u t Permian and Lower a n d M i d d l e T r i a s s i c . N e o g o n d o l e l l a s p . ; 1 2 , lcwer; 1 3 , u p p e r v i e w ; import a n t Perrr,ian and Lower and M i d d l e T r i a s s i c g e n u s . S w e e t o g n a t h u s w h i t e i ( R h o d e s ) ; X60; Lower P e r m i a n s D-~~~ ecies.
15 16
17 18
--
19
20
-
21
22
-
23
24 - 25 26
21 28 29 30 31 32 - 33
T :e 3 s t z e t oq r.a t k od u c s u 1c c p 1i c a tn s ( V c u r q 7 u i c E , :iav: 1ey and ? : i ; l e r ) ; : : i d d i e Perr:.ian s c e c i e s . A n c h i g n a t h o e s p . ; X23; P e r m i a n and Lower T r i a s s i c genus. I d i o g n a t h o d u s a c u t u s E l l i s o n ; X29; Upper C a r b o n i f e r o u s genus. S t r e p t o g n a t h o d u s e c c e n t r i c u s E l l i s o n ; X29; 1 8 , u p y e r ; 19, l o w e r v i e w ; C a r b o n i f e r o u s g e n u s . Gnathodus p u n c t a t u s ( C o o p e r ) ; X29; 20, l a t e r a l ; 21, upper view; C a r b o n i f e r o u s genus. C a v u s g n a t h u s c o n v e x a R e x r o a d ; 22, l a t e r a l ; 2 3 , u p p e r v i e w ; Lower C a r b o n i f e r o u s s p e c i e s . M e s t o g n a t h u s beckmsnni B i s c h o f f ; 24, u p p e r ; 2 5 , l o w e r v i e w ; Lower C a r b o n i f e r o u s s p e c i e s . S i p h o n o d e l l a q u a d r u p l i c a t a ( B r o n s o n and P l e h l ) ; Lower Carboniferous species. P a l m a t o l e p i s p e r l o b a t a s c h i n d e w o l f i M u l l e r ; Upper Devonian s p e c i e s . S p a t h o g n a t h o d u s s t e i n h o r n e n s i s Z i e g l e r ; Lower Devonian species. I c r i o d u s w o s c h m i d t i Z i e g l e r ; Lower Devonian s p e c i e s . I c r i o d u s l a t e r i c r e s c e n s b i l a t e r i c r e s e n s Z i e q l e r ; Lowe r Devonian s p e c i e s ; g e n u s i m p o r t a n t t h r o u g h o u t Devonian. A n c y r o d e l l a l o b a t a ( B r a n s o n and M e h l ) ; Upper D e v o n i a n species. P o l y g n a t h u s p e n n a t a H i n d e ; 32, u p p e r ; 33, l o w e r v i e w ; genus i m p o r t a n t t h r o u q h o u t Devonian.
116 Siegenian
Species of Icriodus are the important Siegenian indicators, including 2. pesavis and f. latericrescens in North America and I.huddlei in Europe, Spathognathodus sulcatus is important in North American faunas but has not been found in Europe (Ziegler, 1971). Emsian As in the other Lower Devonian units, species of Icriodus and Spathognathodus. are the most important elements of the Emsian fauna, In addition, Polygnathus appears in the Emsian in both Europe and North America and is particularly important in the latter. Polygnathus dehiscens, Spathognathodus steinhornensis s-. and Icriodus species including huddlei and I. bilatericrescens are most important for the Emsian. In general, Emsian correlations of the Atlantic borderlands area (e.g., New York, Spain and France) are better understood than those of the earlier Lower Devonian units. Middle Devonian Eif elian Conodonts of this interval are well defined in Belgium, Spain, and Germany and can be differentiated into three zones (Ziegler, 1971). Jcriodus coringer marks the base of the Eifelian on both sides of the Atlantic (New York and Belgium) but is succeeded upward by Icriodus latericrescens n. subsp. A (Klapper, et al., 1971) in North America, a form that has not been found in Europe. Polygnathus species dominate the Atlantic borderland faunas and correlations are fiiirly secure, Givetian Icriodus and Polygnathus dominate the biostratigraphically important Givetian faunas. Specific differences are common between the European and North American section but sufficient common species (e.g., Polygnathus varcus) as well as associated megafossils allow some confidence in correlaticn. .The sections in Germany, Belgium and Spain and the New York section have become standrads. Some taxonomic problems with potentially significant Icriodus species are yet to be resolved (Ziegler, 1971, p. 258).
s.
.;
117
I
I
t
North America
Western Europe
Polygnothus vorcus
Polygnothus spp.
lcriodus obliquimorginotus
lcriodus spp.
-9 .4
P,
A *%
Q)
Polygnothus kockelionus C
Spothognothodus bidentotus
2
P o l y g n o t h u s spp.
i2 lcriodus corniger
1
Non- lotericrescid
c
Icriodus-Polygnothus
8
I. b. blloterlcrescens-S.
..0u, lu
I
-
steinhornensis -Polygnothus
Polygnathus foveolotus Spothognothodus exiguus exiguus Spothognothodus steinhornensis Polygnothus dehiscens lenzi Spothognothodus exiguus exiguus Spothognothodus steinhornensis
I
Polypnothus dehiscens lenzl I
1 :.
$ ..
$1
1
Spothognothodus sulcotus (lote forms)
I. huddlei curvicoudo
Icr iodus later icrescens n. su bsp. B lcriodus cf. I. n.sp.A.
I.huddlei huddlei
crj
I -
I
lcriodus huddlei curvicoudoto rectonguloris s.1.-ongustoldes
lcriodus pesovis -Spothog nothodus johnsoni - S.tronsitons
Ancyrodelloideslcriodus pesovis
Spothognothodus n. sp. C
lcriodus w.postwoschmidti
Spothognothodus n. sp. 0
lcriodus w. woschrnidti
lcriodus w.woschmidt1
C
.? C
.s
8-
Fig. 2 .
Conodont z o n a t i o n of the Lower and Middle Devonian.
118
Upper Devonian By a l m o s t any s t a n d a r d o f m e a s u r e , t h e h i g h p o i n t of conod o n t e v o l u t i o n w a s reached d u r i n g t h e L a t e Devonian.
More t h a n
1 0 0 0 names h a v e b e e n p r o p o s e d f o r L a t e D e v o n i a n form t a x a ,
t w i c e a s many a s f o r any o t h e r s i m i l a r i n t e r v a l and e v o l u t i o n was more r a p i d t h a n a t any o t h e r t i m e ( C l a r k , 1 9 7 2 ) .
A t least
1 2 0 " i n d e x s p e c i e s " h a v e b e e n r e c o g n i z e d f o r t h e Upper Devonian
and t h e d e t a i l e d z o n a t i o n r e c o g n i z e d f i r s t i n Germany h a s b e come a n i d e a l model f o r b i o s t r a t i g r a p h y and a w o r l d s t a n d a r d f o r conodonts ( Z i e g l e r , 1971) ( F i g . 3 ) . S u b d i v i s i o n o f t h e Upper Devonian i n t o F r a s n i a n and Famenn i a n s t a g e s o r i n t o ammonoid u n i t s f o r c l a s s i f i c a t i o n i s unn e c e s s a r y now b e c a u s e t h e c o n o d o n t z o n e s h a v e p r o v i d e d a d e t a i l e d standard t h a t can stand a l o n e .
S p e c i e s of P o l y g n a t h u s ,
A n c y r o d e l l a and A n c y r o g n a t h u s a r e t h e i m p o r t a n t e a r l y Upper Devonian s p e c i e s b u t t h e s e t a x a a r e r e p l a c e d by a l i t e r a l f l o o d of P a l m a t o l e p i s s p e c i e s t h a t form the- s t r u c t u r a l u n i t y of t h e Upper D e v o n i a n z o n a t i o n . b e d e t e r m i n e d i n much of
Ancestor-descendant. r e l a t i o n s h i p s can t h e zonation.
A few s p e c i e s o f S p a t h -
o g n a t h o d u s a n d P r o t o g n a t h o d u s a r e i m p o r t a n t f o r t h e l a t e Upper Devcnian f a u n a s . CARBONIFEROUS Introduction The m o s t c o m p l e t e Lower C a r b o n i f e r o u s ccjnodont f a u n a s a r e t h o s e r e c o g n i z e d i n Germany and t h e Upper M i s s i s s i p p i V a l l e y . The A t l a n t i c b o r d e r l a n d s Lower C a r b o n i f e r o u s s e q u e n c e s c o n t a i n f a u n a s t h a t , i n p a r t , c o r r e l a t e w i t h t h e German and M i s s i s s i p p i V a l l e y s e c t i o n s , b u t w h i c h h a v e d i f f e r e n c e s , as w e l l (Rhodes and A u s t i n , 1 9 7 1 ) .
These l a t t e r f a u n a s a r e from F r a n c e ,
B e l g i u m , S p a i n , I t a l y , N o r t h A f r i c a , B r i t a P n and I r e l a n d , f o r t h e e a s t e r n A t l a n t i c b o r d e r l a n d s and t h e c e n t r a l A p p a l a c h i a n s for the western borderlands, Upper C a r b o n i f e r o u s c o n o d o n t s a r e p o o r l y known. American f a m a s i n t h e c e n t r a l A D r a l a c h i a n s
North
( P e n n s y l v a n i a , Ohio
and West V i r g i n i a ) h a v e b e e n o b t a i n e d from a p p r o x i m a t e l y 5% o f t h e s e p r e d o m i n a n t l y non-marine
sequences (Lane e t a l . , 1 9 7 1 ) .
C e n t r a l and w e s t e r n N o r t h American s e q u e n c e s a r e more m a r i n e a n d , h e n c e , more c o m p l e t e . complete.
The E u r o p e a n s e c t i o n i s q u i t e i n -
Conodont f a u n a s from t h e e a r l y Upper C a r b o n i f e r o u s
119
I Protognathodus UPPER MIDDLE LOWER UPPER MIDDLE LOWER
Spathognathodus costatus Polygnathus st yr iacus Scaphignathus velifer
1
Palmatolepis marginifera Palmatolepis rhomboidea
I
P a Ima t o leD is cr e D id a
[
I Pal matolepis triangularis Palmatolepis gigas
I
A . triangularis
I
1
I UPPER Polyg nat hus asym metricus
I
I
LOWER UPPER LOWER UPPER LOWER UPPER MIDDLE LOWER UPPER MIDDLE LOWER UPPER MIDDLE LOWER
Schmidtognathus hermani Polygnathus cristatus
Figure 3 .
I I
MIDDLE LOWER LOWE RMOST UPPER LOWER
Conodont z o n a t i o n of t h e Upper D e v o n i a n .
I
I
120
of Belgium, France, Britain and Ireland, comprise the record. The relatively poor conodont record for the Upper Casboniferous is related to ecologic constraints that may have as their source the shifting continental plates that closed the "protoAtlantic" Ocean during this period of different continental elevations. Lower Carboniferous Tournaisian-Kinderhookian
These roughly correlatable Lower Carboniferous time-rock units are characterized by similar to identical species of Siphonodella in both Europe and Yorth America. The m l t i p l e Siphonodella zones in North America are not as well represented in Europe but are correlated with Spathognathodus and Polygnathus species that are present in both sections (Collinson g al. - , 1971) (Fig. 4 ) Visean-Valmeyeran These intervals are distinguished by a diverse fauna of Gnathodus, Bactrognathus, Taphrognathus, Apatognathus, and Cavusgnathus species in North America and many of the same species plus important Polygnathus, llestognathus and Scaliognathus in Europe. Correlation among the various European and North American sections is good. The British Avonian sequence, in particular, is easily correlatable witpi the Upper Mississippi Valley section (Rhodes and Austi.n, 1971; Collinson et al., 1971). Namurian-Chesterian Although the late Lower Carboniferous interval is incornplete in the British Avonian, tht German and Mississippi Valley sections appear complete. All are characterized by Gnathodus, the German and North American sections by Paragnathodus, Kladognathus and Streptognathodus. Upper Carboniferous Namurian-Morrowan The Belgium Namurian sequence contains the best eastern Atlantic fauna (Eiggins and Bouckaert, 1968). Gnathodus and Idiognathodus dominate the faunas in Belgium and species are at least similar to those in the Morrowan of North America. Because of the rather poor state of knowledge concerning Upper Pennsylvanian conodont biostratigraphy, few correlations exist
.
121
(Rhodes and Austin, 1971). Morrowan falmas are better known than others from the Upper Carboniferous, and Lane et al. (1971) have described Midcontinent species of Spathognathodus, Idiognathodus, and Gnathodus that define seven biostratigraphic units. The same species are recognized in western North America and the central Appalachians. ~
British Avonian
M i s s i s s i p p i Valley R e g i o n
C b
Streptognathodus unicornis
*\
L
s
Kladognathus-Cavusgnothus naviculus
9
Kladognathus primus G. bilineatus-Kladognathus mehli
Gnat hod us g i r t y i co I I i n son i
G.bilineatus-Cavusgnathus altus
Gnathodus mononodosus \
no conodonts Mestognath. beckmanni-Poly. bischoffi
G. antetexanus-Polygnathus lacinatus
Gnathodus texanus-Taphrognathus
Boctrognathus-~aphrognathus
-Gnathodus s e m i g l a b e r -
G. semiglaber-Pseudo.multistriatus
Pseudopolygnath. multistriatus
c
Spath.costatus costatus-
Siphonodella isosticha-S.cooperi
b
Spat h ,robustus- S .t rid ent at us
-2- Gnathodus delicatus -2 C
S.quadruplicata-S.crenulata
Siphonodella-Polygnathus inornatus
t’ Patrognathus vario bilisSpathognothodus plurnulus
Siphonodella duplicota Siphonodella sulcato
Figure 4 .
Conodont zonation of the Lower Carboniferous.
122
Post-Morrowan C a r b o n i f e r o u s T h i s Upper C a r b o n i f e r o u s i n t e r v a l i s o n e of t h e more poorl y d e f i n e d u n i t s b a s e d on c o n o d o n t s . Post-Morrowan f a u n a s a r e known i n Belgium ( W e s t p h a l i a n ) a1.d i n c e n t r a l and w e s t e r n N o r t h America.
F o r t h e most p a r t , G n a t h o d u s , A d e t o g n a t h u s ,
I d i o g n a t h o d u s and Anchignathodus d o m i n a t e t h e f a u n a and many of t h e s p e c i e s r a n g e t h r o u g h o u t t h e Upper C a r b o n i f e r o u s s e c t i o n (Fig. 5 ) . R a t e s of c o n o d o n t e v o l u t i o n d u r i n g most of t h e P a l e o z o i c a p p a r e n t l y were h i g h e r t h a n d u r i n g t h e L a t e C a r b o n i f e r o u s and t h e u n i f o r m c o n o d o n t t a x a of th3.s i n t e r v a l r e f l e c t t h i s .
An
a l t e r n a t i v e i d e a t o t h i s i s t h a t o u r c o n c e p t o f L a t e Carboni f e r o u s t i m e , p a r t i c u l a r l y i n t h e post-Morrowan
i n t e r v a l , nay
b e i n e r r o r and t h i s i n t e r v a l may r e p r e s e n t o n l y a p a r t of what 'is p r e s e n t l y t h o u g h t t o b e i n c l u d e d . Whatever t h e e x p l a n a t i o n , Upper C a r b o n i f e r o u s c o n o d o n t s a r p e a r t o b e l e s s d i v e r s e t h a n t h o s e of any o t h e r s i m i l a r P a l e o z o i c o r Trf-a s s i c i n t e r v a l . PERMIAN
Introduction Marine Permian s t r a t a i s r a r e i n t h e A t l a n t i c b o r d e r l a n d s . T h e r e i s a l i m i t e d s e c t i o n i n G r e e n l a n d and c o n s i d e r a b l y more i n t h e A r c t i c I s l a n d s and wester;, N o r t h A m e r i c a .
There i s
l i t t l e known of t h e s e c t i o n i n S o u t h America and A f r i c a b u t
f a i r l y good, i f i n c o m p l e t e , s e c t i o n s a r e found i n s o u t h e r n and n o r t h e r n Europe.
The Dutch-German Permian B a s i n s have Lower
P e r m i a n , a t l e a s t some of which i s m a r i n e , and Upper Permian ( Z e c h s t e i n ) i r ! some a b u n d a n c e .
The Z e c h s t e i n Sea may h a v e had
c o n s i d e r a b l e e x t e n t i n n o r t h e r n Europe (Thomas, 1 9 7 5 ) and t h e German s u r f a c e s e c t i o n s c o n t a i n Upper Permian f a u n a s . E a r l i e s t Permian c o n o d o n t s a r e known o n l y i n w e s t e r n North America
( C l a r k and Behnken, 1 9 7 1 ) b u t l a t e s t Permian f a u n a s a r e
known now i n K a s h m i r - P a k i s t a n
and p a r t s of t h e mid-East
as w e l l
a s w e s t e r n N o r t h America ( F i g . 6 ) . Lower Permian Sakmarian-Wolfcampian E a r l i e s t Permian c o n o d o n t s , c o r r e s p o n d i n g t o e a r l y Wolfcampian d e t e r m i n a t i o n s i n w e s t e r n N o r t h A m e r i c a , e s s e n t i a l l y a r e t h e same a s t h o s e of t h e L a t e C a r b o n i f e r o u s , i . e . , Gnathodus, I d i o g n a t l i o d u s , A d e t o g n a t h u s , e t c . ( C l a r k , 1 9 7 4 ) .
123
Anchignathodus typicalis
0
$
3-.
7 Neogondolella orientalis
Gnathodus,Adetognathus,
7 Neogondolella rosenkrantzi
Idiognothodus,Anchignothodus SPP
::I
Q Neogondolella serrata serrota Neostreptognathodus clinei ldiognothodus n sp A
8 9s
* I S S , 42 C
Q
Neostreptognathodus proyi Neostreptognathodus sulcoplicatus Neostreptognathodus pequopansis
ldiognathodus n o d u l i f e r u s ldiognathodus ellisonl
Fig, 5.
Conodont z o n a t i o n o f Fig. 6. t h e Upper C a r b o n i f e r o u s .
Conodont z o n a t i o n of t h e Permian.
A c r i s i s i n ccncdont e v o l u t i o n d u r i n g t h e E a r l y Permian pro-
duced a d r a m a t i c c h a n g e i n a l l y o u n g e r f a u n a s ( C l a r k , 1 9 7 2 ) . To d a t e , t h e p r e - c r i s i s
E a r l y Permihn f a u n a h a s o n l y been rep o r t e d from w e s t e r n N o r t h A m e r i c a . Younger E a r l y P e r m i a n c c n o d o n t s a r e known f r o m N o r t h and S o u t h A m e r i c a ( C l a r k . 1 9 7 4 ) and i n c l u d e S w e e t c g n a t h u s , N e o g o n d o l e l l a , A n c h i g n a t h o d u s , and Ellisonia. Artinskian-Leonardian Conodonts o f t h i s i n t e r v a l a r e d o m i n a t e d by s p e c i e s of Neostreptognathodus, a d e s c e n d a n t of t h e o l d e r Permian Sweetcgnathus.
The b e s t f a u n a s o f t h i s i n t e r v a l a r e from w e s t e r n
N o r t h America b u t some s o u t h e r n E u r o p e a n m a t e r i a l i s known. N e o g o n d o l e l l a s p e c i e s showing good r e l a t i o n s h i p s w i t h o l d e r Wclfcampian m a t e r i a l a r e u s e f u l f o s s i l s i n t h i s i n t e r v a l (Behnken, 1 9 7 5 ) . Upper P e r m i a n Guadalupian S p e c i e s of N e o g o n d o l e l l a , E l l i s o n i a and N e o s p a t h o d u s a r e t h e i m p o r t a n t c o n o d c n t s of t h L s i n t e r v a l .
Good s e q u e n c e s i n
w e s t e r n N o r t h America form t h e s t a n d a r d of r e f e r e n c e w h i c h h a s been c o r r e l a t e d w i t h n o r t h e r n E u r o p e a n and G r e e n l a n d f a u n a s . The e m e r g e n c e of N e o s p a t h o d u s and N e o g o n d o l e l l a a s e x c e l l e n t
124
i n d e x forms i n t h e G u a d a l u p i a n i s t h e b e g i n n i n g o f a t r e n d t h a t continues u n t i l the Triassic e x t i n c t i o n . Dzhulfian Youngest m a r i n e Permian s t r a t a a r e e x t r e m e l y r a r e and a r e unknown i n t h e A t l a n t i c b o r d e r l a n d s a r e a ( F u r n i s h , 1 9 7 3 ) . G r e e n l a n d c o n o d o n t s a p p e a r t o b e G u a d a l u p i a n a s do Z e c h s t e i n s p e c i m e n s from n o r t h e r n E u r o p e .
K a s h m i r , P a k i s t a n and T r a n s -
Caucasus areas have p r o d u c e d t h e o n l y p u b l i s h e d d a t a on l a t e s t Permian conodonts (Sweet, 1970, 1 9 7 3 ) .
L a t e s t Permian t a x a
probably are p r e s e n t i n undescribed s e c t i o n s i n t h e western U n i t e d S t a t e s , as w e l l . Latest Permian faunas c o n s i s t of Neogondolella s p e c i e s
i n t e r m e d i a t e b e t w e e n t h o s e o f t h e G u a d a l u p i a n and E a r l y T r i a s s i c , A n c h i g n a t h o d u s , and E l l i s o n i a s p e c i e s s i m i l a r t o b o t h Permian and T r i a s s i c t a x a .
I n f a c t , Sweet (1973) concluded
t h a t t h e l a t e s t Permian t a x a d i d n o t e x p e r i e n c e t h e PermoT r i a s s i c c r i s i s and a r e d i s t i n q u i s h e d o n l y b e c a u s e o f t h e i r s i m i l a r i t y t o E a r l y T r i a s s i c forms. TRIASSIC Lower T r i a s s i c E a r l i e s t T r i a s s i c c o n o d o n t s " r e n o t p r e s e n t l y known from t h e A t l a n t i c borderlands b u t should be p r e s e n t i n Greenland,
a t least.
The f a u n a s from Lower T r i a s s i c s t r a t a a r e w e l l
known i n w e s t e r n Pu'orth America and i n t h e S a l t Range and T'ransI n d u s Ranges o f West P a k i s t a n .
The d e t a i l e d z o n a t i o n o f t h e
Lower T r i a s s i c i n c l u d e s s p e c i e s of N e o g o n d o l e l l a and Neospatho-
g , evolutionary
descendants of Permian s p e c i e s , as w e l l a s
new forms s u c h a s Parachirognathus-Furnishius and P l a t y v i l l o sus (Sweet -
e t al., 1971).
The e a r l i e s t f a u n a i s t h e same a s
t h a t f o u n d i n l a t e s t P e r m i a n s t r a t a b u t by t h e m i d d l e p a r t o f t h e Lower T r i a s s i c t h e f a u n a s a r e u n i q u e l y T r i a s s i c .
Species
of E l l i s o n i a a r e m o s t common b u t a r e n o t a s s t r a t i g r a p h i c a l l y r e s t r i c t e d a s t h e N e o g o n d o l e l l a and Neospathodus s p e c i e s (Fig. 7 ) . Middle T r i a s s i c Plosher ( 1 9 7 3 ) r e c o g n i z e d t h r e e f a u n a l a s s o c i a t i o n s f o r t h e M i d d l e and Upper T r i a s s i c .
-
The f i r s t , t h e g l a d i o g o n d o l e l -
l i d group, has only been recognized i n t h e Alpine-Tethyas b e l t .
125
R hae t ia n
Conodonts present but not diagnostic Epigondolella bidentata
Norian
Epigondolella multidentata
I
Epigondolella abneptis
Paragondole Ila p o l yg n a t h ifor mis
Karnian Neospathodus new passensis Epigondolella mungoensis
Ladinian Neogondolella mombergensis Neogondolella constricta
An isia n Neogondolella r e g a l e I
I Spat h ia n
Griesbachian Figure 7.
I
Neospathodus timorensis Neogondolella jubata unnamed zone
Anchignathodus typicalis
Conodont zonation of t h e T r i a s s i c .
1
126
The second, a group dominated by Neogondolella, has been recognized on both sides of the Atlantic as has the third, the Epigondolella group. Mosher (1973) suggested that, in spite of Triassic continental separation, with the exception of group 1, the Middle and Upper Triassic conodorits were not seriously affected. Species of Keogondolella, (r;. constricta and N_. mombergensis) dominate the Middle Triassic fai;nas, The German, Spanish and northern Italy faunas are well known as well as those from western North America. In addition, species described from the Cameroons in West Africa apparently are widespread (e.g., Epigondolella nungoensis). Upper Triassic Upper Triassic faunas include genera whose stratigraphic value is well documented in earlier rocks, e.g., Neospathodus, Epigondolella, and Paragondolella. Distinctive species are widespread in Europe and North America, although the best faunas have been obtained from the Alpine region of Europe. These species represent the end of the conodcnt lineage and their extinction at the close of the Triassic marks the end of conodont b iostra tigraphy UPPER PALEOZOIC AND TRIASSIC C.)NODOKT EVOLUTION The biostratigraphlc application of conodonts throuqh this ~ 1 5 million 0 year interval is based largely on morrhologic modifications of specimens that are interpreted as ancestors
.
and descendants.
Thus, approximately a dozen genera and sever-
al times that number of species provide the framework of a very sound biostratigraphy (Fig. 8 ) . Species of Icriodus mark the base of the Devonian and their evolution throughout the Devonian provide nine or ten zones of great importance. Their range is overlapped by that of Polygnathus, a taxon that alpeared in the late Early Devonian and provided a dozen or so important Middle and Upper Devonian to Lower Carboniferous zones. The stratigraphic significance of these two genera is overshadowed by species of Palmatolepis that dominate the Upper Devonian and provided more than a dozen zones. The Carboniferous began wi.th new genera, including Siphonodella, a definite Polygnathus descendant,
127
Epigondolella eospothodus
I
I
I
I
whose species provide four or more Lower Carboniferous (and one Late Devonian) zones. Thus, three genera and their descendants (Icriodus,Polyqnathus-Siphonodella, and Palmatolepis) are the basis of the Devonian-earliest Carboniferous biostratigraphy (40-50 million years), This homogeneous biostratigraphy breaks down in the remainder of the Carboniferous and earliest Permian, and spe
tion of biostratigraphically in- such as Gnathodus, Cavusgnaportant genera from the thus, Adetognathus, IdiognaDevonian-Tr ia ssic thodus, and Streptognathodus are the basis of the biostratlgraphy. Polygnathus, one of the important Devonian genera, is also important for the Carboniferous, and may be ancestral to some of the other important Carboniferous forms. A marked change, recently described as a c0:iodont "crisic:", occurred in the Early Permian (Clark, 1972). All of the important Carboniferous species became extinct and all postEarly Permian conodonts can be traced to a handful of srecies that were obscure, to some extent, prior to the crisis. Thus, the Sweetognathus-Neostreptoqnathodus line provided five or so Lower Permian zones, and evolved from Anchignathodus, a less important Carboniferous genus that survived until Early Triassic. Also important was Neogondolella, a genus that provides approximately a dozen Permian and Triassic zones, and nay be related to the relatively unimportant Carboniferous genus Gondolella. Neospathodus was new in the Permian and provided species for eight or so Permian and Triassic zones. Neospathodus was ancestral to Epigondolella and Paragondolella plus several other important Triassic forms. This post-Early Permian interval apparently was a return to the honoqeneous biostrati-
.
128
graphy of the Devonian-earliest Carboniferous, and three genera, Sweetognathus, Neogondolella and Neospathodus were the ancestors of most of the later Permian and Triassic biostratigraphically important species. ATLANTIC BORDERLANDS ON A DRIFTING CRUST The present North Atlantic Ocean began forming ~ 1 8 0 - 2 0 0 m.y. ago as part of the Russian platform rotation (Ostenso and Wold, 1973). This produced the opening of the North-AtlanticLabrador Sea that has resulted ii, the present separation of North America from most of Europe. North America and most of central and northern Europe had been in fairly close positions since at least the Early Paleozoic when the Russian platform collided with the North American-Greenland plate, causing the Caledonian orogeny. The proximity of Nortkl America and Europe during the duration from Early Paleozcic to the Early Triassic has been portrayed in a series of maps by Smith g . (1973). Unfortunately, details necessary to unambiguously define precise paleogeographic details of North America and Europe during this interval are lacking, Relationship of the Southern Atlantic borderlands continents is better known and during this interval, South America and Africa formed a single mass, separation of which during the Early Mesozoic produced tkle present South Atlantic. Thus, the Late Devonian configuration of the present Atlantic borderland areas included a North AmericanEurope proximity and a "proto-Atlantic" separating North America-Europe from Africa and South America. Durinq the Carboniferous the African-South Amei-ican block approached the North American-Middle and Nocthern Europe block, destroying the "proto-Atlantic" and by the Permian and Triassic there was no major oceanic area separating a tenuously fused South AmericaAfrica-North America-Europe. The Triassic collision of the Russian platform with the Siberian platform formed the Eurasian continent and produced rotation that opened the North AtlanticLabrador Sea. If this can be accepted as a factual description of the Late Paleozoic-Triassic paleogeographic framework, Devonian to Early Triassic conodont faunas in North America and Europe might be expected to be more similar than faunas of the Late
129
Triassic because of the formation of the modern Atlantic Ocean that separated the areas during the Triassic. Similarly, Devonian and Carboniferous conodont faunas of North AmericaEurope and those of southern Europe-Africa and South America should be less similar when they were separated by a "protoAtlantic" during the Devonian an? Carboniferous than during the Permian and Early Triassic when the continents were together. Late Triassic faunas might be expected to reflect the separation related to formation of the modern Atlantic. This might be considered a working hypothesis in spite of the many difficulties of reconstructing continental positions and the relatively unknown ecologic requirements of conodonts. In order to test this hypothesis, at least in part, the similarity index for faunas at various times and geographic positions has been calculated. This has been done using the similarity coefficient 2w/a+b, where w = number of conodont taxa common to two cmtinents, and a + b = total number of species on both continents. This has been based on form taxa recorded in the Wisconsin I B M conodont catalogue for all intervals indicated, except the Permian. The Permian figures are based on natural or multielement species. Resulting figures should be comparable, at least. These comparisons have been made with the warning of Cook and Taylor (1975) in mind, These students have emphasized the dana-r of unequivocal acceptance of the idea that faunal similarity = geographic proximity. Other factors (e.g., thermocline barriers) must be considered. The results are tabulated in Table 1. Apparent immediately is the incompleteness of the data matrix for many of the areas that could be expected to test the hypothesis. That is, South America and Africa have a unique roll in Late Paleozoic and Triassic continental patterns but there is no conodont data for most of this time interval that can be used. Indeed, the lack of data may be taken to indicate adverse ecology during this interval, Nicoll (1975) has suggested that conodonts could not tolerate the cold water associated with Permian glaciation, at least for western Australia. A Permian paleolatitude of S O 0 6 5 O for this part of Australia may suggest temperature tolerances for conodonts of that time interval. The more complete European-North American data is not inconsistent with
130
c o n t i n e n t a l p a t t e r n s (Fig. 9 ) b u t t h e drop i n s i m i l a r i t y index of . 2 9 0 - .194 - ,191 as t h e A t l a n t i c formed d u r i n g t h e T r i a s s i c c a n n o t by i t s e l f b e c o n s i d e r e d of e x t r a o r d i n a r y s i g n i f i -
c a n c e . The European-North American d a t a r e f l e c t s C a r b o n i f e r o u s e c o l o g i c problems t h a t a r e n o t n e c e s s a r i l y r e l a t e d t o c o n t i n e n t a l p o s i t . i o n ( e . g . , V a l l e n t i n e and Moores, 1 9 7 2 ) .
Indices for
o t h e r a r e a s ( T a b l e 1) may r e f l e c t t h e need f o r more d a t a more t h a n t h e y d o t h e c o n f i r m a t i o n of s h i f t i n g p o s i t i o n of t h e L a t e P a l e o z o i c and T r i a s s i c A t l a n t i c b o r d e r l a n d s . I n a d d i t i o n t o t h e form-taxa
s i m i l a r i t y index, a similar-
i t y i n d e x b a s e d on name z o n a l s p e c i e s was c a l c u l a t e d ( T a b l e 2 ) . T h i s s i m i l a r i t y m a t r i x h a s many b u i l t i n b i a s e s , n o t t h e l e a s t i m p o r t a n t of which i s t h e f a c t t h a t t h e r e a s o n a t a x o n i s t h e f o r a z o n e , i s b a s e d on i t s w i d e s p r e a d o c c u r r e n c e ,
name-giver
Hence, any c a l c u l a t i o n b a s e d on z o n a l s p e c i e s c o u l d b e expect e d t o g i v e a h i g h e r d e g r e e of s i m i l a r i t y t h a n t h e c o m p a r i s o n of s i m i l a r i t y i n d i c e s b a s e d on t o t a l t a x a ( T a b l e 1 ) . T h i s i s c o n f i r m e d , b u t t h e much h i g h e r i n d e x numbers c a n b e a t t r i b u t e d t o t h e b i a s i n s e l e c t i o n of z o n a l s p e c i e s . This zonal s i m i l a r i t y index i s p l o t t e d i n Figure 1 0 f o r N o r t h A m e r i c a and E u r o p e .
Whether o r n o t t h e L a t e T r i a s s i c
dropoff i s due t o t h e s e p a r a t i o n of t h e c o n t i n e n t s d u r i n g t h a t interva.'; i s u n c e r t a i n .
Certainly, it i s not inconsistent with
t h e p l a t e - t e c t o n i c moc;el now p r o p o s e d .
C a r b o n i f e r o u s low v a l -
u e s p r o b a b l y r e f l e c t t h e w e l l known C a r b o n i f e r o u s e c o l o g i c c h a n g e s more t h a n c o n t i n e n t a l p r o x i m i t y .
'
U
DEVONIAN
L
U MISSISSIPPIAN
L
U
PENNSYLVANIAN
L
U
PERMIAN
L
M
U
TRIASSIC
DEVONIAN
CARBON- PERMIAN TRIASSIC IFEROUS
Fig. 9. P l o t of d i v e r s i t y i n d e x F i g . 10. P l o t of z o n a l s i m i of f o r m - s p e c i e s f o r Europe and l a r i t y i n d e x of f o r m - s p e c i e s f o r Europe and N o r t h America. North A m e r i c a .
131 ACKNOWLEDGMENTS T h i s s t u d y was made p o s s i b l e b y NSF G r a n t GA-40454.
J i m Gamber, U n i v e r s i t y o f W i s c o n s i n g r a d u a t e s t u d e n t , a i d e d i n s i m i l a r i t y i n d e x c a l c u l a t i o n and u p d a t i n g o f t h e I B M f o r m t a x a Ed L a n d i n g , U n i v e r s i t y of M i c h i c a n a r a d u a t e s t u -
catalogue.
d e n t , m a i n t a i n e d t h e I B M c a t a l o g u e a n d made some p r e l i m i n a r y calculations for .this study,
P a u l Dombrowski d r a f t e d t h e f i g -
u r e s a n d t h e m a n u s c r i p t was t y p e d by C a t h e r i n e M . Ward. REFERENCES Austin, R.
1 9 7 3 , M o d i f i c a t i o r i of t h e B r i t i s h A v o n i a n
L.,
c o n o d o n t z o n a t i o n and a r e a p p r a i s a l o f European D i n a n t i a n c o n o d o n t z o n a t i o n and c o r r e l a t i o n .
Ann.
SOC. Gdol.
B e l g i q u e , v . 9 6 , p . 523-532. 1 9 7 5 , L e o n a r d i a n and G u a d a l u p i a n ( P e r m i a n )
Behnken, F . H . ,
conodont b i o s t r a t i g r a p h y ted States. Clark, D.
49, p.
284-315.
1972, E a r l y Permian c r i s i s and i t s b e a r i n g on
L.,
Permo-Triassic Volume 1, p .
,
i n w e s t e r n a n d s o u t h w e s t e r n Uni-
Jour. Paleont. v.
c o n o d o n t taxonomy.
Geol. P a l a e o n t . ,
Sp.
147-158.
1 9 7 4 , F a c t o r s of E a r l y P e r m i a n c o n o d o n t p a l e o e c o -
l o g y i n Nevada.
,
Jour. Palecnt. v.
and Behnken, F . H . ,
g r a p h y of
t h e Permian.
48, p.
710-720.
1 9 7 1 , Conodonts and b i o s t r a t i -
Geol. S O C . A m e r i c a M e m .
127,
p . 415-439. Collinson, Charles,Rexroad, C. B.
and Thompson, T . L . ,
1971,
Conodont z o n a t i o n of t h e N o r t h American M i s s i s s i p p i a n . G e o l . SOC. America Mem. Cook, H . E . ,
and T a y l o r , M .
127, p. E.,
353-394.
1975, E a r l y P a l e o z o i c continen-
t a l m a r g i n s e d i m e n t a t i o n , t r i l o b i t e b i o f a c i e s , and t h e thermocline, western United States.
Geol.,
v.
3 , p . 559-
562. F u r n i s h , W. M . ,
1973, Permian S t a g e N a m e s ,
i n , The P e r m i a n and
T r i a s s i c S y s t e m s and t h e i r m u t u a l b o u n d a r y , e d . Logan a n d C a n a d i a n S O C . P e t r o l . G e o l . Mem. 2 , p .
Hills.
522-548.
H i g g i n s , A . C . and B o u c k a e r t , J . , 1 9 6 8 , Conodcnt s t r a t i g r a p h y and p a l a e o n t o l o g y of t b l e Namurian o f B e l g i u m .
Expl.
C a r t e s G e o l . e t M i n e r e s B e l g i q u e , Men. 10, 64 p . Klapper
,
G.
,
O r r , W.
Sandberg, C . A . K.,
,
Collinson, C.
Rickard, L. V . ,
,
H u d d l e , J. W .
Schumacher, D . ,
,
Seddon, G .
and
132
Uyeno, T. T . , stratigraphy. Lane, H. R . , G.
D.,
1 9 7 1 , N o r t h American Devcnian conodoxit b i o -
Geol. SOC. A m e r i c a M e m .
127, p.
285-316.
M e r r i l l , G . K . , S t r a k a , J . J . , 111, and W e b s t e r , 1 9 7 1 , N o r t h American P e n n s y l v a n i a n c o n o d o n t s t r a t i -
graphy.
G e o l . SOC. A m e r i c a Men. 1 2 7 , p . 395-414.
N i c o l l , R . S . , 1 9 7 5 , The e f f e c t of L a t e C a r b o n i f e r o u s - E a r l y P e r m i a n g l a c i a t i o n on t h e d i s t r i b u t i o n o f P e r m i a n conodonts i n A u s t r a l a s i a . V.
G e o l . SOC. America A b s t r a c t s ,
7 , p . 828-829.
O s t e n s o , N . A . a n d Wold, R . J . , 1 9 7 3 , A e r o m a g n e t i c e v i d e n c e f o r o r i g i n of A r c t i c Ocean B a s i n , i n , A r c t i c Geology, e d . M.
G.
Pitcher.
A s s o c . P e t r o l . Geol. M e m .
Amer.
1 9 , p.
506-516. Rhodes, F . H . T. and A u s t i n , R . L . , 1 9 7 1 , C a r b o n i f e r o u s cono d o n t f a u n a s of E u r o p e . G e o l . SOC. A m e r i c a M e m . 1 2 7 , p . 317-352. Sandberg, C . A. and Z i e g l e r , W . ,
1973, Refinement of s t a n d a r d
Upper Devonian c o n o d o n t z o n a t i o n b a s e d on s e c t i o n s i n Nevada a n d West Germany. 122, Smith, A. G . ,
G e o l . P a l a e o n t . , v . 7 , p . 97-
B r i d e n , J . C . and Drewry, G . E . ,
1973, Phanero-
z o i c w o r l d maps, i n Organisms and c o n t i n e n t s t h r o u g h t i m e , e d . N . F . Hughes. Palaeont. Assoc. Spec. Paper 1 2 , p.
1-42.
Sweet, W. C . ,
1 9 7 0 , Uppermost P e r m i a n and Lower T r i a s s i c
c o n o d o n t s o f t h e S a l t Range and T r a n s - I n d u s R a n g e s , West Pakistan.
,
Univ. K a r s a s S p e c . Pub. 4 , p . 207-275.
1973, L a t e Permian and E a r l y Triassic ccnodont
f a u n a s , i n , The P e r m i a n and T r i a s s i c S y s t e m s and t h e i r m u t u a l b o u n d a r y , e d . Logan a n d H i l l s . P e t r o l . Geol. Mem. Sweet, W.
C.,
2 , p. 630-646.
and B e r g s t r o m , S . Pl., 1 9 7 1 ,
conodont b i o s t r a t i g r a p h y .
C a n a d i a n Soc.
e.Symposium o n
G e o l . SOC. America Men. 1 2 7 ,
449 p .
,
Mosher , L . C .
Hasenmueller, W. A . , Triassic. Thomas, T . M . ,
,
C l a r k , D. L.
,
Collinson, J . W.
,
and
1 9 7 1 , Conodont b i o s t r a t i g r a p h y of t h e
G e o l . SOC. A m e r i c a M e m .
1 2 7 , p . 441-465.
1975, Search f o r hydrocarbons i n s h e l f areas of
N o r t h w e s t E u r o p e : P r o g r e s s and p r o s p e c t s .
Amer.
ASSOC.
133
Petrol. Geol. Bull., v. 59, p . 573-617. Vallentine, J. W. and Moores, E. M., 1972, Global tectonics and the fossil record. Jour. Geol., v , 80, p . 167-184. Ziegler, W., 1971, Conodont stratigraphy of the European Devonian. Geol. SOC. America Mem. 127, p . 227-284.
Table 1.
S i m i l a r i t y i n d e x f o r conodont f o r m s p e c i e s d u r i n g i n d i c a t e d t i m e a n d f o r d e s i g n a t e d g e o g r a p h y . ~-
U.
Dev
U. L. M i s s
Eu & NA
.348
.136
Eu ti Af
.268
.017
Eu
&
.249
.312
Eu
& Asia
.lo9
.071
Aus
Miss L.
.231
L. P e n n U. P e r m M. T r i Penn L. P e r m L. T r i U. T r i -~ _ _ _ _ _ 0 .019 .34 .37 -290 .194 .191
x
X
X
x
X
X
X
X
0
0
.57
-53
.239
X
X
X
0 .065
-088 X
.239
x = i n s u f f i c i e n t data
T a b l e 2. Z o n a l s i m i l a r i t y i n d e x f o r N o r t h America and E u r o p e . __._
Upper T r i a s s i c
.75
Middle T r i a s s i c
.85
Lower T r i a s s i c
-76
Upper P e r m i a n
.67
Lower P e r m i a n
.29
Upper C a r b o n i f e r o u s
.25
Lower C a r b o n i f e r o u s
.57
Upper Devonian
-89
Middle Devonian
.50
L o w e r Devonian
-60
.-
.021 X
.270
Perm
Dev
Carb
Tri
.
.348
.204
.355
-268
.004
.249
-078
x
.lo9
.077
-55
x
.225 .03 6 X
.191
135
Discussion D r . F. M.
Swain:
To what e x t e n t may f a c i e s c o n t r o l have p l a y e d
an i m p o r t a n t r o l e i n l a t e P a l e o z o i c - T r i a s s i c c o n o d o n t d i s t r i bution i n c o n t r a s t t o continental separation o r proximity? D.
Dr.
L. Clark:
Much of o u r r e s e a r c h d u r i n g t h e p a s t 7 y e a r s
h a s been r e l a t e d t o c o n o d o n t p a l e o e c o l o g y . e v i d e n c e t h a t c o n o d o n t s behaved
T h e r e i s good
much l i k e most o t h e r g r o u p s
of o r g a n i s m s : some were v e r y c l o s e l y r e l a t e d t o f a c i e s f a c t o r s , b u t many were p a r t of t h e p e l a g i c r e a l m .
The s i m i l a r -
i t y index c a l c u l a t i o n s d i d not consider f a c i e s c o n t r o l .
The
s i m p l i f i e d a s s u m p t i o n was t h a t a g r e a t e r s i m i l a r i t y may i n d i c a t e p r o x i m i t y of e n v i r o n m e n t s .
I t was r e c o g n i z e d t h a t geo-
g r a p h i c a l l y a d j a c e n t b u t d i f f e r e n t e n v i r o n m e n t s may s u p p o r t completely d i f f e r e n t taxa.
The c a l c u l a t i o n s f o r t h i s r e p o r t
w e r e made s i m p l y t o see i f t h e s i m i l a r i t y i n d e x f o r c o n o d o n t s (most o f which were n e k t o n o r p l a n k t o n ) c o r r e l a t e d w i t h t h e b e s t guesses f o r c o n t i n e n t a l s e p a r a t i o n s ) . I would answer t h e Holdsworth q u e s t i o n more s i m p l y
--
d i f f e r e n t i a l s o l u b i l i t y of a p a t i t e d u r i n g d i a g e n e s i s of c a r b o n a t e s (and c l a s t i c s ) h a s n o t been r e c o g n i z e d .
Only re-
worked c o n o d o n t s show p r e s e r v a t i o n a l problems and t h i s c a n be e x p l a i n e d more e a s i l y by t h i n g s o t h e r t h a n s o l u t i o n of c a l cium p h o s p h a t e . D r . M.
C.
Keen:
Could you comment on t h e s t a t u s of t h e cono-
d o n t s r e p o r t e d from t h e C r e t a c e o u s of W . A f r i c a ? C l a r k : The " C r e t a c e o u s " c o n o d o n t s from A f r i c a a r e p a r t o f t h e upper Middle T r i a s s i c f a u n a r e c o g n i z e d worldwide. F u r t h e r , most o f t h e c o n o d o n t e l e m e n t s from t h e A f r i c a n sample became e x t i n c t ( i n normal s e q u e n c e s ) b e f o r e t h e c l o s e of t h e T r i a s sic.
The same i s t r u e f o r t h e J a p a n e s e " J u r a s s i c " m a t e r i a l .
T h e r e a r e no unambiguous p o s t - T r i a s s i c c o n o d o n t s . Dr.
B. K . Holdsworth:
I s t h e r e any p o s s i b i l i t y t h a t a p p a r e n t
f a c i e s r e s t r i c t i o n of c o n o d o n t s c o u l d b e c o n n e c t e d w i t h d i f f e r e n c e s i n s u s c e p t i b i l i t y t o s o l u t i o n of p a r t i c u l a r forms during diagenesis? Clark:
To t h i s I would s a y t h a t a s f a r a s I know, t h e r e a r e no
n o t a b l e d i f f e r e n c e s i n r e l a t i v e s o l u b i l i t y of conodont s p e c i mens i n t h e s e d i m e n t s w e g e n e r a l l y work w i t h .
Under normal
c i r c u m s t a n c e s , t h e c o n o d o n t s a r e v e r y w e l l p r e s e r v e d and show
136
no i n d i c a t i o n s of b e i n g p a r t i a l l y d i s s o l v e d .
Accordingly, L
do n o t t h i n k t h a t s e l e c t i v e p r e s e r v a t i o n f o r t h a t r e a s o n , i f a t a l l p r e s e n t , i s l i k e l y t o be a problem.
One b i g problem
h a s , however, a p p e a r e d and it i n v o l v e s t h e l a b o r a t o r y p r e p a r a t i o n . U p t o r e c e n t l y , some l a b o r a t o r i e s were u s i n g monoc h l o r i c a c i d f o r p r e p a r a t i o n o f conodont samples b u t u n f o r t u n a t e l y , it h a s t u r n e d o u t t h a t when t r e a t e d w i t h t h i s a c i d f o r some t i m e , some forms were d i s s o l v e d and o t h e r u n a f f e c t e d , t h e r e b y producing b i a s e d f a u n a s as l a b o r a t o r y a r t i f a c t s . F o r t u n a t e l y , a s f a r a s I know, t h e s e e a r l y - d i s s o l v e d e l e m e n t s have n o t p l a y e d a n i m p o r t a n t r o l e i n t h e d i s c u s s i o n s of conod o n t biogeography and b i o s t r a t i g r a p h y .
137
NOTES ON PALEOZOIC ACRITARCHS FROM THE ATLANTIC MARGINS F. M. Swain University of Minnesota, Minneapolis, Minnesota; and University of Delaware, Newark, Delaware Abstract Stratigraphically useful acritarch assemblages of Atlantic margins have been described from Ordovician rocks of Europe and North Africa; Silurian of Europe, North Africa, and North America; Devonian of Europe, North and South America; Carboniferous of North America, Europe, and North Africa; and Permian of Europe, North Africa, and North America. Zusammenfassung Formationskundig, ndtzliche Versammlungen der Acritarchen von Atlantischen Rdnder sind von Ordovizischen Felsen des Europas und Nordafrikas; dem Silur des Europas, Nordafrikas und Nordamerikas; Devon des Europas,,Nord- und Siidamerikas: Karbon des Nordamerikas, Europas, und Nordafrikas; und dem Perm des Europas, Nordafrikas, und Nordamerikas beschreibt.
Introduction The following brief discussion will deal with a few selected acritarch assemblages from localities bordering the Atlantic Ocean, together with references that will hopefully guide the reader to additional studies on these useful microfossils. A specialist was not available to prepare an acritarch article for this volume and the present writer has depended completely on publications from which a few notes are presented herein. Cambrian A few acritarch assemblages have been recorded from Cambrian rocks but the writer has not reviewed the literature on them. Ordovician Acritarchs of Ordovician rocks of Europe and North Africa have been studied by Burmann (1968), Cramer (1964), Cramer &. (1974a, 1974b) , Deunff (1961, 1968) , Martin (1968), Vavrdovd (1965, 1972) , and Eisenack, et al. (1973). Cramer et al. (197433) recorded 10 species which they be-
138
l i e v e have s h o r t s t r a t i g r a p h i c r a n g e s from t h e l a t e A r e n i g i a n s h a l e s o f t h e T a d l a B a s i n o f Morocco: Vavrdovd 1 9 7 2 ( T e x t - f i g .
Aureotesta c l a t h r a t a
1-8), Morrocanium s i m p l e x C r a m e r ,
K a n e s , D i e z , and C h r i s t o p h e r , 1 9 7 4 , Neoveryhachium c a r m i n a e (Cramer, 1964) and r e l a t e d forms, P r i s c o t h e c a s i e m p r e p l i c a t a Cramer e t a l . ,
1 9 7 4 , Rugulidium microrugulatum ( M a r t i n , 1 9 6 8 ) ,
g. v a r i r u g u l a t u m C r a m e r , e t a l . , 1 9 7 4 , g.
al.,
1974 ( T e x t - f i g .
1-9),
angulatum C r a m e r e t a l . dre, 1946).
,
5.
s c a b r a t u m (Crarner, 1 9 6 4 ) ,
5.
G-
1974 , and Veryhachium l a i r d i (Def l a n -
Most o f t h e s p e c i e s a r e l a t e A r e n i g i a n t o e a r l y
Llanvirnian i n range but
late Arenigian age.
g.
s i e m p r e p l i c a t a i s of Tremadocian t o
S e v e r a l of t h e forms a l s o o c c u r i n Belgium,
northern Florida subsurface, Spain, Libya, Tunisia, Algeria, and S a u d i A r a b i a . I n a c o m p a r a t i v e s t u d y Cramer e t a l .
(1974a), seven late
Aren g i a n t o e a r l y L l a n v i r n i a n a c r i t a r c h s a l s o a r e b e l i e v e d t o have r e s t r i c t e d r a n g e s : 1-1)
fig. 1-2),
M_.
P i r e a d u b i a VavrdovA 1 9 7 2 ( T e x t - f i g .
Multiplicisphaeridium hoffmanensis C r a m e r e t a l .
.:
m o r o q u e n s e Cramer e t a l . 1974 ( T e x t - f i g .
r a y i i C r a m e r e t a l . 1974 ( T e x t - f i g .
C r a m e r e t a l . 1974 ( T e x t - f i g . et a l . 1974 ( T e x t - f i g . 1 - 6 ) . -
1-5)
,
(Text-
1-3),
1 - 4 ) , Oodium mordidum
Coryphidium e l e g a n s C r a m e r
P_. d u b i a h a s a l s o b e e n r e c o r d e d
from Bohemia, L i b y a a n d i n p r e s u m a b l y r e w o r k e d S i l u r i a n d e p o s -
i t s o f NW S p a i n . Other than t h e occurrences i n t h e r e l a t i v e l y undisturbed Ordovician rocks of northern F l o r i d a subsurface r e f e r r e d t o a b o v e , l i t t l e i s known o f O r d o v i c i a n a c r i t a r c h s o f t h e w e s t e r n margin of t h e A t l a n t i c Ocean. Silurian S t u d i e s of S i l u r i a n " h y s t r i c h o s p h e r e s "
( a c r i t a r c h s ) of t h e
E u r o p e a n A t l a n t i c m a r g i n s i n c l u d e t h o s e by E i s e n a c k ( 1 9 3 4 , 1 9 5 4 , 1 9 5 5 , 1 9 5 9 , 1 9 6 2 , 1 9 6 5 , 1 9 7 0 , 1 9 7 1 , 1 9 7 2 1 , Downie ( 1 9 5 9 , 1 9 6 3 ) , D e f l a n d r e ( 1 9 4 2 , 1 9 4 5 ) , Deunff
(1954) , C r a m e r (1964,
1 9 6 6 a , 1 9 6 6 b ) , Bachmann a n d Schmid ( 1 9 6 4 ) , M a r t i n ( 1 9 6 5 , 1 9 6 6 1 , Stockmans a n d W i l l i h r e ( 1 9 6 3 ) , a n d L i s t e r ( 1 9 7 0 ) . The L l a n d o v e r i a n V i s b y Marl o f t h e B a l t i c
(Gotland) con-
t a i n s a r i c h a c r i t a r c h assemblage with Multiplicisphaeridium s p p . a n d o t h e r f o r m s , many of w h i c h a l s o r a n g e i n t o t h e Wenlockian elsewhere (Eisenack,
2.& . ) .
139
An assemblage of acritarchs from the type Wenlockian Wenlock Shales of Wenlock, England comprises, among other longer ranging forms, 38 species or subspecies that Downie (1963) believed are confined to the Wenlockian. Several of the forms he cited, however, are now known to occur in older or younger deposits. Correlation of the Wenlockian assemblage with Wenlockian beds in France (Montagne Noire, Deflandre, 1945) and in the Baltic region (Eisenack, 2 . G.) is shown by certain of the species. Three assemblages are recognized by Downie in the type Wenlockian: Assemblage I of the Buildwas Beds, lower Wenlock, typified by Deunffia and Domasia,.Assemblage I1 of the middle Wenlock Coalbrookdale Beds with "Veryhachium" bulbiferum and "V_." elongatum as typical species; and Assemblage I11 of the upper Coalbrookdale and Tickhill Beds, upper Wenlock, have abundant Micrhystridium, leiofusid acritarchs and Multiplicisphaeridium granulatispinosum. The attempt has been made here to justify Downie's generic assignments with those of the acritarch catalogue of Eisenack et al. (1973) but some of Downie's species have not yet been listed in the published volume of the catalogue. Silurian acritarch floras are also known from the San Pedro and equivalent formations of northern Spain (Cramer 1966, 1967, 1968, 1970; Cramer and Diez 1968), Tarannonian and late Wenlockian? of Belgium (Martin, 1965, 1968); Tunisia and Libya (Hoffmeister, 1959; Cramer, 1970) and Shropshire England, (Lister, 1970) Silurian acritarchs of the western Atlantic margins have been recorded by Fisher (1953), and especially by Cramer and Diez de Cramer in a series of papers. A comprehensive examination of Silurian acritarchs of eastern North America was made by Cramer and Diez de Cramer (1972). Localities pertinent to the present volume are those in Maine, New York, Ontario, Pennsylvania, Virginia, Georgia, and Florida. The authors recognized five acritarch biofacies which they believe to be time-transgressive in the Silurian. The five, probably temperature-controlled biofacies are: (1) Neoveryhachium carminae (Text-fig. 12) ; ( 2 ) Domasia (Text-fig. 18) ; (3) Deunffia (Text-fig. 15) (with D. eisenacki subf acies) ; (4) Gloecocapsamorpha prisca , (5) Pulvinosphaeridium-
.
140
Estastra biofacies
( t o l e r a n t of warm c l i m a t e ) ; when t h e d i s -
t r i b u t i o n of t h e b i o f a c i e s i s p l o t t e d on a Pangaea a r r a n g e m e n t of t h e S i l u r i a n c o n t i n e n t s .
The a u t h o r s r e c o g n i z e s e v e r a l a c -
r i t a r c h realms a l l i n t h e S i l u r i a n s o u t h e r n hemisphere.
The
proposed t i m e - t r a n s g r e s s i v e d i s t r i b u t i o n o f t h e b i o f a c i e s t h r o u g h t h e S i l u r i a n p e r m i t s t h e a u t h o r s t o s u g g e s t a r a t e of c o n t i n e n t a l d r i f t of 3 c m . p e r y e a r d u r i n g t h e Wenlockian. The Gondwana p o l e , assumed t o b e t h e S o u t h P o l e of t h a t t i m e , p r o b a b l y l a y i n n o r t h e r n South A f r i c a .
An i n t e r e s t i n g s u g g e s t i o n is made that t h e l o n g - p r o b l e m a t i c a l e a r l y P a l e o z o i c b l o c k of
n o r t h e r n F l o r i d a s u b s u r f a c e o r i g i n a l l y l a y between B r a z i l and A f r i c a a s a s m a l l s l i v e r of c o n t i n e n t a l c r u s t , and s u b s e q u e n t l y drifted t o its present position. Cramer and D i e z d e Cramer l i s t t h e t o t a l c h r o n o s t r a t i g r a p h i c r a n g e s of s p e c i e s o r s p e c i e s - g r o u p s t h a t o c c u r abund a n t l y and more o r l e s s world-wide i n t h e S i l u r i a n , a s f o l l o w s : Neoveryhachium c a r m i n a e
(Text-fig. 1 2 )
,
middle e a r l y
L l a n d o v e r i a n ( g r a p t o l i t e zone 1 8 o r 1 9 ) t o b a s a l e a r l y Gedinnian. Domasia t r i s p i n o s a , m i d d l e e a r l y L l a n d o v e r i a n t o t o p Wenl o c k i a n ( 9 . z. 3 1 ) . Domasia e l o n g a t a ( T e x t - f i g .
18)
,
e a r l y l a t e Llandoverian
( 9 . z. 2 2 o r 2 3 ) t o t o p Wenlockian. Deunffia
(2.
monospinosa,
g.
eoramusculosa,
g.
furcata,
( T e x t - f i g . 1 5 ) , m i d d l e l a t e L l a n d o v e r i a n ( 9 . z. 23) t o t o p Wenlockian. D e u n f f i a e i s e n a c k i , e a r l y L u d l o v i a n ( 4 . z. 33 t o p ) t o end Ludlovian ( p o s t
-
g . z. 36)
Quadraditium fantasticum, l a t e e a r l y Llandoverian t o top S i l u r i a n . (Text-fig. 13) Duvernaysphaera a r a n a i d e s
(Text-fig.
14)
,
l a t e early
L l a n d o v e r i a n t o e a r l y Emsian. Baltisphaeridium denticulatum
(Text-fig.
16)
( s . 1 . ) b a s e of
S i l u r i a n ( 9 . z. 1 6 ) t o t o p G e d i n n i a n .
Eupoikilofusastriatifera ( T e x t - f i g . 17) , % ( w e l l s c u l p t u r e d f o r m s ) , b a s e of S i l u r i a n t o b a s a l e a r l y Gedinnian. Devonian A c o l l e c t i o n of a c r i t a r c h s from g r e e n s h a l e s of t h e San Pedro F o r m a t i o n , e a r l y G e d i n n i a n ? a t O b l a n c a , C a n t a b r i c Moun-
141
t a i n s , n o r t h e r n S p a i n was r e c o r d e d by C r a m e r ( 1 9 4 6 3 3 ) .
Twenty-
f i v e s p e c i e s o c c u r i n f r e q u e n c i e s g r e a t e r t h a n 1%, and t h e f o l lowing a r e p r e s e n t i n numbers g r e a t e r t h a n 1 0 % of t h e f a u n a : Veryhachium t r i s u l c u m D e u n f f , V_.
reductum D e u n f f , V_. c a r m i n a e
Cramer, M i c r h y s t r i d i u m s t e l l a t u m D e f l a n d r e , and M_. f r a g i l e Deflandre.
T h e r e a r e f i v e new s p e c i e s of L e i o f u s a .
five species,
L.
One o f , t h e
b e r n e s g a Cramer i s s t a t e d t o be w i d e l y d i s t r i -
buted i n t h e San P e d r o F o r m a t i o n of t h e C a n t a b r i c Mountains; t h e o t h e r f o u r a r e a p p a r e n t l y more r e s t r i c t e d i n d i s t r i b u t i o n . Some a d d i t i o n a l r e f e r e n c e s t o s t u d i e s of Devonian a c r i t a r c h s i n S p a i n and o t h e r p a r t s of w e s t e r n Europe and N o r t h A f r i c a i n c l u d e t h o s e of Cramer ( 1 9 6 6 a , 1 9 6 7 ) , J a r d i n g e t a l .
(1972) ,
Lanzoni and M a g l o i r e ( 1 9 6 9 ) , k u s c h e r ( 1 9 6 9 ) , and Stockmans and
W i l l i k r e (1960, 1 9 6 2 a , b , 1 9 6 9 ) . A s m a l l c o l l e c t i o n of a c r i t a r c h s from t h e e a r l y Devonian
of Uruguay, Department o f Durazno ( M a r t i n e z - M a c c h i a v e l l o , 1 9 6 8 ) , c o m p r i s e s 11 s p e c i e s , i n c l u d i n g a new s p e c i e s of Leoni-
e l l a and
s e v e r a l new s u b s p e c i e s of B a l t i s p h a e r i d i u m s i m p l e x
Stockmans and Willi\ere, Veryhachium l e g r a n d i S t . and W .
1. exasperatum D e u n f f .
,
and
The c o l l e c t i o n was o b t a i n e d from s h a l e s
t h a t have a l s o y i e l d e d sporomorphs. B r i t o ( 1 9 6 7 ) d e s c r i b e d b o t h S i l u r i a n and Devonian a c r i t a r c h s from t h e Maranhao B a s i n of n o r t h e r n B r a z i l .
The a c r i -
t a r c h - b e a r i n g s e q u e n c e o c c u r s w i t h i n a b o u t 2,500 meters of P a l e o z o i c r o c k s r a n g i n g from Cambro-Ordovician t o Permian i n age and l i t t o r a l - t o b a t h y a l - t o c o n t i n e n t a l i n l i t h o f a c i e s . The S i l u r i a n i s r e p r e s e n t e d by t h e I t a i m F o r m a t i o n ( m i d d l e and upper p a r t s ) .
Two p a l y n o l o g i c a l zones termed S ( p o o r l y d e v e l o p e d )
and T l i e i n t h e I t a i m F o r m a t i o n ; Zone T c o n t a i n s D a c t y l o f u s c a m a r a n h e n s i s B r i t o and S a n t o s ( T e x t - f i g . m u l l e r i B r i t o and S a n t o s . t o m i d d l e Devonian.
19)
,
The o v e r l y i n g P i c o s S h a l e i s e a r l y
Zones Lower Q and R a r e w i t h i n t h i s forma-
t i o n ; R has V e l i f e r i t e s tenuimarginatus B r i t o E v i t t i a sommeri B r i t o
and L e i o f u s a
(Text-fig. 18)
,
Cramer; Lower Q h a s T a s m a n i t e s s p . a f f .
(Text-fig.
211,
and T r i a n g u l i n a a l a r g a d a
2.
mourai Sommer,
L e i o f u s a b a c i l l u m Deunff and L. b r a z i l e n s i s B r i t o and S a n t o s (Text-fig. 2 0 ) . The lower and m i d d l e p a r t s of t h e o v e r l y i n g Cabesas F o r m a t i o n , m i d d l e Devonian a r e r e p r e s e n t e d by Upper Zone Q and Zone P ; Upper Zone Q h a s P s e u d o l u n u l i d e a i m p e r a t r i -
142
zensis Brito and Santos (Text-fig. 231, and Zone P has Maranhites sp., Duvernaysphaera radieta Brito (Text-fig. 22) as well as Polyedrixum sp., Netromorphitae and Chitinozoa. The upper Cabeqas Formation, middle to late Devonian has Maranhites brazilensis Brito, M_. mosesi (Sommer) (Text-fig.. 24) , and Pterospermopsis brazilensis Brito (Text-fig. 25), but not Chitinozoa or Netromorphitae. Other western Atlantic Devonian acritarch floras were described from Canada by Deunff (1955, 1961). Devonian sporomorphs are well developed around the Atlantic margins but are not dealt with herein. Carboniferous A major assemblage of acritarchs and prasinophycean algae was recorded from late Devonian and early Mississippian rocks in a bore hole near Barberton, northeastern Ohio, by Wicander (1974). He described 15 new genera and 56 new species from the late Devonian Cleveland and Chagrin Shales and the early Mississippian Bedford Shale. Of these the following were only recorded from the Mississippian: Cymatiosphaera velicarina Wicander (Text-fig. 26) , Conradidium firmamentum W. (Text-fig. 27), Diexallophasis absona W. (Text-fig. 2 8 ) , D. cuspidus W., Exilisphaeridium simplex W. (Text-fig. 29) , Multiplicisphaeridium verrucarum W. (Text-fig. 30) , Navifusa drosera W. (Textfig. 3 1 ) 1 , and Stellinium cristatum W. (Text-fig. 3 2 ) . Carboniferous acritarchs were described from north Africa (Algerian Sahara) by Lanzoni and Magloire (1969) and from Belgium by Stockmans and Williere (1966). There is also extensive literature on Carboniferous sporomorphs of the Atlantic margins not considered herein. Permian Acritarchs of particularly small size and included in Baltisphaeridium ('Text-fig. 33) , Micrhystidium (Text-fig. 34) , Verhyachium (Text-fig. 35) , and Leiofusa (Text-fig. 36) , were found in Lower Permian Marls of Yorkshire England (Wall and Downie 1962). A striking feature of the assemblages is a variable plexus of forms ranging between those of Veryhachium type and those of Micrhystridium type. The former are polygonal tests with as few as four processes and represent a Veryhachium? irregulare complex. The latter are small spherical forms with
--
143
as many as 20 spines of Micrhystridium type. Other Permian acritarchs are known from the Permian of Yugoslavia and from the Sahara (Jekhowsky, 1961), but in western Atlantic, only from Oklahoma (Wilson, 1960). References Bachmann, A., and Schmid, M., 1964. Mikrofossilien aus dem osterreichischen Silur., Verh. Geol. Bundesanst., H. 1, p. 63, 64. Brito, I. M., 1967. Silurian and Devonian acritarchs from Maranhso Basin, Brazil, Micropaleontology, v. 13, p. 473-482. Burmann, G., 1968. Diachrodien aus dem unteren Ordovizium, Palaont Abh., v. 2, no. 4, p. 639-652. Cramer, F. H., 1964a. Microplankton from three Paleozoic Formations in the Province of Le6n, NW Spain, Leidse Geol. Meded., v. 30, p. 253-361, p l s . 1-24. , 1964b. Some acritarchs from the San Pedro Formation (Gedinnien) of the Cantabric Mountains in Spain, Bull. SOC. belge G & o l . , v. 73, p. 33-38. I 1966a. Palynomorphs from the Siluro-Devonian boundary in NW Spain, Notas y Communs. I. G . M. EspaKa, v. 85, p. 71-82. I 1966b. Hoegispheres and other microfossils incertae sedia of the San Pedro Tormation (Siluro-Devonian boundary) near Valporquero, Leon, Spain, Notas y Communas, I. G. M. Espa’ia, v. 86, p. 75-94. I 1967. Palynology of Silurian and Devonian rocks of Northwest Spain, Bol., I. G. M. Espasa, no. 77, p. 225286. I 1968. Silurian Falynologic microfossils and paleolatitudes, Neues Jahrb. Geol. Palaont., Jg. 1968/10, p . 591-597. I 1970. Distribution of selected Silurian acritarchs, Rev. EspaEola Micropaleont., Num. Extraord. VI, p. 1-203. Cramer, F. H., Allan, B., Kanes, W . H., and Diez, M. d. C. R., 1974a. Upper Arenigian to lower Llanvirnian acritarchs from the subsurface of the Tadla Basin in Morocco. Palaeontographia Abt. B., v. 145, Lfg. 5, 6, p. 182-190. Cramer, F. H., and Diez de Cramer, M. d. C. R., 1968, Considerationes taxonomicas sobre las acritarcas del Silurico Medio y Superior del Norte de Espaxa. Primera Parte: Las acritarcas acantomorfiticas, Bol. I. G. M. Espaza, nr 79/6, p. 541-574. , 1972. North American Silurian palynofacies and their spatial arrangement: acritarchs, Palaontogr. Abh. B., v. 138, Lfg. 5-6, p. 107180. Cramer, F. H., Kanes, W. H., Diez, M. d. C. R., and Christopher, R. A., 197433. Early Ordovician acritarchs from the Tadla Basin of Morroco. Palaeontoqraphica, Abt. B., v. 146, Lft. 3-6, p. 57-64. Deflandre, G., 1942. Sur les Hystrichosphkres des calcaires siluriens de la Montagne Noire, Acad. Sci. Paris, C. R., 215, p. 475-476.
.
144
, 1945. Microfossiles des calcaires siluriens de la Montagne Noire, Ann. Palgont., v. 31, p. 41-76. Deunff, J., 1954. Sur le microplancton du Gothlandien armoricain, SOC. Ggol. France, C. R . , Somm. v. 3,,p. 54, 55. , 1955. Un microplancton fossile Devonien a Hystrichosph'eres du continent Nord-Am&icain, Bull. Microscopie Appliqube, ser. 2, v. 5 , p. 138-149, pls. 1-4. , 1961a. Un microplancton 'a Hystrichosphbres dans le TrGmadoc du Sahara, Rev. Micropaleont., v. 4, p. 37-42. ., 1961Q. Quelques pre'cisions concernant les hystrichosphaeridees du Devonien du Canada, C. R. SOC. Geol. France, v. 8 , p. 216-218. , 1968. Arbusculidium,genre noveau d'Acritarche du Tremadocien marocain, C. R. Somm., Seances SOC. Geol. France, 1968, p. 101, 102. Downie, C., 1959. Hystrichospheres from the Silurian Wenlock Shale of England, Palaeontology, v. 2, p. 56-71. , 1963. "Hystrichospheres" (Acritarchs) and spores of the Wenlock Shales (Silurian) of Wenlock, England. Palaeontology, v. 6, pt: 4, p. 625-652, pls. 91, 92. Eisenack, A., 1934. Neue Mikrofossilien des baltischen Silurs I11 und neue Mikrofossilien des b8hmischen Silurs I, Palsont 2, V. 16, p. 52-76. , 1954. Hystrichospharen aus dem baltischen gotlandium, Senckenbergiana, v. 34,,p. 205-211. , 1959. Neotypen baltischen Silur-Hystrichosphxren und neue Arten, Palaeontographica ( A ) , v. 112, p. 193-211. , 1962. Einige Bemerkungen zu neuen Arbeiten Gber Hystrichosphsren, Neues Jahrb. Geol. Palaont., m.n., 1962, p. 52-101. . 1965. Mikrofossilien aus dem Silur Gotlands, Hystrichospharen, Problematika, Neues Jahrb. Geol. Palaont. Abh. 122, p. 257-274. , 1970. Mikrofossilien aus dem Silur Estlands und der Insel &el, Geol. Foren., Stockholm. Forh., v. 92, p. 302322. , 1971. Weitere Mikrofossilien aus dem Beyrichienkalk (Silur), Neues Jahrb. Geol. Palaont. mn., p. 449-460. , 1972. Chitinozoen und andere Mikrofossilien aus der Bohrung Leba, Pommern, Palaeontogr. Abt. A., v . 139, p. 64-87. Eisenack, A., Cramer, F. H. and Diez, M. d. C. R., 1973. Katalog der fossilen Dinoflagellaten, Hystrichosphsren und verwandten Mikrofossilien., v. 3, Acritarcha (I), E. Schweizbart'sche Verlagbuchlandlung, Nagele und Obermiller, Stuttgart, 1973, p. 1-1104. Fisher, D. W., 1953. A microflora of the Maplewood and Neagher Shales, Buffalo SOC. Nat. Sci., Bull. 21, p. 13-18. Hoffmeister, W. S., 1959. Silurian plant spores from Libya, Micropaleontology, v. 5, p. 331-334. Jardine', S., Combaz, A., Magloire, L., Peniguel, G., and Vachey, G., 1972. Acritarches du Silurien terminal et du D&vonien du Sahara AlgLrien, 2 . R. Sept. Congr. internat. Stratigr. Gebl. CarbonifLre, Krefeld 1971, v. 1, p. 295-310. Jekhowsky, B. de, 1961. Sur quelques hystricosphbres PermoTriasiques d'Europe et d'Afrique, Rev. Micropaleontologie, V. 3, p. 207-212, pls. 1, 2.
145 L a n z o n i , E . , and M a g l i o r e , L . , 1 9 6 9 . A s s o c i a t i o n s p a l y n o l o giques e t l e u r s a p p l i c a t i o n s s t r a f i g r a p h i q u e s dans le D ~ v o n i e n s u p g r i e u r e t C a r b o n i f & r e i n f e r i e u r d e Grand e r g Occid e n t a l ( S a h a r a A l g e r i e n ) , Rev. d e L ' I n s t i t u t F r a n q a i s du P e t r o l e , v . 2 4 , p . 441-468, p l s . 1 - 8 . L i s t e r , T . R . , 1 9 7 0 . A monograph of t h e A c r i t a r c h s and C h i t i nozoa from t h e Wenlock a n d Ludlow S e r i e s of t h e Ludlow and M i l l i c h o p e a r e a s , S h r o p s h i r e , P a l a e o n t o g r . S o c . Monographs, 1, p . 1 - 1 0 0 . M a r t i n , , F . , 1 9 6 5 . L e s a c r i t a r c h s du s o n d a g e d e l a b r a s s e r i e L u s t a K o r t r i j k ( C o u r t r a i ) ( S i l u r i e n b e l g e ) , B u l l . SOC. b e l g e G e o l . P a l . H y d r . , t . 7 4 , p . 351-400. , 1 9 6 8 . L e s A c r i t a r c h e s d e 1 ' 0 r d o v i c i e n e t du S i l u r i e n belges. Dgtermination e t v a l e u r s t r a t i g r a p h i q u e , I n s t . R o y . S c . N a t . ( 1 9 6 9 ) M e m . 1 6 0 , p . 1-175. M a r t i n e z - M a c h i a v e l l o , J . C . , 1968. Q u e l q u e s A c r i t a r c h e s d ' u n 4 c h a n t i l l o n du De'vonien i n f e ' r i e u r ( C o r d o b 6 s ) d p B l a n q u i l l o , D e p a r t e m e n t d e D u r a z n o , Uruguay, Rev. M i c r o p a l e o n t o l o g i e v . 11, n o . 2 , p . 77-84. Rauscher, R;, 1969. P r e s e n c e d ' u n e forme n o u v e l l e d ' A c r i t a r c h s d a n s l e Devonien d e Normandie, C . R . Acad. S c i . P a r i s , v . 268, s e r . D . , p . 34-36, p l . 1. S t o c k m a n s , F . , and W i l l i & r e , Y. , 1 9 6 0 . H y s t r i c C o s p h L r e s du Dgvonien b e l g e ( S o n d a g e d e 1 ' A s i l e d ' a l i e n e s a T o u r n a i ) , S e n c k . L e t h . , v . 4 , p . 1-11, p l s . 1, 2 . , 1 9 6 2 a . H y s t r i F h o s p h k r e s du DBvonien b e l g e (Sondage d e 1 ' A s i l e d ' a l i e n e s a T o u r n a i ) , B u l l . SOC. B e l g e G 6 0 1 . P a l e ' o n t . H y d r o l . , v . 7 1 , p . 41-77, D l S . 1, 2 . , 1 9 6 2 b . H y s t r i c h o s p h k r e s dy Devonien b e l g e (Sondage d e Wepion, B u l l . S O C . B e l g e Ggol. P a l e ' o n t . H y d r o l . , v . 7 1 , p . 83-89, p l s . 1, 2 . , 1 9 6 3 . H y s t r i c h o s p h g r e s ou mieux l e s A F r i t a r c h e s du S i l u r i e n b e l g e , Sondage d e l a b r a s s e r i e L u s t a C o u r t r a i ( K o r t r i j k ) , B u l l . S O C . B e l g . d e Ggol. , t . 7 1 , f a s c . 3 , p . 450-481, p l . 1-3 ( B r u s s e l s ) . , 1966. L e s a c r i t a r c h e s du D i n a n t i e n du S o n d a a e d e l ' a s i l e d ' a l i e n e s Tournai ( B e l c- r i -s u e ) ., B u l l . S O C . Belqe G e b l . v . 7 4 , p . 462-477, p l . .l. , 1 9 6 9 . . A c r i t a r c h e s du Fammenie n I n f g r i e u r , M e m . Acad. R . B e l g i q u e s , C 1 . S c i . , v . 3 8 , p . 1-63, p l s . 1-5. V a v r d o v i , m . , 1 9 6 5 . O r d o v i c i a n a c r i t a r c h e s from c e n t r a l Bohev . 4 0 , p . 351-357. mia, y e s t n i k U . U . G . , V a v r d o v a , M . , 1 9 7 2 . A c r i t a r c h s from K l a b a v a S h a l e s ( A r e n i g i a n ) , Vestnik U.U.G., v . 4 7 , p . 79-86. Wall, D . , and Downie, C . , 1 9 6 2 . P e r m i a n h y s t r i c h o s p h e r e s from B r i t a i n , P a l e o n t o l o g y , v . 5 , p t . 4 , p . 770-784, p l s . 1 1 2 - 1 1 4 . W i c a n d e r , E . R . , 1 9 7 4 . Upper Devonian-Lower M i s s i s s i p p i a n a c r i t a r c h s and p r a s i n o p h y c e a n a l g a e from O h i o , U.S.A., P a l a e o n t o g r a p h i c a , A b t . B . , v . 1 4 8 , L f g . 1 - 3 , p . 9-43, 1 9 p l s . W i l s o n , L . R . , 1 9 6 0 . A P e r m i a n h y s t r i c h o s p h a e r i d from Oklahoma, Okla. Geol. n o t e s , v . 2 0 , p . 7 , 1 7 0 .
146
Text-figures 1. Pirea dubia Vavrdova, X495, late Arenigian to early Llanvirnian shales, Tadla Basin, Morocco (after Cramer et al., 1974) 2. Multiplicisphaeridium hoffmanensis Cramer, et al., X825, late Arenigian to early Llanvirnian, Tadla Basin, Morocco (after Cramer et al., 1974) 3 . ~~u~tip:icisphaeridiummaroquense Cramer, et al., X450, late Arenigian to early Llanvirnian, Tadla Basin, Morocco, (after Cramer et al., 1974) 4 . Multiplicisphaeridium rayii Cramer et al., X495, late Areni gian to early Llanvirnian, Tadla Basin, Morocco (after Cramer et al., 1974) 5 . Oodium mordidum Cramer et al., X900, late Arenigian to early Llanvirnian, Tadla Basin, Morocco (after Cramer al. , 1974) 6. Zryphidium elegans Cramer et al., X825, late Arenigian to early Llanvirnian, Tadla Basin, Morocco (after Cramer al., 1974) 7. mcostella perforata Cramer et al., X495, late Arenigian to early Llanvirnian, Tadla Basin, Florocco (after Cramer 5 al. , 1974) 8. Ereotesta clathrata Vavrdova, X495, late Arenigian, Tadla Basin, Morocco (after Cramer et al., 1974) 9. Rugulidium rugulatum Cramer et al., X495, late Arenigian, Tadla Basin, Morocco (after Cramer, et al., 1974) 10. Xcanthodiacrodium s p . , X495, late Arenigian, Tadla Basin, Morocca (after Cramer & al., 1974) 11. Polygonium gracile Vavrdoz, X825, late Arenigian, Tadla Basin, Morocco (after Cramer & al., 1974) 12. Neoveryhachium neocarminae C r a m e s two different specimens, X495, middle early Llandoverian to basal early Gedinnian, eastern U.S.A. (after Cramer and Diez de Cramer, 1972) 13. Quadriditium fantasticum Cramer, X495, late early Llandoverian to top Silurian, eastern U.S.A. (after Cramer and Diez de Cramer, 1972) 14. Duvernayosphaera aranaides Cramer, X495, late early Llandoverian to early Emsian, eastern U.S.A. (after Cramer and Diez de Cramer, 1972) 15. Deunffia furcata Cramer, X495, middle late Llandoverian to top Wenlockian, eastern U.S.A. (after Cramer and Diez de Cramer. 1972) 16. Eupoikilofusa striatifera stericula Cramer, X495, species ranqes from base of Silurian to basal early Gedinnian, eastern U.S.A. (after Cramer and. Die2 de CTamer, 1972) 17. Dactylofusa maranhensis Brit0 and Santos, X240, Itaim Formation, Silurian, Maranhao Basin, Brazil (after Brito, 1967) 18. Evittia sommeri Brito, X240, Picos Shale, early to middle Devonian, Maranhao Basin, Brazil (after Brito, 1967) 19. Baltisphaeridium denticulata indianae Cramer, X495, base Silurian to top Gedinnian (range of species) eastern U.S.A., (after Cramer and Diez de Cramer, 1972) 20. Leiofusa brazilensis Brito and Santos, X300, Picos Shale, early to middle Devonian, Maranhao Basin, Brazil (after Brito, 1967) 21. Veliferites tenuimarginatus Brito, X750, Picos Shale, early to middle Devonian, Maranhao Basin, Brazil (after Brito, 1967)
147
22. Duvernayosphaera radiata Brito, X300, Cahecas Formation, middle Devonian, Maranhao Basin, Brazil (after Brito, 1967) 23. Pseudolunulidea imperatrizensis Brito and Santos, X240, Cabecas Formation, middle Devonian, Plaranhao Basin, Brazil (after Brito, 1967) 24. Maranhites mosesi (Sommer), X375, Cabecas Formation, middle Devonian, Plaranhao Basin, Brazil (after Brito, 1967) 25. Pterospermopsis brazilensis Brito, X240, Cabecas Formation, middle Devonian, Maranhao Basin, Brazil (after Brito, 1967) 26. Cymatiosphaera velicarina Wicander, X825, Bedford Shale, early Mississippian, Ohio (after Wicander, 1974) 27. Conradidium firmamentum Wicander, X525, Bedford Shale, early Mississippian, Ohio (after Wicander, 1974) 28. Diexallophasis absona Wicander, X750, Bedford Shale, early Mississippian, Ohio (after Vicander, 1974) 29. Exilisphaeridium simplex Wicander, X750, Bedford Shale, early Mississippian, Ohio (after Wicander, 1974) 30. Multi licis haeridium verrucanum Wicander, X712, Bedford Shale: earl: Mississippian, Ohio (after Wicander, 1974) 31. Navifusa drosera Wicander, X450, Bedford Shale, early Mississippian, Ohio (after Flicander, 1974) 32. Stellinium cristatum Flicander, X637, Bedford Shale, early Mississippian, Ohio (after Wicander, 1974) 33. Baltisphaeridium debilispinum Thlall and Downie, X750, Lower Permian Marl, Yorkshire , Enqland (after bJall and Downie, 1962) 34. Micrhystridium stellatum Deflandre, X750, Lower Permian Marl, Yorkshire, Fngland (after Val1 and Downie, 1962) 35. Veryhachium? irregulare Jekhowsky, X750, Lower Permian Marls, Yorkshire, England (after Wall and Downier 1962) 36. Leiofusa jurassica Cookson and Eisenack, X750, Lower Permian Marls, Yorkshire, England (after Wall and Downie, 1962) 37. Domasia elongata Downie, X1050, Buildwas Beds, early Wenlockian, Wenlock, England (after Eisenack et al., 1973)
148
This Page Intentionally Left Blank
151
PALEOBIOGEOGRAPHY OF CHITINOZOA BY Donald W. Z a l u s k y Glassboro State College G l a s s b o r o , New J e r s e y 08028 Abstract B i o g e o g r a p h i c d i s t r i b u t i o n of O r d o v i c i a n t o S i l u r i a n Chit i n o z o a of t h e a n c i e n t A t l a n t i c b o r d e r l a n d s i s d i s c u s s e d . The c l a s s i f i c a t i o n s y s t e m f o l l o w e d i s t h a t of J a n s o n i u s (1970). The g e n u s C o n o c h i t i n a i s r e s t r i c t e d t o forms w i t h a mucro o r c o p u l a ; some forms d e s i g n a t e d C o n o c h i t i n a by p r e v i o u s a u t h o r s are a s s i g n e d t o E u c o n o c h i t i n a , t h e r e f o r e , C o n o c h i t i n a , s e n s u s t r i c t o , h a s r a r e l y b e e n r e p o r t e d from N o r t h America. The gen u s C y a t h o c h i t i n a i s s u b d i v i d e d i n t o two i n f o r m a l g r o u p s : Qa t h o c h i t i n a I which i n c l u d e s t h e c o n i c a l f o r m s , a n d , C y a t h o c h i t i n a I1 which i n c l u d e s t h e c y l i n d r i c a l forms. Cosmop o l i t a n o r p r o v i n c i a l g e n e r a and s p e c i e s are n o t e d . Rgsurne' La d i s t r i b u t i o n b i o g g o g r a p h i q u e d e s C h i t i n o z o a i r e s Ordov i c i e n s aux C h i t i n o z o a i r e s S i l u r i e n s d e s a n c i e n s c o n f i n z A t l a n t i q u e s est d i s c u t g e . Le systeme d e c l a s s i f i c a t i o n s u i v i e s t c e l u i d e J a n s o n i u s ( 1 9 7 0 ) . L e g e n r e C o n o c h i t i n a est l i m it; a u x formes c o n t e n a n t un mucro o u un c o p u l a ; c e r t a i n s f o r mes a t t r i b u g e s a u p a r a v a n t % C o n o c h i t i n a p a r q u e l q u e s a u t e u r s , s o n t a t t r i b u d e s 'a E u c o n o c h i t i n a , d o n c , C o n o c h i t i n a , s e n s u s t r i c t o , o n t 6 t 6 r a r e m e n t r a p p o r t 6 d e l ' h d r i q u e du Nord. L e g e n r e C y a t h o c h i t i n a se s u b d i v i s e e n deux g r o u p e s i n f o r m e l s : C y a t h o c h i t i n a I q u i comprend l e s formes c o n i q u e s , e t Cyathoc h i t i n a I1 q u i comprend l e s formes c y l i n d r i q u e s . D e s g e n r e s e t d e s e s p e c e s c o s m o p o l i t e s ou p r o v i n c i a u x s o n t n o t & . Introduction C h i t i n o z o a , d i s c o v e r e d i n S i l u r i a n Baltic g l a c i a l errat-
i c s by E i s e n a c k i n 1929, are now documented from a l l c o n t i n e n t s e x c e p t A n t a r t i c a i n t h e O r d o v i c i a n t h r o u g h t h e Devonian interval.
I n i t i a l f i n d s were g e n e r a l l y b l a c k and opaque and
were i l l u s t r a t e d o n l y as s i l h o u e t t e s , of well-preserved
Subsequent d i s c o v e r i e s
m a t e r i a l p r o v i d e d some i n f o r m a t i o n on i n t e r -
n a l s t r u c t u r e s and a d d i t i o n a l i n f o r m a t i o n on s u r f a c e ornamen-
152
tation.
I t i s now r e c o g n i z e d t h a t s p i n e s a n d o t h e r ornamen-
t a t i o n o r i g i n a t e from a n e x t e r n a l tegument o r p e r i d e r m which i s f r e q u e n t l y d e g r a d e d and l o s t more r e a d i l y t h a n t h e p r i n c i p l e w a l l l a y e r o r ectoderm. The p r i m a r y m o r p h o l o g i c a l f e a t u r e s u t i l i z e d i n c h i t i n o zoan taxonomic a s s i g n m e n t a r e g e n e r a l o v e r a l l s h a p e , p r e s e n c e o r a b s e n c e of s p i n e s , and o t h e r o r n a m e n t a t i o n .
Loss of t h e
p e r i d e r m g e n e r a l l y r e s u l t s i n t h e l o s s of a l l smaller s p i n e s , a l t h o u g h some g e n e r a p o s s e s s l a r g e r p e r i a b o r a l s p i n e s which may b e r e t a i n e d .
Undoubtedly t h i s h a s c a u s e d some c o n f u s i o n
a s many d e g r a d e d s p i n o u s forms t h e n m i m i c smooth forms of similar o v e r a l l shape, Chitinozoan i n v e s t i g a t i o n s have demonstrated t h a t C h i t i nozoa were g e o g r a p h i c a l l y w i d e s p r e a d and e v o l v e d r a p i d l y i n t h e O r d o v i c i a n t h r o u g h Devonian P e r i o d s t h e r e b y p r o v i d i n g t h e p o t e n t i a l f o r a n extremely u s e f u l c o r r e l a t i o n t o o l .
Work i n
p r o g r e s s and r e c e n t i n v e s t i g a t i o n s w i l l u n d o u b t e d l y c l a r i f y much of t h e taxonomic and b i o s t r a t i g r a p h i c c o n f u s i o n .
Recom-
mended i s t h e two volume work o f t h e Commission I n t e r n a t i o n a l e d e M i c r o f l o r e du P a l e o z o i q u e ( c i t e d i n b i b l i o g r a p h y ) f o r a l l C h i t i n o z o a r e s e a r c h p r i o r t o 1965.
Subsequent t o t h i s d a t e ,
i m p o r t a n t p u b l i c a t i o n s ( E i s e n a c k , 1968, 1970, 1971, 1 9 7 2 a , 1972b, 1973; Boneham and Masters, 1973; Cramer, 1966, 1968, 1969, 1973; Cramer and Cramer, 1 9 7 2 a , 1972b; J e n k i n s , 1967,
153
1969, 1 9 7 0 a , 1970b; L a u f e l d , 1967, 1974; Lange, 1967, 1974; Urban, 1 9 7 2 ; Urban and K l i n e , 1970; Urban and Newport, 1973; and o t h e r s ) h a v e p r o v i d e d a s u b s t a n t i a l d e g r e e of m a t u r i t y t o C h i t i n o z o a knowledge.
E a r l y w o r k e r s r e p o r t i n g f i r s t occur-
r e n c e s from a p a r t i c u l a r g e o g r a p h i c e n t i t y c o n t r i b u t e d much t o t h e e x t e n s i o n of t h e p a l e o b i o g e o g r a p h y o f C h i t i n o z o a b u t und o u b t e d l y f o r c e d some i d e n t i f i c a t i o n s d u e t o t h e r e a s o n s d i s cussed earlier.
I t i s hoped t h a t i n v e s t i g a t o r s i n o t h e r spe-
c i a l i t i e s e n c o u n t e r i n g c h i t i n o z o a n s i n t h e i r samples w i l l req u e s t t h e a s s i s t a n c e of c h i t i n o z o a n workers t o a v o i d i n v a l i d taxonomic a s s i g n m e n t s . The s t r a t i g r a p h i c r a n g e of C h i t i n o z o a i s c o n f i n e d t o t h e P a l e o z o i c b u t i s o n l y known w i t h some d e g r e e o f c e r t a i n t y i n t h e O r d o v i c i a n t h r o u g h Devonian i n t e r v a l .
I s o l a t e d Cambrian
o c c u r r e n c e s h a v e b e e n r e p o r t e d b u t are (1) d o u b t f u l c h i t i n o z o a n s , ( 2 ) found i n r o c k s whose Cambrian a g e is q u e s t i o n a b l e , o r (3) p o s s i b l y r e p r e s e n t s t r a t i g r a p h i c l e a k s .
C h i t i n o z o a n s have
b e e n r e p o r t e d p r e s e n t i n P a l e o z o i c r o c k s younger t h a n Devonian b u t g e n e r a l l y o n l y t h e i r p r e s e n c e w a s noted--forms identified or illustrated. t r a t e d well-preserved
were n o t
Tasch ( 1 9 7 3 ) , however, h a s i l l u s -
C h i t i n o z o a r e p r e s e n t i n g two new g e n e r a
from t h e Permian F o r t R i l e y Fm. of Kansas (USA), b u t h a s n o t published f u r t h e r d e t a i l s .
154
Taxonomic C o n s i d e r a t i o n s The genus C o n o c h i t i n a w a s r e c o g n i z e d as s o b r o a d l y def i n e d t h a t many similar though d i s t i n c t f o r m s had b e e n assigned therein.
A s emended by Taugourdeau (1 9 6 6 ), C o n o c h i t i n a
was r e s t r i c t e d t o e x c l u d e forms w i t h o u t a mucro o r c o n s p i c u o u s callus.
For t h e e x c l u d e d f o r m s , Taugourdeau e r e c t e d t h e g e n u s
Euconochitina. I t i s g e n e r a l l y c o n s i d e r e d t h a t most i f n o t a l l C h i t i n o -
zoa formed c h a i n s a l t h o u g h c h a i n l e n g t h must h a v e v a r i e d between and w i t h i n g e n e r a .
D e s m o c h i t i n i d s and r e l a t e d forms a r e
f r e q u e n t l y found i n s h o r t t o l o n g c h a i n s w h e r e a s o t h e r forms
a r e g e n e r a l l y found i s o l a t e d o r n o t i n f r e q u e n t l y i n c h a i n s of o n l y two o r t h r e e t e s t s a t t a c h e d o r a l p o l e t o a b o r a l p o l e . F r e q u e n t l y more o r l e s s d i s t i n c t c o n c e n t r i c c i r c l e s , t h e call u s , a r e found c e n t e r e d o n t h e a b o r a l p o l e of many g e n e r a , p r o b a b l y t h e r e s u l t of c h a i n forming.
T h e r e f o r e t h e "conspic-
uous" c a l l u s becomes a matter of i n d i v i d u a l judgement i n ass i g n i n g a form t o C o n o c h i t i n a o r t o E u c o n o c h i t i n a .
This auth-
o r r e l i e s p r i m a r i l y o n t h e p r e s e n c e of a mucro t o d i f f e r e n t i a t e forms f o r a s s i g n m e n t .
On t h i s b a s i s , most N o r t h American
forms p r e v i o u s l y a s s i g n e d t o C o n o c h i t i n a are c o n s i d e r e d t o b e members of E u c o n o c h i t i n a . "...Conochitina
J a n s o n i u s (1970, p. 803) s t a t e d ,
s e n s u s t r i c t 0 i s common i n the B a l t i c area,
b u t s o f a r n o t r e c o r d e d from North America."
A review o f t h e
155
p u b l i s h e d i l l u s t r a t i o n o f North American " c o n o c h i t i n i d s " d i s c l o s e s o n l y a few forms w i t h a mucro.
I l l u s t r a t i o n s of
c. -
m i c r a n c a n t h a ( C o l l i n s o n and Schwalb, 1955, P1. 2 , F i g . 20) and
C.
d a c t y l u s (Pl. 2 , F i g s . 16-19) e a c h show more o r l e s s c l e a r -
l y a mucro; f u r t h e r , i n t h e d i a g n o s i s t h e y d e s c r i b e t h e p r e s e n c e of a s m a l l p a p i l l a (mucro) f o r e a c h s p e c i e s .
Boneham and
Masters (1973) i l l u s t r a t e d a specimen of C . d a c t y l u s which p o s s e s s e s a mucro ( F i g . 3 , n o , 8 ) .
C. m i n n e s o t e n s i s
i s shown
t o have a " b a s a l p r o c e s s " by J e n k i n s (1969, P1. 3 , F i g s . 11-
12).
J e n k i n s (p. 1 5 ) i n r e f e r e n c e t o
5.
r o b u s t a n o t e d , "A few
specimens i n e a c h p o p u l a t i o n b e a r a s h o r t c y l i n d r i c a l o r d i s t a l l y t a p e r e d r o d - l i k e b a s a l p r o c e s s ( c o p u l a , E i s e n a c k , 1959; 'mucro,'
L a u f e l d , 1967) . . . . I t
With t h e s e e x c e p t i o n s as n o t e d above, C o n o c h i t i n a s e n s u s t r i c t o i s n o t a b l y scarce i n N o r t h America w h e r e a s Euconochi-
tina is
r a t h e r common.
P o s s i b l y c o n t i n u e d work w i l l s h e d more
l i g h t o n t h i s anomaly. I t i s n o t t h e a u t h o r ' s i n t e n t t o s u b s t i t u t e i n f o r m a l no-
m e n c l a t u r e , however, a u s e f u l p u r p o s e would b e s e r v e d i f a t t e n t i o n i s c a l l e d t o t h e g e n e r i c d e s c r i p t i o n of C y a t h o c h i t i n a , E i s e n a c k , 1955.
E i s e n a c k d e s c r i b e d t h i s genus as h a v i n g a
more o r less c y l i n d r i c a l n e c k and a c o n i c a l chamber.
Such i s
t h e o v e r a l l s h a p e found i n t h e d e s i g n a t e d t y p e s p e c i e s , C. campanulaeformis.
The d i s t i n g u i s h i n g c h a r a c t e r i s t i c of Cya-
156
t h o c h i t i n a i s t h e p e r i a b o r a l c a r i n a i n a s s o c i a t i o n w i t h t h e above mentioned s h a p e .
E i s e n a c k and o t h e r s have s u b s e q u e n t l y
i n c l u d e d w i t h i n C y a t h o c h i t i n a forms w i t h l i t t l e o r no f l e x u r e t o d i s t i n g u i s h t h e neck, o r , t h a t is, e s s e n t i a l l y c y l i n d r i c a l forms.
Without d e s i r i n g t o change t h e a f f i l i a t i o n t o t h i s ge-
nus, t h e author proposes t h a t i n v e s t i g a t o r s d e s i g n a t e those c y a t h o c h i t i n i d s p o s s e s s i n g e s s e n t i a l l y a c o n i c a l chamber and c y l i n d r i c a l n e c k as C y a t h o c h i t i n a I and t h o s e e s s e n t i a l l y cyl i n d r i c a l i n o v e r a l l s h a p e as C y a t h o c h i t i n a 11; f o r example,
C. c y l i n d r i c a 11.
T h i s a u t h o r b e l i e v e s s u c h a p r o c e d u r e may
s e r v e a u s e f u l p u r p o s e i n b o t h p h y l o g e n e t i c and p a l e o b i o g e o g r a p h i c a l d i s t r i b u t i o n s t u d i e s p a r t i c u l a r l y i n t h o s e cases where i l l u s t r a t i o n s do n o t accompany c h i t i n o z o a n f a u n a l l i s t s . P e r h a p s t h e b e s t example of how t h i s may b e a p p l i e d i s found on P l a t e VII, Vol. 1, The C h i t i n o z o a n (Combaz, e t a l . ,
1 9 6 7 ) w h e r e i n a l l s p e c i e s of C y a t h o c h i t i n a known t h r o u g h 1965 are i l l u s t r a t e d .
These s p e c i e s are c a t a g o r i z e d a s Type I o r
I1 i n T a b l e 1 of t h i s r e p o r t .
O f t h e 23 s p e c i e s and subspe-
c i e s i l l u s t r a t e d , 11 are Type I and 1 2 are Type 11.
a r e p r o v i n c i a l t o t h e Northwest Africa--2 One of t h e f o r m e r ,
C.
Ten forms
Type I, 8 Type 11.
i n f u n d i b u l i f o r m i s , Taugourdeau and d e
Jekhowsky, 1960, w a s q u e s t i o n a b l y a s s i g n e d t o C y a t h o c h i t i n a by Taugourdeau and d e Jekhowsky b e c a u s e i t d i d n o t p o s s e s s a n observable carina.
157
European p r o v i n c i a l forms i n c l u d e 4 Type I and 4 Type 11. The l a t t e r i n c l u d e C. c o n i c a , Taugourdeau, 1 9 6 1 , from t h e Aquitane (S.W.
F r a n c e ) and
2.
e l e n i t a e , Cramer, 1964 from t h e
a d j a c e n t area o f Northwest S p a i n .
The a c r i t a r c h and c h i t i n o -
zoan f a u n a of Northwest S p a i n h a v e b e e n d e s c r i b e d by Cramer (1966) as more c l o s e l y a l l i e d w i t h t h e S a h a r a n Fauna t h a n t h e Baltic.
C.
s t e n t o r and
C. s t r i a t a ,
t h e o t h e r two s p e c i e s of
Type 11, a r e u n u s u a l l y l a r g e s p e c i e s from t h e B a l t i c area o r i g i n a l l y d e s c r i b e d by E i s e n a c k (1937) i n t h e genus Conochit i n a and s u b s e q u e n t l y t r a n s f e r r e d t o C y a t h o c h i t i n a as i n c l u d e d s p e c i e s by E i s e n a c k i n 1955. Cosmopolitan s p e c i e s o c c u r r i n g i n two o r more c o n t i n e n t s c o n s i s t of 5 Type I i n c l u d i n g t h e r a t h e r u b i q u i t o u s c a l i x -
campanulaeformis-kuckersiana complex. I t would t h u s a p p e a r t h a t C y a t h o c h i t i n a I1 i s r e s t r i c t e d t o t h e p r e s e n t s o u t h e r n l a t i t u d e s i f t h e a t y p i c a l forms s t e n t o r and C. s t r i a t a a r e e x c l u d e d .
C.
Cyathochitina I appear
t o have a more c o s m o p o l i t a n d i s t r i b u t i o n w i t h some s p e c i e s res t r i c t e d t o a t r a n s i t i o n a l zone between t h e S a h a r a n and t h e B a l t i c faunas.
I n p a r t i c u l a r , forms c l o s e l y r e l a t e d t o
2.
campanulaeformis are p r e s e n t i n B a l t i c , S a h a r a n , and North American s t r a t a . T h i s somewhat r e s t r i c t e d example u t i l i z i n g o n l y cyathoc h i t i n i d s known p r i o r t o 1965 a p p e a r s t o h o l d w i t h r e s p e c t t o
158
post-1965 r e p o r t s o f c y a t h o c h i t i n i d o c c u r r e n c e s .
Excluding a
p o s s i b l y p r e m a t u r e r e s t r i c t i o n of t h i s genus a n o t a t i o n o f t y p e would s e r v e t o e l i c i t u s e f u l p a l e o b i o g e o g r a p h i c a n d phylogenetic information. T a b l e 1. P a l e o b i o g e o g r a p h i c d i s t r i b u t i o n of pre-1965 c y a t h o c h i t i n i d s , Type I and 11. C y a t h o c h i t i n a Eisenack, 1955 P r o v i n c i a l Forms NW A f r i c a
Europe
Type I1 I1 I1 I1 I1 I1 I I II I1
Species Type S p e c i e s alata I1 c o n i c a cylindrica I1 e l e n i t a e djadoensis I granulata elongata I hymenophora fistulosa I novempopulanica fusiformis I regnelli hymenophora n i g a r i c a I1 s t e n t o r ?infundibuliformis I1 s t r i a t a koumeidaens i s obtusa Cosmopolitan Forms
Type I I I
Species c a l i x (NW A f r i c a , B a l t i c ) campanulaeformis (NW A f r i c a , B a l t i c , N. America) d i s p a r (NW A f r i c a , B a l t i c ) d i s p a r v e r r u c a t a ( A q u i t a n e , N. America) k u c k e r s i a n a (NW A f r i c a , B a l t i c , N. America)
I I
Ordovician and S i l u r i a n Biogeography The a n a l y s i s of t h e p a l e o b i o g e o g r a p h y of C h i t i n o z o a i s a t p r e s e n t c o m p l i c a t e d by s e v e r a l f a c t o r s .
A decided tendency
f o r c h i t i n o z o a n s t o r e a c t t o as y e t unknown e n v i r o n m e n t a l f a c tors exacerbates the picture.
The s u d d e n d i s a p p e a r a n c e a n d
159
r e a p p e a r a n c e a t h i g h e r i n t e r v a l s by some s p e c i e s h a s b e e n d i s c u s s e d by Mannil (1972).
C r a m e r (1970) n o t e d t h e sudden d r a -
matic i n c r e a s e s i n narrow z o n e s ,
t
2 mm, c h a r a c t e r i z e d by o n l y
one o r two s p e c i e s , a phenomenon which h e compared t o a l g a l blooms.
Rapid d e c r e a s e s i n abundance w i t h o u t a c o n c o m i t a n t
change i n l i t h o l o g y have b e e n d e s c r i b e d by L a u f e l d (1967). C e r t a i n areas o f t h e w o r l d have emerged a s q u a s i - r e f e r e n c e s e c t i o n s f o r c h i t i n o z o a n s due t o t h e i n t e n s i v e work of o n e o r more i n d i v i d u a l s .
Foremost among t h e s e i s t h e B a l t i c
area as t h e r e s u l t o f E i s e n a c k ' s a c t i v i t y s p a n n i n g more t h a n f o u r decades.
Material o b t a i n e d from areas g r e a t l y removed
from t h e B a l t i c h a v e o f n e c e s s i t y have b e e n compared t o t h e h o s t o f s p e c i e s d e s c r i b e d by E i s e n a c k . A s o u t h e r n r e f e r e n c e f a u n a came i n t o b e i n g due p r i m a r i l y
t o t h e e f f o r t s o f Taugourdeau, and Taugourdeau and d e Jekhowsky.
Material from c o r e samples s p a n n i n g O r d o v i c i a n t h r o u g h
Devonian s t r a t a p e r m i t t e d Taugourdeau and d e Jekhowsky (1960) t o i d e n t i f y highty-four
s p e c i e s ( f o r t y - n i n e new) i n two new
g e n e r a and t e n known g e n e r a .
A t t h e t i m e o f t h i s work t h e
p r e c i s e a g e of t h e s t r a t a w a s i n d e f i n i t e .
Cramer ( 1 9 6 4 ) , 1966) r e p o r t e d on t h e o c c u r r e n c e of Chit i n o z o a i n S i l u r i a n s t r a t a o f Northwest S p a i n a d d i n g a n o t h e r assemblage f o r comparative purposes.
L a u f e l d (1967) r e p o r t e d
on t h e C a r a d o c i a n C h i t i n o z o a of S h r o p e s h i r e , Wales.
Andress,
160
Cramer, and G o l d s t e i n (1969) r e p o r t e d on O r d o v i c i a n and S i l u r i a n c h i t i n o z o a n s o b t a i n e d from t h r e e w e l l c o r e s i n F l o r i da. O t h e r s i g n i f i c a n t works o f i n t e r e s t h e r e i n c l u d e Cramer (1968), c h i t i n o z o a n f a u n a o f the Red Mountain FIIL ( S i l u r i a n ) of Alabama and L a u f e l d ' s (1974) e x t e n s i v e r e p o r t on t h e S i l u r i a n C h i t i n o z o a of G o t l a n d .
Although Oklahoma i s geo-
g r a p h i c a l l y removed from t h e p r e s e n t A t l a n t i c Ocean, r e p o r t s by J e n k i n s (1967, 1969) on C h i t i n o z o a o f t h e O r d o v i c i a n formations.
V i o l a L s . F e r n v a l e L s . and t h e S y l v a n S h a l e of Okla-
homa p r e s e n t i m p o r t a n t c o n s i d e r a t i o n s f o r p a l e o b i o g e o g r a p h i c distribution.
J e n k i n s ' c o n t r i b u t i o n s a r e enhanced by h i s p r e -
vious e x t e n s i v e experience w i t h Baltic faunas. The f o r e g o i n g c o n s t i t u t e t h e p r i m a r y r e p o r t s which h a v e c o n t r i b u t e d most t o t h e p r e s e n t knowledge of t h e O r d o v i c i a n S i l u r i a n c h i t i n o z o a n b i o g e o g r a p h i c d i s t r i b u t i o n of the p e r i p h e r a l areas o f t h e A t l a n t i c Ocean.
S o u t h American c h i t i n o -
zoan f a u n a s a r e p r e s e n t l y l i t t l e known and do n o t c o n s t i t u t e a r e f e r e n c e i n t h e s e n s e used h e r e .
Many a d d i t i o n a l r e p o r t s
( o v e r 300) c o v e r i n g t h i s r e g i o n and o t h e r s supplement t h o s e p r e v i o u s l y mentioned b u t f o r v a r i o u s r e a s o n s a r e n o t germane to t h i s discussion The p a l e o b i o g e o g r a p h i c p a t t e r n of c y a t h o c h i t i n i d s p r e v i o u s l y d i s c u s s e d i n d i c a t e s a n o r t h e r n o r "Baltic" c h i t i n o z o a n
161
p r o v i n c e and a s o u t h e r n o r "Saharan" p r o v i n c e w i t h some deg r e e of b o t h p r o v i n c i a l i s m and c o s m o p o l i t a n i s m .
The i n t e r -
v e n i n g boundary i s b l u r r e d a n d t h e boundary i t s e l f may b e t h e r e s u l t o f a l i m i t i n g environmental f a c t o r r a t h e r than a physical barrier.
I t i s l i k e l y t h a t d i r e c t communication between
t h e two p r o v i n c e s e x i s t e d i n some t i m e i n t e r v a l s d u r i n g t h e l o n g t i m e s p a n under c o n s i d e r a t i o n . The c h i t i n o z o a n f a u n a o f t h e O r d o v i c i a n S y l v a n S h a l e w a s c o n s i d e r e d by J e n k i n s (1970) t o b e r e l a t e d t o t h e B a l t i c f a u n a on t h e b a s i s o f s p e c i e s i n common which r e p r e s e n t e d twentyf i v e p e r c e n t of t h e twelve s p e c i e s i d e n t i f i e d i n t h e S y l v a n S p e c i e s i n common were C o n o c h i t i n a cactacea, 6. ele-
Shale
D. s c a b i o s a . gans Y Desmochitina minor and -
(1969
Previously, Jenkins
had found t h e V i o l a and F e r n v a l e Limestones (Ordovi-
c i a n , Oklahoma) t o c o n t a i n c o s m o p o l i t a n e l e m e n t s c o n s i s t i n g of e l e v e n p r e v i o u s l y d e s c r i b e d s p e c i e s , and n o t e d a s t r i k i n g r e s e m b l a n c e t o f a u n a s from t h e Cardoc-Ashgill of E s t o n i a , Sweden, B r i t a i n , and t h e O s t e r k a l k of t h e B a l t i c r e g i o n .
The
cosmopolitan s p e c i e s c o n s i s t e d of Angochitina c a p i l l a t a , C. w e s e n b e r g e n s i s , Conochitina miracantha, -
robusta,
C. m i n n e s o t e n s i s ,
9c h i t i n a -minor
C. h i r s u t a , C.
C y a t h o c h i t i n a k u c k e r s i a n a , Desmo-
D. l a t a , R h a b d o c h i t i n a u s i t a t a , and R.
turgida.
L a u f e l d (1967) c o n c l u d e d t h a t t h e C a r a d o c i a n C h i t i n o z o a from D a l a r n a Sweden show a s t r i k i n g r e s e m b l a n c e t o E s t o n i a n
162
f a u n a s o f e q u i v a l e n t a g e (p. 292, " . . . a s might b e expected.") and s u g g e s t e d s i m i l a r i t i e s , l i m i t e d by s p a r s e i n f o r m a t i o n i n t h e l i t e r a t u r e , w i t h f a u n a s from t h e e a s t e r n United S t a t e s . Study o f t h e S h r o p s h i r e Caradocian c h i t i n o z o a n fauna by J e n k i n s (1967) l e d him t o conclude t h a t B a l t i c f a u n a s were c l o s e l y comparable whereas Saharan f a u n a s were o n l y r e m o t e l y comparable, c o n t a i n i n g o n l y two s p e c i e s r e c o g n i z e d i n Shrops h i r e (Rhabdochitina magna and S i p h o n o c h i t i n a p e l l u c i d a ) . However, as J e n k i n s p o i n t e d o u t (p. 485), t h r e e a d d i t i o n a l s p e c i e s known from t h e O r d o v i c i a n of B r i t i a n and t h e B a l t i c
are i n c l u d e d i n a Range C h a r t of Saharan forms by B e n o i t and Taugourdeau (1961) a l t h o u g h n o t f i g u r e d o r d e s c r i b e d by them. The p r e s e n c e of C l a t h r o c h i t i n a i n s e v e r a l Saharan assemblages and i t s a b s e n c e i n S h r o p s h i r e is b e l i e v e d by J e n k i n s (p. 485) t o emphasize t h e d i f f e r e n c e s between t h e s e two f a u n a s .
Some-
what anomalous i s t h e o c c u r r e n c e of c h i t i n o z o a n s b e l o n g i n g t o t h e genus S i p h o n o c h i t i n a i n B r i t i a n and t h e Sahara b u t a b s e n t i n the Baltic. I n a comparative s t u d y of G o t l a n d i c and Saharan S i l u r i a n chitinozoans.
Taugourdeau and d e Jekhowsky (1964) found a
h i g h d e g r e e of a f f i n i t y between t h e two b u t n o t e d a t i m e l a g i n c e r t a i n cases i n which some forms appeared much e a r l i e r i n t h e Sahara.
T h i s t i m e l a g w a s a t t r i b u t e d t o t h e l a r g e migra-
t i o n d i s t a n c e involved.
The B a l t i c c h i t i n o z o a n s were found by
163
Taugourdeau and d e Jekhowsky t o g e n e r a l l y be l a r g e r t h a n Saharan forms p a r t i c u l a r l y Conochitina.
Desmochitinids
were
remarkably s i m i l a r and a n c r y o c h i t i n i d s appeared t o f o l l o w p a r a l l e l e v o l u t i o n a l t h o u g h t h e y were much more d i v e r s i f i e d i n t h e Saharan r e g i o n .
F i n e l y p i l o s e a n g o c h i t i n i d s were noted
t o be a b s e n t i n A f r i c a .
Saharan assemblages w e r e c o n s i d e r e d
by Taugourdeau and de Jekhowsky t o b e more v a r i e d and evolved than t h e i r European c o u n t e r p a r t s .
Cramer (1973) c o n s i d e r e d F l o r i d a S i l u r i a n c h i t i n o z o a n s t o be q u i t e similar t o e q u i v a l e n t material from P o r t u g u e s e Guinea. Laufeld (1974) i n c l u d e d w i t h i n t h e genus G o t l a n d o c h i t i n a , Laufeld, 1974, t h r e e p r e v i o u s l y d e s c r i b e d C h i t i n o z o a (AncyroA . milanenc h i t i n a a e q u o r i s , A n g o c h i t i n a c a l l a w a y e n s i s , and -
sis)
p e c u l i a r t o t h e n o r t h e r n t i e r of s t a t e s of t h e C e n t r a l
United S t a t e s .
T h i s genus h a s n o t been r e c o g n i z e d from t h e
"Saharan" p r o v i n c e . From t h e f o r e g o i n g , i t i s e v i d e n t some d e g r e e of provincialism e x i s t s i n Ordovician-Silurian chitinozoan faunas i n s p i t e of some cosmopolitan e l e m e n t s which g i v e s r i s e t o n o r t h e r n and s o u t h e r n p r o v i n c e s as h a s been demonstrated f o r o t h e r organisms.
The n o r t h e r n o r "Baltic" p r o v i n c e encompasses
l a n d s b o r d e r i n g on t h e p r e s e n t B a l t i c Sea, t h e B r i t i s h I s l e s , A n t i c o s t i I s l a n d , Ontario, t h e northern states of t h e Central
164
United S t a t e s and i n t o Oklahoma.
The s o u t h e r n o r "Saharan"
p r o v i n c e i n c l u d e s .the Northwest A f r i c a n r e g i o n , P o r t u g u e s e Guinea, Northwest S p a i n , a n d F l o r i d a .
The i n t e r v e n i n g b o r d e r
between t h e two p r o v i n c e s may b e t r a n s i t i o n a l , however, " B a l t i c " and "Saharan" a s s e m b l a g e s a r e d i s t i n g u i s h a b l e b a s e d
on t h e c h a r a c t e r of t h e f a u n a as w e l l as c o m p o s i t i o n . References Cited Andress, N. E . , Cramer, F. H . , and G o l d s t e i n , R. F., 1969. O r d o v i c i a n c h i t i n o z o a n s from F l o r i d a w e l l s a m p l e s : T r a n s a c t . Gulf C o a s t Assoc. Geol. S O C . , v. X I X . B e n o i t , A , , and Taugourdea, P . , 1961. S u r q u e l q u e s c h i t i n o z o a i r e s d e l ' o r d o v i c i e n du S a h a r a : i n s t . F r a n s a i s P g t r o l e Rev., v. 1 6 , p. 1403-1421, 2 f i g s . , 2 p l s . C o l l i n s o n , C . , and Schwalb, H . , 1955. North American Paleozoi c C h i t i n o z o a : I l l . Geol. S u r v e y , Rpt. I n v e s t . 1 8 6 , 31 p . , 12 f i g s . , 2 p l s . Combaz, A . , e t a l . , 1967. M i c r o f o s s i l e s o r g a n i q u e du p a l g o z o i q u e ; p t . 2 , l e s c h i t i n o z o a i r e s , morphographie: C e n t r e N a t i o n a l Recherche S c i . Ed., 4 2 . , 8 f i g s . , 5 p l s . Cramer, F. H . , 1964. M i c r o p l a n k t o n from t h r e e P a l e o z o i c f o r m a t i o n s i n the P r o v i n c e o f Leon (NW-Spain): L e i d s e Geol. Mededel., v. 30, p . 253-261, 56 f i g s . , 24 p l s . , 1966. C h i t i n o z o a n s of a c o m p o s i t e s e c t i o n of upper L l a n d o v e r i a n t o b a s a l l o w e r G e d i n n i a n s e d i m e n t s i n n o r t h e r n Leon, S p a i n ; a p r e l i m i n a r y r e p o r t : S O C . Belg. G e o l o g i e B u l l . , v. 75, p . 69-146, 7 f i g s . , 5 p l s . , 1968a. C o n s i d e r a t i o n s p a l g o g e b g r a p h i q u e s 'a p r o p o s d ' u n e a s s o c i a t i o n d e m i c r o p l a n c t o n t e s d e l a se'rie g o t h l a n d i e n n e d e Birmingham ( A l a . , USA): SOC. Geol. F r a n c e B u l l . , ser. 7 , v. 1 0 , p. 126-131, 1 f i g . , 1969. P o s s i b l e i m p l i c a t i o n s f o r S i l u r i a n pal e o g e o g r a p h y from p h y t o p l a n c t o n a s s e m b l a g e s o f t h e Rose H i l l and T u s c a r o r a F o r m a t i o n s o f Pa.: J o u r . P a l e o n . , v. 43, p . 485-491, 2 f i g s . , p l . 70. , 1973. Middle and Upper S i l u r i a n c h i t i n o z o a n s u c c e s s i o n i n F l o r i d a s u b s u r f a c e : J o u r . P a l e o n . , v. 47, p . 279-288, 2 f i g s . , 2 p l s . Cramer, F. H . , and Diez d e Cramer, C. R . , 1972. S u b s u r f a c e s e c t i o n from P o r t u g u e s e Guinea d a t e d by palynomorphs as Midd l e S i l u r i a n : Am, Assoc. P e t r o l e u m G e o l o g i s t s B u l l . , v. 56,
165
p. 2271-2272, 1 f i g . E i s e n a c k , A , , 1937. Neue M i k r o f o s s i l i e n d e s b a l t i s c h e n S i l u r s , I V : P a l a e o n t . Z e i t s c h r . , v. 1 9 , p. 217-243, 22 f i g s . , p l s . 15, 16. , 1955. Neue C h i t i n o z o e n a u s dem S i l u r d e s Baltikums und dem Devon d e r E i f e l : S e n c k e n b e r g i a n a L e t h a e a , v. 36, p. 311-319, 3 f i g s . , 1 p l . , 1968. h e r C h i t i n o z o e n d e s b a l t i s c h e n G e b i e t e s : P a l a e o n t o g r a p h i c a , Abt. A, v. 131, p. 137-198, 1 3 f i g s . , p l s . 24-32. , 1970. M i k r o f o s s i l i e n a u s dem S i l u r E s t l a n d s und d e r I n s e l Oesel: Geol. Foren. Stockholm F o r h . , v. 9 2 , p. 302-322, 7 f i g s . , 1971. Weitere M i k r o f o s s i l i e n a u s dem Beyrichi e n k a l k ( S i l u r ) : Neues J a h r b . G e o l o g i e u. P a l a e o n t o l o g i e Monatsh., p. 449-460, 34 f i g s . , 1972a. C h i t i n o z o e n und a n d e r e M i k r o f o s s i l i e n a u s d e r Bohrung Leba, Pommern: P a l a e o n t o g r a p h i c a , Abt. A , v. 139, p. 64-87, 6 f i g s . , p l s . 16-20. , 1972b. B e i t r a g e z u r Chitinozoen-Forschung: P a l a e o n t o g r a p h i c a , Abt. A . v. 1 4 0 , p . 117-130, 1 f i g . , p l s . 32-37. , 1973. K l e i n o r g a n i s m e n a l s Z e r s t o e r e r s a e u r e f e s ter o r g a n i s c h e r S u b s t a n z e n und von B i o p h o s p h a t e n : P a l e o n t . Z e i t s c h . , v. 47, p. 8-16, 2 p l s . J a n s o n i u s , J a n , 1964. Morphology and c l a s s i f i c a t i o n o f some C h i t i n o z o a : Canadian P e t r o l e u m Geology B u l l . , v. 1 2 , p. 901-918, 2 f i g s . , 2 p l s . , 1967. S y s t e m a t i c s o f t h e C h i t i n o z o a : Rev. P a l e o b o t a n y and P a l y n o l o g y , v. 1, p. 345-360, 2 f i g s . , 1 p l . , 1970. C l a s s i f i c a t i o n and s t r a t i g r a p h i c a p p l i c a t i o n of C h i t i n o z o a : J o u r . P a l e o n t o l o g y , v. 43, p . 889. J e n k i n s , W. A. M . , 1967, O r d o v i c i a n C h i t i n o z o a from Shrops h i r e : P a l e o n t o l o g y , v. 1 0 , p. 436-488, 1 2 f i g s . , p l s . 6875. , 1969. C h i t i n o z o a from t h e O r d o v i c i a n : V i o l a and F e r n v a l e L i m e s t o n e s o f t h e A r b u c k l e Mountains Oklahoma: P a l a e o n t . Assoc. Spec. P a p e r s P a l a e o n t o l o g y 5, 44 p . , 1 0 f i g s . , 9 p l s . , 1970. C h i t i n o z o a from t h e O r d o v i c i a n S y l v a n S h a l e of t h e A r b u c k l e Mountains, Oklahoma: P a l e o n t o l o g y , v. 1 3 , p. 261-288, 7 f i g s . , p l s . 47-51. Lange, F. W . , 1967. S u b d i v i s a o b i o e s t r a t i g r a f i c a e r i v i s a o d a c o l u n a Siluro-Devoniana d a b a s i a do b a i x o Amazonas: s i m p o s i o s o b r e a B i o t a Amazonica A t l a s , v. 1, p. 215-326, 4 f i g s . , 12 p l s .
166
L a u f e l d , Sven, 1967. C a r a d o c i a n C h i t i n o z o a from D a l a r n a , Sweden: Geol. Foren. Stockholm F o r h . , v. 8 9 , p. 275-349, 34 f i g s . , 1974. S i l u r i a n C h i t i n o z o a from G o t l a n d , Foss i l s and S t r a t a , no. 5 , p . 1-128. Mannil, R., 1972. The z o n a l d i s t r i b u t i o n of c h i t i n o z o a n s i n t h e O r d o v i c i a n o f t h e East B a l t i c area. I n t . Geol. Congr. 2 4 t h Sess., S e c t . 7, 569-571. Tasch, P a u l , 1973. P a l e o b i o l o g y o f t h e I n v e r t e b r a t e s : John Wiley and Sons, I e c . , New York, 923 p. Taugourdeau, P h i l i p p e , 1961. C h i t i n o z o a i r e s du s i l u r i e n d ' A q u i t a i n e : Rev. M i c r o p a l g o n t o l o g i e , v. 4 , p . 135-154, 8 figs., 6 pls. , 1966. L e s c h i t i n o z o a i r e s : t e c h n i q u e s d ' g t u d e s , m o r p h o l o g i e e t c l a s s i f i c a t i o n : SOC. Geol. F r a n c e Mem., n. s e r . , v. 45, mem. 1 0 4 , 64 p . , 4 p l s . Tougourdeau, P h i l i p p e , and Jekhowsky, Benjamen, d e , 1960. Repartition et description des chitinozoaires silur-devonien d e l a C.F.P.A. e t d e d e q u e l q u e s s o n d a g e s d e l a C.R.E.P.S., l a S.N. Repal a u S a h a r a : I n s t . F r a n s a i s P g t r o l e Rev., v. 1 5 , p. 1199-1260, 1 9 f i g s . , 13 p l s . , 1964. C h i t i n o z o a i r e s S i l u r i e n s d e G o t l a n d ; comparison avec l e s Formes S a h a r i e n n e s , I n s t . F r a n f a i s P g t r o l e , v. 1 9 , p. 845870. et _ a1' 9 1967. M i c r o f o s s i l e s o r g a n i q u e s Taugourdeau, P h i l i p p e , d e palGozoique, pt,. 1; les c h i t i n o z o a i r e s ; a n a l y s e b i b l i o g r a p h i q u e i l l u s t r e e : C e n t r e N a t i o n a l Recherche S c i . Ed., 96 p . , 4 f i g . , 11 p l s . Umnova, N . , 1973. Methods of I n v e s t i g a t i o n , U s e of I n f a r e d L i g h t f o r t h e S t u d y o f C h i t i n o z o a : P a l e o n t . J o r . , 1973, no. 3, p. 394-400. Urban, J . B . , 1 9 7 2 , A r e e x a m i n a t i o n o f C h i t i n o z o a from t h e Cedar V a l l e y F o r m a t i o n o f Iowa w i t h o b s e r v a t i o n s o n t h e i r morphology and d i s t r i b u t i o n : B u l l s . Am. P a l e o n t o l o g y , v. 63, no. 275, p. 1-43, 9 f i g s . , 8 p l s . Urban, J. B . , and K l i n e , J . E . , 1970. C h i t i n o z o a of t h e Cedar C i t y F o r m a t i o n , Middle Devonian of M i s s o u r i : J o u r . P a l e o n t o l o g y , v. 44, p. 69-76, 2 f i g s . , p l . 18. Urban, J. B . , and Newport, R. L . , 1973. C h i t i n o z o a o f t h e W a p s i p i n i c o n (Middle Devonian) of Iowa: M i c r o p a l e o n t o l o g y , v. 1 9 , p . 239-246, 2 f i g s . , 2 p l s .
167
P A L E O Z O I C RAD IOLARIA : STRAT IGRP.PH IC DISTRIBUTION I PI ATLANTIC RORDF. P,LAI!DS B . K . H O L D S W O R T H , D e p a r t m e n t o f G e o l o g y , The U n i v e r s i t y , K e e l e ST5 5BG, E n g l a n d ABSTRACT: A few well-preserved, well-dated faunas suggest t h a t Ordovician, S i l u r i a n , Devonian and Carboniferous Systems rlay each possess c h a r a c t e r i s t i c Radiolaria populations, with a major change i n fauna occurring i n t h e Uoper S i l u r i a n or Lower Devbnian. Known Paleozoic and post-yaleozoic taxa appear t o be wholly unrelated. RESUHE: Ouelnues faunes bien pre'servges e t bien datges donnent 1 'impression que chacun des Systkrres de l ' n r d o v i c i e n , d u S i l u r i e n , d u De'vonien e t du Carbonif8re peut avoir des populations Radiolaires c , a r a c t f r i s t i q u e s , u? changment, radical dc faune ayant l i e u au, S i l u r i e n Superieur ou, ail Devonien I n f e r i e u r . Les taxa connus d u Paleozoioue e t Ju Postnaleozoioue paraissent compl'etement sans rapport, INTRODUCTION: For re1 i a b l e information on 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 of Radiolaria i n Paleozoic rocks we depend o n a few wellpreserved, well-dated faunas. Nevertheless, understanding of Paleozoic evolution i n the Subclass has undergone a n a j o r revolution in the l a s t 2 5 years. The new information cannot e a s i l y be summarized. Some important gerera a n d even f a m i l i e s remain unnamed a n d the majority of important assemblaqes a r e b u t p a r t l y described or wholly undescribcd. Vhat follows i s an extended a b s t r a c t of r e l a t i v e l y recent work, much o f i t unoublished. Space allows only inadequate i - l l u s t r a t i o n and p r i o r i t y i s given to important, h i t h e r t o unfigured f o r n s . SYSTCPATIC FRAFEWORK: The systematic framework adopted in t h i s account, outlined below, d i f f e r s in imoortant resnects from those of previous authors, b u t space precludes f u l l discussion. Subclass RADIOLARIA F?iller Order POLYCYSTINA Ehrenberg (emended Riedel) Suhorder SPUEIELL4RIA Ehrenberg I . "PALAEOACTII..IO?IIlIDS" new informal groupin?. All the Paleozoic Spumellaria with s i n g l e l a t t i c e d shell o r two o r more concentric s h e l l s , sometimes oylomate, lackinq t h e i n t e r n a l sniciilar system of Entactiniacea a n d the c h a r a c t e r i s t i c l a t t i c e struct,ure of Rotasphaerids (below). A l a r g e , diverse group, previously included i n the Actir?omriidae tiaeckel (emended Riedel 1967). T h o u g h Palacoactinomids show c l e a r morohol np i c sinii 1 ar i t i es t o pos t - P a 1 eczoic fic t i nomrii d a e , known Pa 1 eozoi c fauras seem t o show a marked decline i r Palaeoactinomids by Late Paleozoic, and i t i s n o t comnletely c l e a r t h a t they a r e , in f a c t , present in Late Paleozoic rocks. S t r a t i g r a n h i c d i s t r i b u t i o n s u g c e s t s , t h e r e f o r e , t h a t Palaeoactinommids a n d Actinomnidae could be Dhylo?enetically s e n a r a t e . 11. "ROTASPHAEnIDS" new i n f o r m 1 grouninq. Paleozoic Spumellaria with s i n g l e , s p h e r i c a l , l a t t i c e d s h e l l with annular meshes, lackinq the i n t e r n a l s p i c u l a r system of t h e Fntactiniacea and disolayino a point o r points upon t h e s h e l l fror: which r a d i a t e 5 or n'ore strong. s t r a i g h t l a t t i c e b a r $ . 1 . F ' n o t e : S u g g e s t i o n s o f h o l l o w s p i n e s a m o n g s t some O r d o v i c i a n Palaeoactinommids tend t o r e i n f o r c e t h i s view, b u t t h e "grouping" i s a l m o s t c e r t a i n l y p o l y p h y l e t i c and s t r a t i g r a p h i c r a n g e s and p h y l e t i c r e l a t i o n s h i p s a r e very unclear a t p r e s e n t .
168
Superfami l y ENTACTINIACEA P,iedel (Family E n t a c t i n i i d a e Riedel 1967) P a l e o z o i c q a d i o l a r i a , c o n v e n t i o n a l l y - thouclh Derhans i n c o r r e c t l y a l l o c a t e d t o S p u m e l l a r i a , w i t h a s p i c u l e of f o u r o r more r a y s , o o i n t - o r b a r - c e n t r e d , c o n s t i t u t i n g t h e main part of t h e s k e l e t o n o r i n c o r p o r a t e d w i t h i n a l a t t i c e d o r spongy s h e l l , e s s e n t i a l l y s n h e r i c a l , w i t h or w i t h o u t pylome. S h e l l very r a r e l y t u b u l a r , pylomate, and r a r e l y (and d o u b t f u l l y ) c o n i c a l , aylomate. 111, Family ENTPCTIIIIIDAE P i e d e l 1967 (enended h e r e i n ) . Entactiniacea with u n l a t t i c e d s k e l e t o n s , t h e s p i c u l e rays not d i f f e r e n t i a t e d t o d e f i n e a o i c a l and b a s a l hemisoheres. h b r e commonly, s k e l e t o n s l a t t i c e d , e s s e n t i a l l y s p h e r i c z l , l a c k i n g nylome. Tyne Genus: Entactinia Foreran. The E n t a c t i n i i d a e a s d e f i n e d h e r e i n c l u d e s t h e whole l a r a e groun of n r e v i o u s l y r e c o g n i z e d "non-Dylomate E n t a c t i n i i d a e " ( c f . Holdsworth 1 9 7 3 ) . C o n s t i t u e n t genera a r e i n t e r p r e t e d i n t h e s e n s e s o f Foreman (1963) a n d F o r t e y and Holdsworth ( 1 9 7 2 ) . IV. Family PYLENTDMEblIDAE D e f l a n d r e 1963 (emended h e r e i n ) . E n t a c t j n i a c e a with s i n g l e e s s e n t i a l l y s p h e r i c a l , l a t t i c e d ( e x t r e m e l y r a r e l y soongy) s h e l l , r a r e l y n o r e than one s h e l l , t h e o u t e r m o s t s h e l l d i s n l a y i n o an a c t u a l o r i n c i n i e n t nylome w i t h d i f f e r e n t i a t e d r i m . Lackinq t u b u l a r or c o n i c a l e x t e n s i o n a s s o c i a t e d w i t h pylome. Tyne Genus: Pylentonema D e f l a n d r e . ? Superfami l y E1:TPCTIPIIACEA \ I . Family PALAEOSCE>!IDIICAE P i e d e l 1967 (emended h e r e i n ) . P a l e o z o i c s p i c u l a r , s i l i c e o u s s k e l e t o n s comprising 6 t o 8 d i v e r g e n t r a y s , a r i s i n g most c o m o n l y from t h e ends o f a very s h o r t median b a r . 3 t o 4 or more e l a b o r a t e l y ornamented d e f i n e a " b a s a l 4 weaker, u s u a l l y unornamented r a y s d e f i n e a n " a p i c a l hemisphere". Type Genus: Palaeoscenidium D e f l a n d r e . VI. Family POPOFSKYCLLIDPE D e f l a n d r e 1964 L a t t i c e s h e l l p e r f o r a t e , t u b u l a r o r conicP1, w i t h open a n e r t u r e . S p i c u l e s c o n f i n e d t o a d a p e r t u r a l i n t e r i o r of s h e l l , b u t !with n o r t i o n s o f some r a y s i n c o r p o r a t e d i n s h e l l w a l l , Type Genus: Popofskyellum n e f l a n d r e . Of t h e t h r e e c o n s t i t u e n t g e n e r a , C y r t e q t a c t i n i a Foreman - nob c o n s t r u e d a s a n o n o s D e c i f i c Genus r e n r e s e p t e d by C y r t e n t a c t i n i a n r j n o t i c a Foreman - i s an u n q u e s t i o n a b l e e n t a c t i n i a c e i d . The i n c l u s i o n of Popofskyellum and T u s c a r i t e l l u m D e f l a n d r e w i t h i n t h e E n t a c t i n i a c e a i s l e s s c e r t a i n , and amongst s p e c i e s of t h e s e g e n e r a , one s p e c i e s of Fnnofskyellum o n l y i s d e f i n i t e l y known t o p o s s e s s i n t e r q a l s p i c u l e s . Subord,er ALBAILLELLARIA D e f l a n d r e 1953 (emended Holdsworth 1 9 6 o a ) . B i l a t e r a l l y synr;ett-icai P a l e o z o i c R a d i o l a r i a with c l o s e d o r ooen frames i r ? which can be d e t e c t e d a fundamental n a t t p r n o f t h r e e r o d s a r r a n g e d ii: t r i a n g u l a r form. V I I . Family CERATnIKTSCIME Pcldsworth 196oa A l b a i l l e l l a r i a i n which t h e s k e l e t a l frame i s c l o s e d a n d t y n i c a l l y c o n s i s t s o f 3 r o d s , a - r o d , b-rod a n d i n t e r s e c t o r , t h e a - r o d comrionly b e a r i n g p a i r e d s p i n e s ( c a v e a l r i b s ) developed i n a n l a w r.or?al t o t h a t o f t h e framc. I d e a l frame with b n t h ends o f a l l q r i w i n a l rric!s n r e x t r a t r i a n p u l a r e x t e n s i o n s , b u t some e x t e n s i o n s nay be s E r p r c s s Genus: C e r a t o i kisclrm D e f l a n d r e . VIII. Family ALEP~ILLELLIC~~E D e f l a n d r e 1952 ( m . h e r e i n ) . A l b a i l l e l l a r i a w i t h a - r o d and i r t e r s e c t o r l o n p e r than b - r o d , i r n a r t developed a s c o l u i - e l l a e w i t h i n l a r o s l y i m o e r f o r a t e s h e l l , n o t i o i n i n o
169
d i s t a l l y o r w i t h j u n c t i o n tenuous ( s e e Holdsworth 1 9 6 9 ) . Tyoe Genus: A1 bai 11 e l 1 a D e f l a n d r e . RADIOLARIA INCERTAE SEDIS I X . RADIOLARIA FAPIILIA NnVA F o r t e y and Holdsvortti 1972 P a l e o z o i c l a t t i c e d o r spongy c o n i c a l s h e l l s , ooen a t b a s e , with s n i r a l l y d i s p o s e d l a t e r a l s p i n e s and a l s o a p i c a l s p i n e s . Lacking i n t e r n a l s n i c u l e . X . Genus C O R Y T H O E C I A Foreman 1963 Paleozoic t u b u l a r , imperforate, s i l i c e o u s s h e l l s . S t r u c t u r e orobably variable and imperfectly know.
X I . SPICULAR FORllS A d i v e r s e , probably p o l y p h y l e t i c , g r o u p of forms with s k e l e t o n s c o n s i s t i n o
of t h r e a d - l i k e s p i c u l e r a y s , n o t o b v i o u s l y r e l a t e d t o any f a m i l y or aenus l i s t e d above. Note: I m o l i c i t i n t h e above t r e a t n e n t i s a r e j e c t i o n a t t b e moment o f arguments f o r D h y l e t i c c o n n e c t i o n s of t h e E n t a c t i n i a c e a with P l a s s e l l a r i a and e x t a n t s p i c u l a t e s o h e r o i d s , such c o n n e c t i o c s a o o e a r i n q d o u b t f u l i n t h e l i g h t o f p r e s e n t l y a v a i l a b l e mornhologic and s t r a t i g r a p h i c d a t a . O f p o s t - P a l e c z o i c f a m i l i e s c o r s i d e r e d by Riedel (1967) t o Clave Pa1eozo:c r e p r e s e n t a t i v e s , P h a c o d i s c i d a e , Spongodiscidae and L < t h e l i d a c a r e omi:ted, a s no c o m p l e t e l y convincing e v i d e n c e of t h e i r e x i s t e n c e has been found in well-preserved Paleozoic faunas. E Q I E F CESCRIPTIOIdS OF T A X A AI:C OUTLINE OF STQATIGWPHIC DIST~TPUTION C A W 3 ? I A Y : 'lo ~-
adeqliate f a u n a s a r e known from t b e A t l a n t i c Borderlands o r
e l s e w h e r e , b u t Palaeoactinommids a r e probably p r e s e n t and "azarov (1975) r e p o r t e d E n t a c t i n i i d a e E( this oaper). ORnnVICIAr::
E x c e l l e n t l a t e A r e n i q i a n R a d i o l a r i a have been d e s c r i b e d f r n r i
t h e ! ' a l h a l l f o n n a Formation, S p i t s b e r g e n ( F o r t e y and Holdsworth 1 9 7 2 ) . P second fauna i s known froiii t h e e a r l y A r e n i g i a n of t h i s Formation ( B r u t o n e t a l . , i n o r e o a r a t i o n ) and a s i r i l a r o c c u r r e n c e i s r e n o r t e d from t h e Table Head Formation, Rewfoundland ( P e r q s t r d m : 9 7 4 ) . P a l a e o a c t i n o m n i d s dominate both S p i t s b e r q e n h o r i z o n s ( " s n h a e r o i d s " of F o r t e y and Holdsworth 1 9 7 2 ) . A r c h i t e c t u r e i s o a r t i c u l a r l y c l e a r i n t h e e a r l i e r assemblage anc e n t a c t i n i a c e i d s o i c u l e s a r e u n a u e s t i o n a b l y a b s e n t . Comonly t h e i n n e r n o s t s t r u c t u r e i s a s u b s p h e r i c 2 1 n i c r o s o h e r e w i t h s t r a i g h t l a t t i c e b a r s , from t h e j u n c t i o n s of which a r i s e t h e orimary s o i t i e s . !n some c a s e s t h i s i s t h e o n l y complete l a t t i c e , b u t more corvonly i t i s e n c l o s e d by a very much l a r g e r s n h e r i c a l she11 with c i r c u l a r p o r e s , l e s s commonly by two such shells. A second broad qroiin l a c k s innermost n i c r o s p h e r e s , h a v i r p 1 t o 3 e s s e q t i a l l y s o h e r i c a l s h e l l s wit;? pores of i r r e g u l a r s i z e a n d s h a p e . s h e l l s b e i n ? sometimes i l l - d e f i r e d
so t b , a t a sub-snonay s t r u c t i i r e r e s u l t s .
Palaeoactinowid snjnes are i n v a r i a b : y d e l i c a t e , unbladed, n e e d l e - l i k e , s o v e t i - e s s l i g h t l y c u r v e d . Conterocrzneous E n t a c t i q i i d a e sbnw varioi,s a r c h i t e c t u r e s , a 1 1
170
r a d i c a l l y d i f f e r e n t from Palaeoactinommids.
Important a r e F n t a c t i n i i d
Genus Hovum Fortey and Holdsworth, w i t h subspherical s h e l l developed from l a t e r a l spinules of 3 of 6 near orthogonal, point-centred s p i c u l e r a y s , and a f f . Entactinia Foreman (Fo&y and Holdsworth) spp. These l a t t e r possess crude, sometimes incomplete s i n g l e l a t t i c e s h e l l s generated e i t h e r by terminal ramification of 4 o f 6 very s t r o n q , bar-centred s p i c u l e r a y s , o r by l a t e r a l s p i r u l e s developed from 1 or more rays of a s i m i l a r s p i c u l e , a l l rays continuing as main spines, frequently terminating with simple b i f u r c a t i o n . Examples of o t h e r a r c h i t e c t u r e s a r e very r a r e a n d include a f f . . Stiamosphaera I RQst Fortey and Holdsworth and E r t a c t i n i i d Genus Flovum 2 . This l a s t i s d i s t i n c t i v e , t h e sub-spherical skeleton developivq from a barc e n t r e @ , 6-ray spicule? b v reneated hifurcations of a l l r a y s , r e s u l t i n q soinules n o t fusing. Entactiniidae a r e a l s o probably represented by 6ray, point- and bar-centred spicules unassociated with any t r u e l a t t i c e . Conical Radiolaria Fainilia iiova sap. a r e confined t o t h e higher Spi tsberqen horizon. Arenigiai Entactiniidae have some s i m i l a r i t i e s t o S i l u r i a n forms ( a s have Palaeoactinommids) b u t d i f f e r completely from Late Paleozoic forms. Potasohaerids, Pylentonemidac, a l l A l b a i l l e l l a r i a a n d nrohably a l l Palaeoscenidiidae a r e apoarpntly absent in the e a r l i e s t Ordovician. Adequate younger Ordovician material i s u n k n o w n from the A t l a n t i c Borderlands, b u t Dunhan a n d fYirDby ( i n w c s s : --. iR 1 i t t . l shot, braninian-like Palaeoactinommids ard Entactiniidae t o p e r s i s t in t h e Caradocian of Nevada, together with W’P tim 7Palaeoactinommids and possibly Fntactinia Foreman S . S . sp. SILUEIfi’:: Well-preserved, h i t h e r t o undescribed padiolaria of Llandoverian throuclh llerllockian aqe a r e known from q r a o t o l i t i c cnncretions of t h e Cape P h i l l i p s Fornation, Cornwallis I s l a r d s . Preliminary study of assemblaqes frorr 1 2 borizons shows Palaeoactinornmids, Rotasphaerids, Palaeoscenidiidae and Ceratoi kiscum s p p . t o be t h e numerically dominant qrouns, with l a t t i c e d Entactiniidae l e s s common than in Late Paleozoic faunas atld with hlcderi-spine forms absent. Ceratoikiscum. spp. a r e undetectPd bP1nvJ the Cyrtocjraptus r i g i d u s Tone (Wenlockian). The r e l a t i v e iniportances of environriental change, s e l e c t i v e d i s s o l c t i o n and evolutionary chanoe i n aetermining the wide v a r i a t i o n in assemblage make-up cannnt y e t k assesscr’. 4mongs t Palaeoactinormids , smal l e r sing1 e atxi double she1 led species appear c l o s e l y comparable with Arenigian forms, b u t far g r e a t e r d i v e r s i t y i s now founa i n the qroiip. bximum l a t t i c e s h e l l diaiwter v a r i e s
171
from 0.15mni. t o 0.85mm. The mnst d i s t i n c t i v e forms a r e t h e l a r f l e r , w i t h outermost s i m p l e , r e t i c u l a t e l a t t i c e w i d e l y s e p a r a t e d from one o r n o r e much s m a l l e r , r e t i c u l a t e i n n e r l a t t i c e s .
I n v e s t i r a t i o n o f innermost
s t r u c t u r e i s more d i f f i c u l t t h a n a t Ordovician h o r i z o n s , b u t i n no i n s t a n c e so f a r where i t has been n o s s i b l e t o d e t e r m i n e t h e innermost d e t a i l s of a S i l u r i a n forni i . : C c . t h a n one snheroirial l a t t s c e h a < ? P V ~ T P - Q
t r a c e been found of an e n t a c t i n i a c e i d s p i c u l e . P r e s e n t l y i t can o n l y be concluded t h a t t h e common, m u l t i s h e l l e d and r a r e r sponcjy S i l u r i a n s p h e r o i d s a r e Palaeoactinoniniids. : j u q b 2 r and s p a c i n g o f l a t t i c e s h e l l s i s v a r i a b l e i n t h e g r o u p , a s i s main and s u b s i d i a r y s p i n e c o u n t , b u t s o i n e s a r e i n v a r i a b l y unbladed r o d s - e x c e p t i n t h e i n s t a n c e o f a r e l a t i v e l y r a r e group w i t h d e l i c a t e , sub-sponl;y s h e l l s which n o s s e s s 4 t o Drobahly 6 s t r o n g , t h r e e - b l a d e d major s n i n e s . These a r e t h e e a r l i e s t exaninles of bladed s p i n e s d e f i n i t e l y known t o t h e w r i t e r i n t h e P a l e o z o i c r e c o r d . The Ro t a s p h a e r i d s a r e t o t a 11y d i s t i n c t from t h e Pa l a e o a c t i noiimids i n p o s s e s s i n g o n l y one s n h e r i c a l l a t t i c e s h e l l which i n v a r i a h l y shows a t l e a s t p a r t of t h e s h e l l i n c o r p o r a t i n g n o t a b l y s t r a i q h t b a r s which r a d i a t e from a c e n t r e , o f t e n a s s o c i a t e d w i t h a s n i n e b a s e . P o s t commonly, oerhans i n v a r i a b l y , more t h a n one r a d i a t i o n o o i n t i s p r e s e n t . I n t e r i o r s a r e c l e a r l y s e e n and t h e r e i s no t r a c e of s t r u c t u r e w i t h i n t h e s i n g l e s h e l l . S p i n e s are never l e s s than 6 , commonly more, i n v a r i a b l y r o d - l i k e h u t i n very r a r e i n s t a n c e s w i t h a v e r y s l i g h t s u a g e s t i o n of i n c i p i e n t b l a d i n g . The commonly r o b u s t s h e l l s a r e l i k e l y t o have been s o l u t i o n - r e s i s t a n t , so t h a t t h e p r e s e n c e of t h i s g r o u p i n t h e C o r n w a l l i s S i l u r i a n a s s e m b l a o e s , b u t a p p a r e n t l y i n no d e f i n i t e l y o l d e r o r d e f i n i t e l y younger f a u n a s , may be of particular biostratigraphic significance. E n t a c t i n i i d a e o c c u r i n r a t h e r i n s i g n i f i c a n t numbers. A s i n g l e s p e c i e s of E n t a c t i n i i d Genus tlovum i s very s c a r c e , d i f f e r i n ? fro11 t h e A r e n i g i a n forms i n having rounder p o r e s and more d e l i c a t e s h e l l . I n m a t e r i a l so f a r examined E n t a c t i n i a S.S. i s p r o b a b l y r e n r e s e n t e d by no more t h a n 4 s m a l l , r a r e s p e c i e s , a : l with unbladed s p i n e s and d e l i c a t e or v e r v d e l i c a t e i n t e r n a l s p i c u l e s , and E n t a c t i n o s p h a e r a S . S . I.as n o t h ? ? n d e f i n i t e l y r e c o g n i z e d . C f . E n t a c t i n i a i s r e p r e s e n t e d by even s m r c r r specimens w i t h v e r y s t r o n g , ti-ray, b a r - c e n t r e d s p i c u l e and h i r h l y e c c e n t r i c , c r u d e and probably incomplete r e t i c u l a t e s h e l l g e n e r a t r r ’ by l a t e r a l s p i n u l e s from s p i c u l e r a y s which c o n t i n u e a s r o d - l i k e s p i n e s . E n t a c t i n i i d a e l a c k i n g t r u e , r e t i c u l a t e s h e l l a r e s l i g h t l y more abundant a s a g r o u p , b u t i n d i v i d u a l s p e c i e s e q u a l l y r a r e . Amongst t h e s e , E n t a c t i n i i d Genus Novum 2 i s c l o s e l y comparable w i t h t h e form from t h e
172
7owest A r e n i q i a n .
9 f o r m w i t h b a r c e n t r e and 7 v e r y strong r a y s shows
s p i r u l e s a r i s i n g l a t e r a l l y a t m o r e t h a n one l e v e l f r o m r a y s , n r e f e r e n t i a l l y developed on 4 r a y s .
V o s t a b u n d a n t i s a g r o u p w i t h 6 r a y s and s h o r t o r
v e r y s h o r t b a r c e n t r e h a v i n q l a t e r a l s p i n u l e s a r i s i n q f r o m one o r m o r e l e v e l s , e q u a l l y d e v e l o p e d o n each r a y A f f . Entactinia
-
H a p l e n t a c t i n i a Foreman s . 1 . SDO.
snn. o f A r e n i g i a n t y D e a n n e a r a b s e n t .
The
S i l u r i a n E n t a c t i n i a spp. and H a p l e n t a c t i n i a - l i k e f o r m s Seem t o * c n r e s e i t "advance' i n development r e l a t i v e t o t h ,
b i t i:
pro9ini?p.
ap
vtice?t?lc
i c
t h a t t h e C o r n w a l l i s r e t i c u l a t e F n t a c t i n i i d a e s t i l l a p p e a r t o he s c a r c e r and l e s s d i v e r s e t h a n i n known L a t e P a l e o z o i c P $ 5 p ~ ~ i h l ; r o ? , s r d i t
"1uqC
b~
c o n c l u d e d t h a t t h e m a j o r e n t a c t i n i i d r a d i a t i o n was p o s t - W e n l o c k i a n . P a l a e o s c e n i d i i d a e a r e represented b y coimon Palaeoscenidium s . 1 .
snn.
w i t h 3 o r 4 b a s a l s p i n e s and 2 o r 3 a p i c a l s .
They d i f f e r from
Palaeoscenidium S.S. i n l a c k i n g t h e i m p e r f o r a t e , t e n t - l i k e s h e l l s u r r o u n d i n g t h e p r o x i m a l ends o f t h e b a s a l s , b u t b a s a l s a r e ornamented b y n e r d a n t s p i n u l e s , sometimes a m a l g a m a t i n g t o f o r m a n i r r e g u l a r , p e r f o r a t e "tent".
The e n t a c t i n i i d - l i k e b a r f r o m w h i c h b a s a l s and a p i c a l s a r i s e i s
c l e a r and c a n b e d e t e c t e d i n L a t e P a l e o z o i c P a l a e o s c e n i d i u m S.S. SPD.
This
bar i s a l s o c l e a r i n a f u r t h e r , w i d e l y d i v e r s e group o f s p i c u l a r forms here Such f o r m s d i f f e r f r o m
r e f e r r e d t o as " a f f . P a l a e o s c e n i d i u m s p p . " . H a p l e n t a c t i n i a s.1,
i n t h e markedly d i f f e r e n t i a l develonment o f t h e p r i m a r y
s p i c u l e r a y s , and t h e r e i s a c l e a r a p p r o a c h t o t h e b a s a l / a p i c a l p o l a r i s a t i o n c h a r a c t e r i s t i c o f P a l a e o s c e n i d i u v s . 1 . and
S.S.
S i n q l e and
commonly m u l t i p l e b i f u r c a t i o n t e n d s t o mark t h e s t r o n g e r r a y s : l e s s e r l e n g t h , a b s e n c e o f b i f u r c a t i o n o r weak s i n q l e b i f u r c a t i o n marks t h e l e s s developed, a p i c a l analogues. Palaeoscenidium s.1.
Pendant l a t e r a l s p i n u l e s , r e m i n i s c e n t of
a r e f o u n d o n some r a y s .
It i s temntino
f o r r > s a s i n t e r m e d i a t e s between H a p l e n t a c t i n i a s . 1 . Palaeoscenidium s.1.
(Palaeoscenidiidae)
-
to v i e w such
( F n t a c t i n i i d a e ) and
and hence t h e t e p t a t i v e
assignment o f Palaeoscenidiidae t o t h e Suoerfamily Entactiniacea (see S y s t e m a t i c Framework)
I
C e r a t o i k i s c u m sap. and e x c e s s i v e l y s c a r c e R a d i o l a r i a Genus B Foreman sp. r e p r e s e n t t h e f i r s t known a p p e a r a n c e o f C e r a t o i k i s c i d a e and o f A l b a i l l e l l a r i a outside Asia,
(Nazarov
3. (1975)
d e s c r i b e d t h e new,
p o s s i b l y d o u b t f u l C e r a t o i k i s c u m a c a t a n g u l a t u m from t h e L l a n d e i l o - L o w e r Caradoc o f K a z a k h s t a n . )
Absence o f b o t h g e n e r a i n t h e l o w e r p a r t o f t h e
Cape P h i l l i p s F o r m a t i o n ( L l a n d o v e r i a n l e v e l s ) may w e l l b e d u e t o s o l u t i o n . C e r a t o i k i s c u m spp. show i d e a l , r a t h e r d e l i c a t e f r a m e s w i t h no o r weak p a t a g i u m and s l e n d e r c a v e a l r i b s
-
a f e w commonly d e v e l o p e d f r o m t h e d o r s a l
173 end o f t h e b - r o d
-
ornamented o n l y w i t h m i n u t e s e r r a t i o n s .
A v e r y few
" m u t a n t " specimens show d o u b l i n g o f one o r m o r e e x t r a - t r i a n g u l a r r o d s , o r g r o s s d i s t o r t i o n of t h e u s u a l , e s s e n t i a l l y e q u i l a t e r a l f r a m e .
A l l forms
a r e t o t a l l y d i s t i n c t f r o m known L a t e P a l e o z o i c s p e c i e s . O f r a r e , S p i c u l a r Forms, m o s t s i g n i f i c a n t i s a t y o e w i t h 8 c u r v i n g s p i n e s r a d i a t i n g f r o m t h e end o f a s h o r t " s t e m " , r e g u l a r l y spaced s p i n u l e s .
each s n i n e ornamented w i t h
S i m i l a r b!tt s i m p l e r f o r m s a r e o f t e n a b u n d a n t a t
Upner P a l e o z o i c l e v e l s . Forms n o t p r e s e n t i n C o r n w a l l i s assemblages b u t known i n Eurooean S i l u r i a n f a u n a s a r e a p p a r e n t p y l o m a t e P a l a e o a c t i n o m m i d s o f Germany ( S t d r m e r 1966) and A r c h o c y r t i u m D e f l a n d r e s p n . ( D e f l a n d r e 1 9 7 2 ) , p r o b a b l y W e n l o c k i a n , and f r o m B r g t i g n o l l e s , F r a n c e .
These l a t t e r a r e , i n t h e 1
w r i t e r ' s v i e w , t h e e a r l i e s t c o n v i n c i n g P y l e n t o n e m i d a e known.
LATE SILURIAN, EARLY AND MIDDLE D E V O I I I A I I : Adequate assemblages f r o m t h i s c o n s i d e r a b l e i n t e r v a l a r e unknown t o t h e w r i t e r f r o m t h e A t l a n t i c B o r d e r l a n d s , so t h a t , t o i d e n t i f y t h e t i m e o f m a j o r assemblage change w h i c h , i t a p p e a r s , t o & p l a c e b e t w e e n W e n l o c k i a n an? F r a s n i a n , i t i s n e c e s s a r y t o l o o k e l sewhere.
blast s i g n i f i c a n t i s H i n d e ' s ( 1 8 9 9 ) a c c o u n t o f "Tamworth S e r i e s " f o r m s (New S o u t h Wales) w h i c h , i t w o u l d a p o e a r , m u s t b e from r o c k s of E a r l y - M i d d l e D e v o n i a n a g e . Though i t i s i m p o s s i b l e t o D i n - o o i n t t h e h o r i z o n o f a n y d e s c r i b e d s p e c i e s , t h e e n t i r e assemblage i s c l e a r l y m o r e c l o s e l y r e l a t e d t o w e l l - d a t e d L a t e Devonian faunas than t o C o r n w a l l i s S i l u r i a n assemblages. Nazarov ( 1 9 7 5 ) a l s o i n d i c a t e s t y p i c a l L a t e D e v o n i a n forms e x t e n d i n g i n t o t h e b l i d d l e D e v o n i a n of K a z a k h s t a n . I n b o t h r e g i o n s , e s s e n t i a l l y s p h e r o i d a l f o r m s 131;tb i l 2 , l c . i s n i p e s o r e d o m i n a t e - a s i g n i f i c a n t percentage, a t l e a s t , beins E n t a c t i n i i d a e . Thus A b e r d e e n ' s (194U) "Uevoniarl" t a u n a f r o m t h e S a n t i a g o Vember of t h e C a b a l l o s I.lovacu1 i t e , Texas, a m e a r s anomalous i n a p p a r e n t l y l a c k i n g E n t a c t i n i a c e a c o m p l e t e l y ( R i e d e l and Foreman 1 9 6 1 ) . The age o f t h e R o v a c u l i t e and t h e p o s i t i o n w i t h i n i t o f t h e S a n t i a g o F4emher a r e b o t h q u e s t i o n a b l e ( c f . N i g r i n i and N i t e c k i 1 9 6 8 ) . Lower D e v o n i a n c o n o d o n t s from t h e " u p p e r " p a r t o f t h e N o v a c u l i t e ( G r a v e s 1952 : G l e n i s t e r and K l a p o e r ( i n l i t t . ) ) r a i s e t h e p o s s i b i l i t y t h a t some p a r t o f t h e u n i t m i o h t p o s s i b l y b e e v o n i a n , and t h e g r o s s a s p e c t o f t h e S a n t i a F o assemblage, as d e s c r i b e d , seems t o i n v i t e some c o m p a r i s o n w i t b t h e l e s s d i v e r s e o f C o r n w a l l i s p r e - L u d l o v i a n f a u n a s . S a n t i a q o " H e x a s t y l u s " and "He1 i o s p h a e r a " spp. as r e f i g u r e d b y R i e d e l and Foreman ( 1 9 6 1 ) a p p e a r t o b e p o s s i b l e R o t a s p h a e r i d s and t h e l a r g e r mu1 t i s h e l l e d S a n t i a g o s p h e r o i d s compare w i t h C o r n w a l l i s P a l a e o a c t i n o m m i d s . O f S a n t i a g o f o r m s , o n l y " X i p h o s t y l u s " and " S t y l o s p h a e r a " a r e o b v i o u s l y f o r e i g n t o t h e C o r n w a l l i s S i l u r i a n and c l o s e l y c o m p a r a b l e w i t h Tamworth f o r m s . B u t no r e l i a b l e d a t e e x i s t s f o r t h e f i r s t a p p e a r a n c e o f t h e s e and c l o s e l y s i m i l a r f o r m s w i t h s p h e r o i d a l t o s l i g h t l y e l l i p s o i d a l s h e l l s and o n l y two p r o m i n e n t s p i n e s , and i t i s n u i t e p o s s i b l e t h a t they are o f l a t e S i l u r i a n o r i g i n . The u n q u e s t i o n a b l e c o n t r a s t s between p r o v e n b l e n l o c k i a n and p r o v e n F r a s n i a n assemblages ( s e e b e l o w ) c o u l d h a v e been i n i t i a t e d p r i o r t o t h e 1. F ' n o t e : M i d . O r d o v i c i a n " P y l e n t o n e m a " spp. (Plazarov e t a l . 1975) a p p e a r from t h e i r d e s c r i p t i o n s t o b e p y l o m a t e a e o a c t i n o m m i d s .
174
l l i t i d l e D e v o n i a n and q u i t e p o s s i b l y i n l a t e s t S i l u r i a n t i m e .
LATi CEVO':IA:;:
M o s t c o m p l e t e l y d e s c r i b e d a n d i l l u s t r a t e d o f known P a l e o z o i c
assevblages a r e those
-
3f
ti?. F u r q q Eleriber o f t h e O h i o S h a l e (Foreman l C 6 3 )
F w e n n i a n i n a?.: o , i co.iouhni-. w i : ! e i l c e
(Hass 1947 : G l e n i s t e r and K l a n o e r
(-. i n l i t t . ) ) . E n t a c t i n i i d a e doiiiinate t h e t h r e e assemblages s t u d i e d , r e p r e s e n t e d by t h e genera H a n l e n t a c t i n i a , E n t a c t i n i a , Entactinosnhaera F o r e r i a n , P o l y e n t a c t i n i a and T e t r e n t a c t i n i a Foreman.
O f 44 e s s e n t i a l l y
s p h e r i c a l ? l a t t i c e d species, 3 o n l y riay p o s s i b l y l a c k an e n t a c t i n i i d s p i c u l e a n d be r e f e r a b l e t o t h e P a l a e o z c t i n o r i m i d s . absent.
Contrast w i t h Cornwallis S i l u r i a n
Qotiisnktaeri5s a r e
~ ! s s ? - ~ ! ? ~ E :s s
C1-81.l~
-?rL:d ? i d
f u r t h e r under7ined by t h e prevalence o f E n t a c t i n i i d a e w i t h bladed spines and c o n c e n t r i c l a t t i c e s
-
unknown i n t h e Lower P a l e o z o i c .
P a l a e o s c e n i d i i d a e show a m a r k e d d e c l i n e ,
now r e p r e s e n t e d b y o n l y t w o
soecies, Palaeoscenidium cladophorum D e f l a n d r e and q u a d r i r a m o s s Foreman
-
Palaeoscenidium ?
t h e l a t t e r a p p a r e n t l y a remnant o f t h e S i l u r i a n
a f f . P a l a e o s c e n i d i x stocks.
C e r a t o i k i s c i d a e a r e reprosented by 4 snecies
o f C e r a t o i k i s c u m a n d one R a d i o l a r i a Genus B s p e c i e s , a l l v e r y d i s t i n c t f r o i n S i l u r i a n forms.
The m o s t s i g n i f i c a n t i n n o v a t i o n i n t h i s f a m i l y i s t h e
f i r s t a p p e a r a n c e o f a s h e l l e d genus
f i l b a i l l e l l i d a e a r e absent.
-
H o l o e c i s c u s Foreman
-
but
C y r t e n t a c t i n i a p r i m o t i c a Foreman i s h e r e
considered t h e e a r l i e s t r e p r e s e n t a t i v e o f t h e P o p o f s k y e l l i d a e , and F o r i ~ i a n i e l l a c i b d e l o s p h a e r a (Foreman) i s t h e e a r l i e s t known o f t h e Pylentonemidae a p a r t from A r c h o c y r t i c spp. f i g u r e d by Deflandre (1972) from t h e " S i l u r i a n " o f B r c t i g n o l l e s (see above).
The p r o b l e m a t i c
C o r y t b o e c i a i s f i r s t known f r o m t h e H u r o n llember, accompanied b y s e v e r a l S p i c u l a r Forms. P,n o l d e r , e a r l y F r a s n i a n ( P o l y a n a t h u s a s y m m e t r i c u s Zone) assemblage i n e x c e l l e n t n r e s e r v a t i o n i s known f r o m t h e Canol S h a l e , N o r t h
blest T e r r i t o r i e s ( M a c K e n z i e a n d t i o l d s w o r t h , i n p r e p a r a t i o n ) . Pylentonemidae, P o p o f s k y e l l i d a e and Holoeciscus a r e unrenresented, Haplentactinia excessively scarce.
I n other resnocts +ha assaihlan?
compares c l o s e l y w i t h t h e H u r o n f a u n a s , c e r t a i n e n t a c t i n i i d s p e c i e s beinq i d e n t i c a l , others c l o s e l y r e l a t e d .
P a l a e o a c t i n o r m i d s h a v e n o t been
d e t e c t e d and R o t a s p h a e r i d s and A l b a i l l e l l i d a e a r e a b s e n t . P a l a e o s c e n i d i i d a e a r e r e p r e s e n t e d by P . cladonhorum and
0.a f f .
cladonhorum
C e r a t o i k i s c i d a e a r e r i c h e r t h a n i n t h e Huron assemblages, C e r a t o i k i s c u m snn m o r e d i v e r s e , so t h a t t h e a b s e n c e o f H o l o e c i s c u s s t r o n g l y s u g a e s t s t h a t t h e oenus i s o f p o s t - e a r l y F r a s n i a n o r i g i n .
3 o f t h e 4 Huron Ceratoikiscum spa
a r e c l e a r l y r e l a t e d t o Canol s t o c k s , b u t m i n o r s p e c i f i c d i f f e r e n c e s a r e
175
d e t e c t a b l e and probably of b i o s t r a t i g r a p h i c u t i l i t y . P d i s t i n c t i v e Canol v a r i e t y of C e r a t o i kiscum b u j u g u m Foreman i s r e c o g n i z a b l e i n low F r a s n i a n f a u n a s of Nevada and Western A u s t r a l i a . Caveal r i b vane development i n Ceratoikiscum s p i n o s i a r c u a t u m Foreman s . 1 . Group i s g r e a t e r t h a n a t Huron h o r i z o n s and i n marked c o n t r a s t t o Wenlockian development. Within C e r a t o i k i s c u m p l a n i s t e l l a r e Foreman Group e x i s t s a Canol form c l o s e t o Ceratoikiscum avimexpectans D e f l a n d r e ( T o u r n a i s i a n ) , t h e Canol form being unseen a t Huron l e v e l s b u t d e t e c t a b l e i n t h e low F r a s n i a n o f Idestern A u s t r a l i a . C e r t a i n C e r a t o i k i s c u m s t o c k s , a t l e a s t , a p p e a r t o have had a world-wide d i s t r i b u t i o n a t L a t e Devonian t r o p i c a l l a t i t u d e s and a r e r e c o g n i z a b l e i n t h e U r a l s (Nazarov 1973, 1 9 7 5 ) . CARBONIFEROUS: The e a r l i e s t a d e q u a t e fauna i s of t h e Montagne Noire, Southern F r a n c e ( e g . D e f l a n d r e 1 9 6 0 ) , p r o b a b l y T o u r n a i s i a n and c l o s e l y comparable w i t h a proven T o u r n a i s i a n T u r k i s h assemblage (Holdsworth 1 9 7 3 ) . Most i m p o r t a n t i s t h e f i r s t a p p e a r a n c e of t h e A l b a i l l e l l i d a e , r e p r e s e n t e d i n F r a n c e by a t l e a s t 3 A l b a i l l e l l a s p p . and 1 s p e c i e s of Lapidopiscum D e f l a n d r e . Holoeciscus i s now a b s e n t , b u t 5 and perhaps 6 C e r a t o i k i s c u m s p p . were f i g u r e d by D e f l a n d r e ( 1 9 6 0 ) . One may be i d e n t i c a l w i t h t h e Famennian C . s p i n o s i a r c u a t u m s . s . , b u t both French and Turkish T o u r n a i s i a n forms are m o s t l y c l e a r l y d i s t i n g u i s h a b l e from L a t e Devonian s p e c i e s , though r e l a t e d i n some c a s e s . P a r t i c u l a r l y s i g n i f i c a n t i s t h e a n n e a r a n c e of a T u r k i s h s p e c i e s w i t h t h e i d e a l 6 e x t r a t r i a n g u l a r s o i n e s reduced t o 4
"C.
( ? ) apertum" D e f l a n d r e of F r a n c e ) . (possibly Very s t r i k i n g i n French and T u r k i s h f a u n a s i s t h e abundance and d i v e r s i t y of Pylentonemidae - v a s t l y ( I r e d t e r t h a n i n t h e I h r t h h c i - i c a n
l a t e s t Devonian. Deflandre gave t h e names Pylentonerna, Cerarchocyrtium Cyr t i s p ha e r a c t en i um , Cyr t i s p!?a e r o n em i urn, Pa r a a r c h o c y r t i urn a nd I,r c h o c y r t i E t o encompass t h o v a r i a + : n - h i i t classification renains unsatisfactory. F o r m a n i e l l a S . S . a p p e a r s t o be a b s e n t . P o n o f s k y e l l i d a e a r e r e n r e s e n t e d b,v t h e e a r l i e s t known s p e c i e s of Popofskyellum and T u s c a r i t e l l u m , b u t t h e Famennian C y r t e n t a c t i n i a S . S . i s u n d e t e c t e d . Palaeoscenidium cladoDhorum p e r s i s t s from t h e Famennian, b u t t h e o n l y o t h e r r e p r e s e n t a t i v e of t h e P a l a e o s c e n i d i i d a e i s t h e nevi s p e c i e s Palaeoscenidiuri b i c o r n e D e f l a n d r e . Non-pylonate T o u r n a i s i a n s p h e r o i d s a r e v e r y n o o r l y kiown. A nu-hcr oc s p e c i e s a r e E n t a c t i n i i d a e , b u t i n t e r n a l s t r u c t u r e i s seldom c l e a r i n 1
material presently available.
Visebn R a d i o l a r i a remain very p o o r l y known. Re-examination of G.J. H i n d e ' s l a t e Visgan c h e r t s l i d e s from S.W. EnGland (Hinde and Fox 1895) r e v e a l s h i g h l y d i s s o l v e d , p a r t l y c r u s h e d assemblages i n which t h e 1 . F ' n o t e : T r i s e n o s p b a e r a D e f l a n d r e may be a t r u e Palaeoactinommid.
~
-
176 o n l y f o r m s a b l e t o be r e l i a b l y i d e n t i f i e d a r e a few E n t a c t i n i i d a e . H i n d e ' s " L i t h o c a m p e " and " S t i c h o c a p s a " spp. a r e p r o b a b l y A l b a i l l e l l a spp. o u t l i n e s , a n d " P o r o d i s c u s " s p p . a r e f a i r i y c l e a r l y c r u s h e d and D a r t i a l l y dissolved spheroids,
A l a t e Vise/an ( P 2 b )
i n some c a s e s E n t a c t i n i i d a e .
horizon i n black, goniatite-bearing,
shales o f County L e i t r i m , I r e l a n d ,
c o n t a i n s d i s t i n c t i v e , new R a d i o l a r i a Genus B s p p . a c c o m p a n i e d b y t h e e a r l i e s t known i n f l a t e d , b i - w i n g e d A l b a i l l e l l a s p . o f A l b a i l l e l l a u e n n a t a H o l d s w o r t h Group. A . p e n n a t a G r o u p i s a l s o known f r o m a v e r y l o w Namurian ( E l ) g o n i a t i t e h o r i z o n o f t h e F a y e t t e v i l l e Shale, Arkansas, a s s o c i a t e d w i t h a second s p i n o s e A l b a i l l e l l a s n . and s o h e r o i d s w i t h e x t e r n a l c h a r a c t e r i s t i c s t y o i c a l o f Late Paleozoic
E n t a c t i n i i d a e ( N i g r i n i and N i t e c k i 1968).
The Namurian r e c o r d i s a n t i n u e d b y o f t e n e x q u i s i t e l y p r e s e r v e d R a d i o l a r i a
i n t h e b l a c k s h a l e g o n i a t i t e bands o f S t a f f o r d s h i r e and D e r b y s h i r e , - E2b t h r o u g h e a r l y P2a i n a g e ( t i o l d s w o r t h 1964. 19fi6). 1 7 t h e s e
England
bands, E n t a c t i n i i d a e o f t h e g e n e r a P o l y e n t a c t i n i a predominate.
T e t r e n t a c t i n i a i s oresc:!t,
d i v e r s i t y t h a n i n t h e Famennian. are absent.
E n t a c t i n i a , E n t a c t i n o s p h a e r a and b u t shows l e s s
H a p l e n t a c t i n i a and a l l P a l a e o s c e n i d i i d a e
C o n v i n c i n g e v i d e n c e o f P a l a e o s c t i n o m n i d s has n o t y e t been
f o u n d , and t h e r e i s a s t r i k i n g a b s e n c e o f f o r m s w i t h o n l y t r i o , m a j o r s p i n e s , known i n D e v o n i a n f a u n a s ( s e e above, L a t e S i l u r i a n , E a r l y and i ' i d d l e Devonian). Archocyrtium. and
A. c f .
P y l e n t o n e m i d a e a r e known o n l y b y a s i n a l e s n e c i m e n o f
A l b a i l l e l l i d a e a r e r e p r e s e n t e d m o s t commonly b y A . o e n n a t a
pennata (Holdsworth 1966a), spinose sDecies b e i n g e x c e s s i v e l y
r a r e : Lapidopiscum i s absent.
C e r a t o i k i s c i d a e i n c l u d e R a d i o l a r i a Genus K
s p p . , q u i t e d i s t i n c t from a l l o l d e r forms, and C e r a t o i k i s c u m sop., most commonly o f C e r a t o i k i s c u m b i c a n c e l l a t u m t i o l d s w o r t h a n d C e r a t o i k i s c u m t r i c a n c e l l a t u m H o l d s w o r t h Grouus ( H o l d s w o r t h 1 9 6 9 a ) d i s t i n c t f r o m a l l known Dre-IlamLirian s p e c i e s .
-
again, o u i t e
Popofskyellidae are
r e o r e s e n t e d o n l y by e x t r e m e l y r a r e , s p o r a d i c Ponofskyellum snn., and t h e l a s t known o c c u r r e n c e s o f b o t h P o o o f s k y e l l i d a e and C e r a t o i k i s c i d a e a r e i n
All A l b a i l l e l l a r i a a r e r e s t r i c t e d t o r e l a t i v e l y few o f t h e R a d i o l a r i a n h o r i z o n s , and i t i s a t a f e w h o r i z o n s w h e r e t h e r a r e , Zone r i l a .
P o p o f s k y e l lum and sometimes c f . C o r y t h o e c i a specimens a r e f o u n d .
SDicular
Forms a r e f r e o u e n t a t many l e v e l s anti h a v e L a t e D e v o n i a n a f f i n i t i e s . Y o u n g e s t known o f a d e q u a t e P a l e o z o i c assemhlaoes i s t h a t from Eoasianites-bearing,
sponge-rich c o n c r e t i o n s of t h e I n t a r a r e ' Formation,
l i r u q u a y ( s e e e g . Kling a n d ? e i f 1 9 6 9 ) .
Age i s q u e s t i o n a b l e b u t a p p a r e n t l y
w i t h i n t h e i n t e r v a l i<'estohalian ( L a t e Carboniferous) (Pansbottom ners
177 comm.) t o E a r l y P e r m i a n ( B h a r a d w a j 1 9 6 9 ) .
Though o f l e s s e r a g e and
a p p a r e n t l y s i g n i f i c a n t l y h i g h e r l a t i t u d e ( s e e b e l o w ) t h a n E n g l i s h Flamurian f a u n a s , t h e g r o s s a s p e c t o f t h e I n t a r a r e ' a s s e m b l a q e is r a t h e r s i m i l a r . P r e l i m i n a r y s t u d y r e v e a l s a t l e a s t 24 s p e c i e s of E n t a c t i n i i d a e , m a i n l y E n t a c t i n i a a n d E n t a c t i n o s p h a e r a s p p . , b u t a t l e a s t 1 s p e c i e s of
-
Polyentactinia i s present E n g l i s h llamurian species occurs a l s o .
-
c l o s e a n d p e r h a p s i d e n t i c a l t o a common and a n unnamed E n g l i s h F!amurian g e n u s n r o b a b l y
P r e s e n c e o f P a l a e o a c t i n o m m i d s has n o t been e s t a b l i s h e d .
P y l e n t o n e m i d a e , P a l a e o s c e n i d i i d a e , P o n o f s k y e l l i d a e and C e r a t o i k i s c i d a e a r e absent.
A l b a i l l e l l i d a e a r e represented b y a t l e a s t two A l b a i l l e l l a s o p . ,
one a m a r k e d l y s p i n o s e f o r m , b u t A . p e n n a t a G r o u p i s p r o b a b l y a h s e n t . S p i c u l a r Forms a r e c l o s e l y c o m p a r a b l e t o t y p e s known i n t h e E n g l i s h Namurian, a n d t h e r e o c c u r s a l s o a r e m a r k a b l e s c a l a r i f o r m I n c e r t a e S e d i s , f i r s t seen a t E n g l i s h Namurian h o r i z o n s .
P E R M I A N : O t h e r t h a n t h e I n t a r a r e / a s s e m b l a g e , w h i c h c o u l d b e of E a r l y P e r m i a n age, no a d e q u a t e f a u n a s f r o m t h i s System a r e y e t known t o t h e writer.
T h a t a m a j o r r e v o l u t i o n i n R a d i o l a r i a h i s t o r y o c c u r r e d somewhere
between L a t e C a r b o n i f e r o u s and J u r a s s i c i s unauestionable,
but the present
p a u c i t y o f r e l i a b l e i n f o r m a t i o n from t h e Permo-Trias leaves t h e t i m e o f t h i s m a j o r change i n d o u b t .
No A l b a i l l e l l a r i a , E n t a c t i n i a c e a ,
P o c o f s k y e l l i d a e o r P a l a e o s c e n i d i i d a e h a v e , t o t h e w r i t e r ' s k n o w l e d a e , been r e l i a b l y r e p o r t e d f r o n i M e s o z o i c r o c k s , and it has a l r e a d y b e e n n o i n t e d o u t t h a t t h e A c t i n o m m i d - l i k e f o r m s o f t h e P a l e o z o i c ( P a l a e o a c t i n o m m i d s ) may be unrelated t o the post-Paleozoic Actinomidae.
Eut the :lassellaria
-
t h e w r i t e r ' s v i e w u n r e p r e s e n t e d b y a n y r e l i a b l y known P a l e o z o i c form
in
-
were i n e x i s t e n c e by T r i a s s i c ( A n i s i a n - R h a e t i a n ) t i m e , r e n r e s e n t e d by t h e Theoceridae (Riedel 1967).
Thus a f u n d a m e n t a l s t e p i n t h e e v o l u t i o n o f
t h e R a d i o l a r i a had o c c u r r e d as e a r l y , p e r h a p s , a s ) l i d d l e T r i a s , and t h e r e c o r d o f t h e s t r i k i n g p r e - J u r a s s i c e x t i n c t i o n s and i n n o v a t i o n s i s p r o b a b l y t o b e s o u g h t i n L a t e P e r m i a n and E a r l y T r i a s s i c a s s e m b l a g e s . PnLEnLATITUDE AND PALEOOCEAPJOGQAPHY: I t i s u n l i k e l y t h a t t h e f e w f a u n a s discussed above, w i d e l y senarated b o t h i n space and t i n e , p r o v i d e a f u l l y r e n r e s e n t a t i v e r e c o r d of P a l e o z o i c ?aaiolaria evolution. 411 b u t one f a u n a a p p a r e n t l y b e l o n g e d t o a b e l t v t i t h i n 2 5 O o f t h e equator.
Hi?h l a t i t u d e forms a r e r e p r e s e r t e d o n l y i n t h e ? L a t e
C a r b o n i f e r o u s Uruauayan assemblage which c o u l d have o r i g i n a t e d a t l a t i t u d e g r e a t e r t h a n 50's.
I t i s i m p o r t a n t t o n o t e , however, t h a t i n g r o s s a s D e c t
t h i s f a u n a i s c o m p a r a b l e w i t h some t r o p i c a l E n a l i s h bdamurian a s s e m b l a g e s .
178
P a l e o e n v i r o n m e n t c o u l d h a v e been a m o r e i m n o r t a t i t r a c t o r i n d e t e r m i n i n g a s s e m b l a g e make-up.
None o f t h e f a u n a s d e s c r i b e d a b o v e c a n be
considered t o occur i n t r u l y oceanic sediments. t o show maximuni d i v e r s i t y i n open o c e a n s .
Modern R a d i o l a r i a a m e a r
S h a l l o w , e n s i a l i c m a r g i n a l seas
a p p e a r t o h a v e r e s t r i c t e d , endemic p o p u l a t i o n s , s e a s o n a l l y e n r i c h e d b y i n n r e s s o f o p e n - o c e a n f o r m s i n some c a s e s ( B j 6 r k l u n d 1 9 7 4 ) .
P o r t i o n s of
ex t e n s iv e , com p a r a t iv e l y s ha 1 1 ow e p i c o n t inen t a 1 s e a s , r e 1 a t iv e l y d is t a n t from c o n n e c t i o n w i t h c o n t e m p o r a n e o u s oceans m i g h t be e x p e c t e d t o h a v e c o n t a i n e d R a d i o l a r i a p o p u l a t i o n s o f somewhat l o w e r d i v e r s i t y t h a n t h e oceans t h e m s e l v e s , a n d d i v e r s i t y c o u l d w e l l h a v e been c l o s e l y r e l a t e d t o ocean n r o x i n i t y .
S l i g h t b u t i n t r i g u i n g h i n t s o f such a r e l a t i n n s h i n a r e
d e t e c t a b l e i n t h e known P a l e o z o i c r e c o r d . The p r o f u s i o n o f e a r l i e s t C a r b o n i f e r o u s P y l e n t o n e m i d a e i n S o u t h e r n France and Turkey c o n t r a s t s s t a r k l y w i t h t h e v i r t u a l absence o f t h e f a m i l y i n t h e E n g l i s h e a r l y t o m i d rlamurian.
E v o l u t i o n a r y d e c l i n e may b e t h e
e x p l a n a t i o n , b u t f r o m E a r l y P a l e o z o i c t i n i e S o u t h e r n F r a n c e and T a r k e y c a n be i i r t e r p r e t e d a s h a v i n g d i r e c t l y b o r d e r e d a i i a j o r " T e t h y a n " Ocean (Oriden
eal.
1974), whereas t h e E n g l i s h Namurian accumulated i n an
e p i c o n t i n e r t a l embayment (Ranisbottom 1971) t o w h i c h o c e a n i c w a t e r may w e l l have b e e n d e n i e d e a s y a c c e s s .
Conceivably, l o c a t i o n a t t h e "Tethys" [margin
a l s o e x p l a i n s t h e S i l u r i a n apoearance o f Pylentonemidae a t Br;ti?nolles
-
arioi1;alously e a r l y r e l a t i v e t o t h e known f i r s t o c c u r r e n c e i n I.!ortt- A m e r i c a . !.I4t!:ir
t h e s h a l l o w e r and more enclosed o f Pa!eozoic e n i c o n t i n e n t a l seas,
s a ? i n i t y and t e m o e r a t u r e n a y h a v e f a v o u r e d s n e c i a l i s e d n o n u l a t i o n s , d e n t h s
beina i n s u f f i c i e n t f o r t h e establishment o f lower water l a y e r s canable o f accommodatiqg m o r e open-sea f o r m s .
C o m p a r a t i v e l y f e w E n o l i s h ','amurian
m a r i c e bands c o n t a i n s i g n i f i c a n t . n u m b e n o f 4 l b a i l l e l l a r i a , and t h e r e i s s o ~ ei n d e n e n d s n t g e o l o g i c e v i d e n c e t o s u g g e s t t h a t a p p e a r a n c e s o f t h e s u b o r d w c o r r e s n o n d w i t h D e r i o d s o f r e l a t i v e l y h i g h sea l e v e l w i t h i n t h e
Centra; Ergland Pasin. The p o s s i b i l i t y c a n n o t b e i a n o r e d t h a t o t h e r , a p n a r e n t l y a g e dia?nostic,con'rasts
b e t w e e n t h e f e w F a l e o z o i c f a u n a s a v a i l a b l e a r e i n some
c e s e s no niore t h a n a r e f l e c t i o n o f d i f f e r e n c e s
i n t b e environments o f the
asserlblages concerned. Aberdeen. E. Basil:,
!94n. R a d i o l a r i a n f a u n a o f t h e Cabal 10s f o r m a t i o n , ' f a r a t h o t 1
Texas: J o u r . P a l e o n t . . v . 1 4 , 11.127-139.
1974. Th? o l d e s t k w w n w e l l - p r e s e r v e d Q a d i o l z r i a n s f r o m Gergstr8~,1, S."1, "ort!i Aver-ica: A b s t . Pros. Geol. SOC. Am., v.6, ~ ~ 4 9 1 , E h a r a d w o j , D.C. 1?.:9. L o w e r Gondwana Fori9at:ons: Coiilnte ?endue S i x i ' e r i e ? . I n t e r n a t . d e S t r a t . e t de G 6 o l . 1u C a r b o n i f & - e , v . 1 , p.255-278. l u r i d , K.F., 1974. The s e a s o n a l o c c u r r e n c e and d e p t h z o n a t i o n of i o l s r ' a r s i r i Korsfjorc p a y : S a r s i a , v . F , n.13-42. b i d e n . ?.C. , D r e w r y , G.C. S n i t + , A . G . , 1976. Phanerozoic eoual-area pmt-!d ~ l ' a p s : J o u r . G e o l . ,
179
Deflandre, G . , 1960. A propos du d6veloapment des recherches sur l e s R a d i o l a i r e s f o s s i l e s : Rev. Pticropale'ont., v . 2 , p.212-218. ------------- 1972. Le syst'eme t r a b 6 c u l a i r e i n t e r n e chez l$s Pylentongmides e t l e s P o p o f s k y e l l i d & , R a d i o l a i r e s du Paleozoique. Phylogenese d e s N a s s e l l a i r e s : C . R . Acad. S c i . P a r i s , t . 2 7 4 , p.3535-3540. Dunham, J.B. and Murphy, M . A . , i n p r e s s . A n o c c u r r e n c e of well-oreserved R a d i o l a r i a from the Upper Ordovician ( C a r a d o c i a n ) , Eureka County, Nevada: J o u r . P a l e o n t . Foreman, H . P . , 1963. Upper Devonian R a d i o l a r i a from t h e Huron member of t h e Ohio S h a l e : Micropaleont., v . 9 , 0.267-304. F o r t e y , R . A . and Holdsworth, B . K . , 1972. Theoidestl'nown well-oreserved R a d i o l a r i a : B o l l . d e l l a SOC. P a l e o n t . T t a l i a n a , v . 1 0 , r1.35-41. Graves, R . W . , J r . , 1952. Devonian conodonts from t h e Caballos n o v a c u l i t e : J o u r . P a l e o n t . , v . 2 6 , p.610-612. Hass, N . H . , 1947. Conodont zones i n Upper Devonian and Lower V i s s i s s i p p i a n formations of Ohio: J o u r . P a l e o n t . , v.27, p.131-141. Hinde, G.J., 1899. O n t h e R a d i o l a r i a i n t h e Devonian rocks of liew South Wales: Quart. J o u r . Geol. SOC. London, v.55, p.38-64. ----------- and Fox, H . , 1895. On a well-marked horizon o f R a d i o l a r i a n rocks i n t h e Lower Culm Pleasures o f Devon, Cornwall and West Somerset: Q u a r t . J o u r . Geol. S o t . London, v.51, p.609-688. Holdsworth, B . K . , 1964. R a d i o l a r i a n n a t u r e o f t h e t h i c k e r - s h e i l e d g o n i a t i t e f a u n a l phase i n some Navuriari l i m e s t o n e " b u l l i o n s " : Nature London, v.201, p.697-699. ------------__-- 1966. A p r e l i m i n a r y s t u d y o f the oalaeontoloqy and palaeoenvironment o f some Namurian 1 imestone " b u l l i o n s " : Wercian G e o l o g i s t , v.1, p.315-337. - _ _ _ _ _ _ _ _ _ _ _1966a. _ _ _ _ R a d i o l a r i a from t h e h'amurian o f Derbyshire. Palaeontology, v . 9 , p.319-329. - - _ _ _ - _ _ _ - _ _ _1969. _ _ _ The r e l a t i o n s h i p between t h e aenus A l b a i l l e l l a Deflandre and t h e c e r a t o i k i s c i d R a d i o l a r i a : b l i c r o p a l e o n t . , v.15, p. 230-236. ---------------- 1969a. llamurian R a d i o l a r i a of the genus Ceratoikiscui. from S t a f f o r d s h i r e and Derbyshire, England: P l i c r o a a l e o n t . , v.15, p , 221 - 229. ---------------- 1973. The R a d i o l a r i a of the B a l t a l i m a n i Formation, Lower C a r b o n i f e r o u s , I s t a n b u l : i n Kaya, O . , E d i t o r , Paleozoic of I s t a r b u l : Fge U n i v e r s i t e s i Fen F a k u l t e s r K i t a p l a r S e r e s i No04n, p.117-134. Kling, S.A. and R e i f , W . - E . , 1969. The P a l e o z o i c h i s t o r y o f amnhidisc and hemidisc sponges: new evidence from the Carboniferous of Uruguay: J o u r . P a l e o n t . , v.43, p.1429-1434. Nazarov, B . B . , 1973. F i r s t d i s c o v e r y of R a d i o l a r i a E n t a c t i n i i d a e and C e r a t o i k i s c i d a e i n the Upper Devoqizn of t h e Southern U r a l s : Dokl. Akad. Nauk S.S.S.R., v.210, p.696-699. ( I n R u s s i a n ) . ----_-----_-- 1975. Lower and Middle P a l e o z o i c P a d i o l a r i a n s of Kazakhstan: Trudy Geol. I n s t . Akad. Nauk S.S.S.R., v.275, 202pp. ( I n R u s s i a n ) . -_-_--_______ , Popov, L . E . and Apollonov, N . K . , 1975. R a d i o l a r i a from t h e Lower P a l e o z o i c of Kazakhstan: I z v . Akad. Nauk S.S.S.P. S e r . Geol. f o r 1975, no.10, p.96-111. ( I n R u s s i a n ) . N i g r i n i , C . and N i t e c k i , PI.H., 1968. Occurrence of R a d i o l a r i a i n t h e M i s s i s s i p p i a n of Arkansas: F i e l d i a n a . Geology, v . 1 6 , o.255-268. Ramsbottom, N . H . C . , 1971. Palaeogeography and g o n i a t i t e d i s t r i b u t i o n i n t h e Namurian and E a r l y Westphal i a n : Compte Rendue S i x i h e Cong, I n t e r n a t . d e S t r a t . e t de Gebl. d u Carbonif'ere, v . 4 , p.1394-1399. R i e d e l , W.R., 1967. Class Actinopoda: i n Harland, W . B . e t a l . , E d i t o r s , The F o s s i l Record: London ( G e o l o g i c a T S o c i e t y ) , p.291-298. ------____-_ and Foreman, H . P . , 1961. Type specimens o f North American P a l e o z o i c R a d i o l a r i a : J o u r . P a l e o n t . , v.35, p.628-632. Sturmer, W . , 1966. Das Wachstum s i l u r i s c h e r S p h a e r e l l a r i e n und i h r e s p d t e r e n chemischen Umwandlungen: P 8 l e o n t . Z., v . 4 0 , p.257-261,
180
SPECIl1CI:S FIGURED: L i s t e d below a r e t h e s l i d e numbers of a l l f i g u r e d soecirneiis w i t h Enaland F i n d e r c o - o r d i n a t e s Is1 i d e l a b e l t o r i g b t ) i n b r a c k e t s . In t h e c a s e s - o f S i l u r i a n specimens the'numbers (GSC) a s s i g n e d t o t h e specimens i n t h e National Type C o l l e c t i o n o f I n v e r t e b r a t e F o s s i l s , G e o l o o i c a l Survey of Canada, O t t a w a , a r e a l s o i n d i c a t e d . All specimens a r e p r e s e n t l y s t o r e d i n t h e a u t h o r ' s c o l l e c t i o n . P l a t e 1 . 1 . PFIO/P/24. ( K 4 3 . 4 ) ; 2. PllO/P/15. ( J 4 0 . 4 ) ; 3 . PlV/P/17. ( H 3 7 ) ; 4. C W I . C K / P / 2 . ( K 4 3 . 2 ) . C C C 48235: 5 . CLJI,CB/P/13. ( 5 4 0 . 3 ) . GSC 48236: F . w I . c r / r / i 3 , ( ~ 3 5 . 1 ) .GSC 48237. P l a t e 2. 1 . CWI.CS/F/?. (1'43.2). G S C 48238; 2 . CL
181
Plate 1 . 1-3 Early Ordovician Palaeoactinommids (1. Paleontolocisk Puseum Oslo NF.3280 x262, 2. PI10 liF.3281 x262, 3. P l V l NF.3282 ~ 2 8 5 ) . 4-5 Silurian Palaeoscenidiidae (4. A f f . Palaeoscenidium s p . x262, 5. Palaeoscenidium s . 1 , sp. x285). 6 . Silurian Palaeoactinornmid 2 . xl c
182
183
Discussioii Ur. S. M. Berystroirt: Is t h e r e any morphological f d a t u r e by which e n t a c t i tiiid r a d i o l a r i a n s without s h e l l can be s a f e l y distinguished from sponge spicules? I nave s p i c u l e - l i k e f o s s i l s i ~ m i y c o l l e c t i o n s from t h e upper A r e i t i ~ i d ~ofi westerii Newfoulidland which a r e very s i m i l a r indeed to some of tlie r a d i o l a r i a n s you i l l u s t r a t e d , b u c they lack s h e l l . I am not sure i f cney a r e r a d i o l a r i a n s o r sponge s p i c u l e s and wonder i f you can advise ilie tiow t o c l a s s i f y these forms? Dr. B. K. Holdsworth: I n t n e Lower Ordovician t h e s p i c u l e s of t h e Entacti~tiiclde c t i d t I kliow a r e reillarkduly strong. All rnignt well be mistaken f o r some kind of sponge s k e l e t a l element, b u t f o r t h e f a c t t h a t in many cases t n e rays of tne spicules give r i s e t o a more o r l e s s complete l a t t i c e s n e l l , sorliecimes r e t i c u l a t e . B u t I admit t h a t a real problem e x i s t s . Tne presumed e n t a c t i n i i d , Haplentactinia a r r h i n i a Foreman, i s one of r e l a t i v e l y very few Ldte Paleozoic t n t a c t i n i a c e a lacking a t r u e l a L t i c e , tne s p i c u l e rays being ornamented by c i r c l e t s of s h o r t spinules. Superf i c i a l l y i t does look r a t n e r a s tnougn i t could be a sponge element. On cne o t h e r hand, i t i s q u i t e d i f f e r e n t from any kind of s p i c u l e t h a t I know to nave oeen fourid d e f i n i t e l y forming part o f a Late Paleozoic sponge skeleton. I agree wit11 Helen Foreman tiiat t h i s s p i c u l a r form r e a l l y i s an elitac t i n i id radio1 a r i an. &.it in tile t a r l y Paleozoic I sliare your d i f f i c u l t y . There a r e a l o t of often robust s p i c u l a r f o s s i l s shoding the bar-centre s i m i l a r to t h a t SO ofcen found in unquestionable h t a c t i n i a c e a , b u t with no t r u e l a t t i c e d sliell - only s n o r t spinules. Should we consider these to be haplentactiniidae in tne very Droadest sense o r a r e they completely unrelated? Pdssiuly important clues a r e t h e i r s i m i l a r i t y in s i z e and s p i c d l e strength t o unquestiondule Early Paleozoic k n t a c t i n i i d a e ; t h e broadly continuous spectrdili e x i s t i n g wtween t r u l y l a t t i c e d and completely u n l a t t i c e d forms, dnd tne coimon f a i l u r e of obvious sponge megascleres t o appear abundantly in t n e same rocks. I confess, nowever, t o considerable ignorance regardiny t d r l y t'alsozoic s+onges, and I r e a l l y cannot say a t present t h a t I know of an/ i n f a l l i b l e t e s t wliicn can d i f f e r m t i a t e between an u n l a t t i c e d t d r l y Paleozoic e i i t a c t i n i i d aiid a l l kinds of sponge spicules. I d o n ' t know witether Helen has any views on your Problem. Or. H. V. Foreman: No, I r e a l l y have tiothing to add. Dr. F. Grdustein: You s t a t e d chat you hope t h a t Paleozoic Kadiolaria may be of use a s i n d i c a t o r s of ' b l u e ' , deep oceanic water masses. Do you think t n a t i t i s clepin r a t n e r than d i s t a n c e from a shore which you hope t o de1 i n e a w , o r boch? Would the occurrence of Radiolaria-rich d e p o s i t s not ue rattier due t o a "sediment starved" s i t u a t i o n witn l i t t l e o r no d e t r i t a l sl?iliaient iiifldx instead of tlte presence of deep oceanic condi Lions? Holdbworcti: I do not think triat my supposed ' b l u e ' water assemblages necess a r i l y iricricate e i t h e r p a r t i c u l a r l y deep water sediments o r p a r t i c u l a r l y la~iu-reinotesedirnents. I n t i i t ! c i t e d cases of southern France and northd e s t Turkey I incended to suggest t n a i ttie p e s e n c e of unusually d i v e r s e Edrly Larboniferous Hadiolaria asserliulages might be due t o an oceanic infldence induced by subsidence of French and Turkish shallow water, cratonic platforms immediately bordering a deep, 'Tethyan' ocean. To what depth t h a t depositional s u r f a c e would have needed t o have been submerged t o allow ingress of l i t t l e - m o d i f i e d open ocean assemblages, and t h e d i s tance of t h e depositional s i t e from contemporaneous s h o r e l i n e s a f t e r such submergence I would not l i k e t o say. What I do suggest i s t h a t t h e i n fluence on Kadiolaria populations of such ' o v e r s p i l l ' of open ocean water i s l i k e l y t o have declined rapidly in t h e d i r e c t i o n of t h e continent i n t e r i o r s . Certainly, sediments whicn accumulated well within major Late Paleozoic oceans, presumably a t very considerable depth and d i s t a n c e from siiore, should e x h i b i t s i m i l a r , high-diversity assemblages. B u t t h e s t r a -
-
184
t i g r a p n i c contexts of trie French and Turkish occurrences do n o t , I think, i n d i c a t e convincingly t n a t they f a l l i n t o t h i s category, and a s y e t I know with c e r t a i n t y of no occurrences which might. Sediment s t a r v a t i o n in both deep and shallow water will often increase t h e oulk of iiadiolaria in u n i t volume of rock. I t alone i s unlikely to markedly increase d i v e r s i t y - and I s t r e s s t h a t i t i s marked d i f f e r e n c e in d i v e r s i t y of assemblages in Paleozoic rocks with l a r g e Radiolaria f r a c t i o n s t h a t I believe t o be of importance in paleooceanography. I did not mean to imply t h a t simple abundance of Radiolaria i n Paleozoic sediments i s an i n d i c a t i o n of oceanic influence o r of deep water o r i g i n . On the contrary, I am sure t h a t some Paleozoic Radiolaria could occur profdsely in d i s t i n c t l y shallow epicontinental seas f a r removed from t h e deep oceans, and given c e r t a i n conditions - including slow sediment accumulat i o n - they were preserved in considerable numbers. Dr. W . Manspeizer: What independent c r i t e r i a a r e a v a i l a b l e t o support your inference triat the Late Paleozoic Radiolaria a r e ' b l u e ' water fauna? Holdsworth: Tile 'independent c r i t e r i a ' I mentioned do not r e l a t e t o my i n ference t h a t extremely d i v e r s e Late Paleozoic assemblages with a d i v e r s e pilentonemid element a r e ' b l u e ' water faunas. The ' c r i t e r i a ' r e l a t e to iiiy suspicion t n a t in q u i t e shallow epicontinental s e a s , r e l a t i v e l y f a r removed froili deep oceans, comparatively s l i g h t increase in water depth tended to r e s u l t in increase i n Radiolaria d i v e r s i t y w h i l s t not permitting the extreme d i v e r s i t y of the supposed ' b l u e ' water assemblages. I n t h e ' o a s i n f a c i e s ' of tfle European Namurian i t i s common t o f i n d the predominant goniatite-pelecypod macrofauna confined t o r e l a t i v e l y t h i n oeds of very f i n e black shale o r mudstone, a few inches t o a few f e e t t h i c k , separated by very v a r i a b l e thicknesses of u n f o s s i l i f e r o u s s t r a t a . Tile f o s s i l i f e r o u s beds a r e t h e so-called 'marine b a n d s ' , and i t i s exclus i v e l y witnin ttiese bands t h a t hadiolaria a r e known t o occur. The conventional explanation of 'marine bands' i s t h a t t h e rock record r e f l e c t s successive r i s e s and f a l l s of sea l e v e l , probably e u s t a t i c , with 'marine bands' rriarking e u s t a t i c ' h i g h s ' . Falling sea level might be expected t o nave increased basin margin erosion, r e s u l t i n g in t h e i n t e r p o l a t i o n of sands and s i l t s oetween 'marine bands'. Often such i n t e r p o l a t i o n s occur. B u t a t the southern end of t h e Central England Basin i t i s noticeable t h a t c e r t a i n parts of the succession display 'groups' were periods during which average sea level i n the southern area was s u f f i c i e n t l y high t o prevent s i g n i f i c a n t local erosion even during e u s t a t i c ' l o w s ' . (Enhanced local subsidence, higher average e u s t a t i c sea level o r a combination of these f a c t o r s could be invoked in explanation.) Concomitantly, e u s t a t i c ' h i g h s ' uri ngi ng in t n e macrofauna during ttiesz periods should have r e s u l t e d in local r e l a t i v e l y deep s e a s . I n f a c t , presently known occurrences of A l b a i l l e l l a r i a , Ceratoikiscidae, Popofskyellidae and Corythoecia spp. a r e confined t o 'marine bands' of those 'groups' which lack i n t e r p o l a t e d sands and s i l t s . Such bands tend t o have r a t h e r low g o n i a t i t e and sponge cont e n t s - furtiier i n d i c a t o r s , i t can be argued, of r e l a t i v e l y deep water conditions. B u t the Pylentonemidae a r e v i r t u a l l y absent from all 'marine bands I .
185
Autunian and C a r n i a n P a l y n o f l o r u l e s C o n t r i b u t i o n t o t h e Chronology and T e c t o n i c H i s t o r y o f t h e Moroccan P r e - A t l a n t i c B o r d e r l a n d H a r o l d L. Cousminer The American Museum of N a t u r a l H i s t o r y New York, New York Warren l l a n s p e i z e r Newark C o l l e g e o f A r t s and S c i e n c e s Rutgers University Newark, New J e r s e y Abstract
A p a l y n o f l o r u l e from t h e Moroccan Meseta n o r t h o f Khenifra c l o s e l y matches f o s s i l p o l l e n a s s e m b l a g e s from t h e European Autunian Series, as d e s c r i b e d by Daubinger (1974) and from t h e P i c t o u Group o f Nova S c o t i a ( B a r s s and H a q u e b a r d , l 9 6 7 ) . The age o f t h e s e a s s e m b l a g e s i s l a t e s t C a r b o n i f e r o u s t o e a r l i e s t Permian. A second p a l y n o f l o r u l e from t h e C e n t r a l High A t l a s s o u t h of Marrakech i s o f mid-Carnian a g e b a s e d upon t h e c o n c u r r e n c e of a g e - d i a g n o s t i c p o l l e n s p e c i e s t h a t have been r e p o r t e d from t h e Swiss and E n g l i s h m i d d l e Keuper, t y p e C a r n i a n o f A u s t r i a , and North American T r i a s s i c b e d s i n V i r g i n i a , North C a r o l i n a , P e n n s y l v a n i a , New J e r s e y , Texas, New Mexico, and A r i z o n a .
These a g e d e t e r m i n a t i o n s serve t o document a l a r g e - s c a l e u n c o n f o r m i t y t h a t encompasses v i r t u a l l y a l l t h e Permo-Triass i c on t h e Moroccan Meseta and High Atlas. T h i s e p i s o d e of pronounced c r u s t a l t h i n n i n g p r e c e d e d T r i a s s i c r i f t i n g , and u l t i m a t e l y l e d t o t h e f r a g m e n t a t i o n o f Pangaea. Introduction Late P a l e o z o i c and E a r l y Mesozoic r e c o n s t r u c t i o n s of
Pangaea a r e i n f e r r e d l a r g e l y on p a l e o m a g n e t i c , g e o p h y s i c a l and l i t h o l o g i c d a t a ( s e e Pitman and Talwani, 1972; P h i l l i p s and F o r s y t h e , 1972; B a l l a r d and Uchupi, 1975; Schenk, 1971; O l s e n and Leydon, 1973; Dewey, Pitman and Ryan, 1973; and many o t h e r s ) .
Paleontologic data bearing
186
on t h i s q u e s t i o n and t h e t i m e of opening of t h e A t l a n t i c , however, are s p a r s e , This paper p r e s e n t s new p a l y n o l o g i c data from t h e Late Carboniferous and Late T r i a s s i c of Morocco t h a t b e a r on t h e sequence of e v e n t s l e a d i n g up t o t h e opening of t h e North A t l a n t i c Ocean. Geologic S e t t i n g Late Carboniferous sediments of Morocco were depo-
s i t e d i n small i s o l a t e d b a s i n s on t h e Moroccan-Oranian
Meseta, and i n l a r g e P a l e o z o i c t r o u g h s of t h e Tindouf and Colomb Bechar Basins (Fig. 1). According t o Michard and Sougy (1974, c i t e d i n Van Houten, 1976), t h e Meseta w a s formed by t h e Late P a l e o z o i c Hercynian (Variscan) Orogeny. The main d e f o r m a t i o n a l phases o c c u r r e d from t h e Namurian t o t h e end of t h e Westphalian, and i n c r e a s e d i n i n t e n s i t y towards t h e margin of t h e A t l a n t i c Ocean. A new e x t e n s i o n a l phase of c r u s t a l deformation oc-
c u r r e d i n t h e E a r l y Mesozoic t h a t fragmented t h e Meseta. Late Triassic c o n t i n e n t a l sediments of Middle Carnian and younger a g e were l a i d down i n grabens i n t h e w e s t e r n High A t l a s as t h e Meseta began t o s e p a r a t e from t h e A f r i c a n
P l a t e (Cousminer and Manspeizer, 1976).
In the Early
J u r a s s i c (Sinemurian), as North America and A f r i c a began t o move a p a r t , t h e Oranian and Moroccan P l a t e s s e p a r a t e d a l o n g t h e l i n e of t h e Middle A t l a s Mountains (Manspeizer, P u f f e r , and Cousminer, 1976b).
E a r l y J u r a s s i c seas, ema-
n a t i n g from t h e Tethys Sea t o t h e n o r t h and e a s t , t r a n s g r e s s e d t h e Meseta a l o n g t h e High Atlas and Middle A t l a s troughs.
/
TINDOUF
RIF
16
,
BASIN
F i g . 1 . Permo-Carboniferous s e d i m e n t s and p r e s e n t - d a y s t r u c t u r a l p r o v i n c e s . 1 = I d a o u Z a l ; 2 = A l m e z i ; 3 = H a d a d j a ; 4 = Ilechra ben Abbou; 5 = Nzala A r r a r c h a ; 6 = S i d i Kassem; 7 = Chougrane; 8 = K h e n i f r a ; 9 = T e l o u e t ; 1 0 = A i t Z i f f a ; 11 = C h i c h a o u a ; 1 2 = Ben A c h o u c h ; 1 3 = T a z z e k a ; 14 = N a s g o u t ; 15 = J e r a d a ; 16 = T r a r a s .
188
Autunian Palynoflorule:
Moroccan Meseta, Khenifra Basin
A diversified palynoflorule was recently recovered from samples collected by John Lorenz in 1974 from a conglomeratic sandstone on the Moroccan Meseta, about 3 . 5 km. north of the city of Khenifra and along the west bank of the Oued O m er Rbia.
The productive samples were col-
lected about 100 meters above the base of a conglomeratic sandstone that is about 5 0 0 meters thick, folded, and unconformably overlying Namurian (Mississippian to Pennsylvanian) metamorphic rocks.
According to Lorenz (1974),
the formation is principal1.y composed of sandstones intercalated with massively bedded conglomerates, containing poorly sorted, well rounded cobbles of vein quartz, quartzite, and metamorphic rock in a sandy and pebbly matrix. Palynomorphs recovered from the samples are listed on Figure 2, and fall into four groups: I.
Lower vascular plant spores: 11. monosaccate
pollen: 111. disaccate pollen: IV. miscellaneous gymnospermous pollen. Because of the occurrence in the Moroccan samples of Pictou Group zonal markers from both the P o t o n i e i s p o r i t e s and V i t t a t i n a zones, a stratigraphic level is indicated equivalent to that near the boundary between these two zones.
This interpretation signifies that the sample is
of uppermost Stephanian to Lower Permian age. Based on this work a direct comparison can be made between the palynoflorules from the type Autunian sediments of Europe and the Khenifra Basin. In Figure 3 are histograms
showing the percentages of spores, monosac-
cates, disaccates, monocolpates and V i t t a t i n a species
*
* *
*
* * *
ENDOSPORITCS LAEVIGATOSPORITES SCHOPFIPOLLENITES APICULATISPnRIS CYCLOGFANISPORITES LOPHOTRILETES PUNCTATISPORITFS Lycnspnm CALAMOS PDRA RETUSOTRILETES CONVOLUTI SPORA LCIOTRILETES KNOXISPORITES X'TICJLATISPORITES LUNDRLADISPORA VESTISPORA
SPORES
MONOSACCATES
POTONIEISP3RITES MONOSACCITES INI1I:T. FLORINITES CORDAIT I N A CY CADOP I T E S MARS UP I P O L L E N I T E S VITTTATINA VESICASPORA L I M I T IS P O R I T ES SULCATISPORITES ILLINITES S T R I OMONOSACC I T E S PROTOHAPLOXY P I N U S STRIATOABIETITES VESTIGISPORITES DISACCITES INDET. HAMIAPOLLENITES STRIATITES FALC I S P O R I T E S
* Fig.
R
ZONAL MARKERS I N P I C T O U GROUP,
MISC
GYMNOSPERMS
DISACCATES
0
1
5 I
1c
I5
I
FR E Q U E N C Y NOV.4 S C O T I A
2 . F r e q u e n c y h i s t o g r a m o f p a l y n o m o r p h s fro'm t h e M o r r o c a n M e s e t a .
20
25
F i ? . 3 . H i s t o g r a m s o f p o l l e n f r o m E u r o p e a n a n d Moroccan A u t u n i a n s e d i m e n t s (European d a t a from Doubinger, 1 9 7 4 ) .
D
c
I. 8
9. 8
m
+
MISCELLANEOUS
HONOSACCATES
MONOCOLPATES
1. 7
2
1 5 DISACCATES
2 . 6 MONOSACCATES
?6. 7 TRILETES
3
2.
<
6
rn
4 .t
z
L
D
,
D Z
D z I
I
VX)
--a2
zm-
gz:
O r
2 1 DISACCATES
A L L SPORES
< 1 VITTATINA
14
1 VITTATINA
14 D I S A C C A T E S
5 VITTATINA
12 MONOCOLPATES
5 DISACCATES
7 1 MONOSACCATES
A L L SPORES
4 2 MONOSACCATES
DISACCATES
1 VITTATINA
2 2
3 2 A L L Sr'ORES
w
0 ZZ m
N
2 3 VITTATLNA
VITTATINA
A L L SPORES
18
3 HISCELLANEOUS
7
I I 5 DISACCATES
58 HONOSACCATES
1 1 A;LlspOREs
191
characteristic of Autunian 1 (lower series), Autunian 2 (intermediate series) and
Autunian 3 (upper series) sedi-
ments from the type Autunian deposits of Europe, (from data tabulated by Daubinger, 1974).
Included for compari-
son is a similar histogram of the Khenifra palynoflorule. The Morroccan sample is clearly intermediate in character between the Autunian 1 (Stephanian or Late Carboniferous) and the Autunian 2 (Lower Permian) histograms.
Thus, on
the bases of comparisons with both the Pictou Group pollen sequence (Barss and Hacquebard, 1967) and the type Autunian of Central Europe (Daubinger, 1974) the Moroccan palynoflorule from Khenifra is of latest Carboniferous to earliest Permian age (Fig. 4).
A Triassic Palynoflorule from the Central High Atlas Recently (Cousminer and Manspeizer, 1976a) we reported on the biostratigraphic significance of fossil pollen recovered from samples of the Oukaimeden Sandstone, collected in the Ourika Valley south of Marrekech.
According to
Mattis (1975), the Oukaimeden sandstone is about 200 meters thick, and is composed of fine-grained, well-sorted, and well-rounded quartz sands that are cross-bedded, ripplemarked and lensoid.
The productive samples occur about 400
meters above the base of a clastic sequence that overlies Precambrian rock, and about 200 meters below the High Atlas basalts which have an average radiometric age of about 195 m.y.
The base of the Triassic section, in the northern
part of the Ourika Valley, is compsed of an andesitic lava flow and volcanic clasts that rest unconformably (Choubert and Faure-Muret, 1962) on fossil-bearing red beds con-
192
Z x
SINEMURIAN
5
HETTANGIAN
5
KUNGURIAN
1
l
Fig.
4. C o r r e l a t i o n c h a r t .
/
1
1
193
sidered of Stephanian age because they contain W a l c h i a l i n e a r i f o l i a , Mixoneura n e u r o p t e r o i d e s (Clariond and Leca, 19341, and W a l c h i a p i n i f o r m i s (Greber and Proust, 1958). Figure 6 is modified from our previous publication (Cousminer and Manspeizer, 1976a) and gives all cited stratigraphic and geographic occurrences of age-diagnostic palynomorphs, recovered from the Central High Atlas of Morocco.
They include the following comparatively long-
ranging forms that are either entirely restricted to Triassic sediments or have their maximum distribution in sediments of Triassic age: Aratrisporites species, C h o r d a s p o r i tes species , - T r i a d i s p o r a species, O v a l i p o l l i s species, P o r c e l l i s p o r a longdonensis, E n z o n a l a s p o r i t e s v i g e n s
.
The Moroccan sample from the Central High Atlas
is dated as of mid-Carnian age by the concurrent range of three fossil pollen species (Groups 1 and 5 on the Range Chart) that have overlapping and ptrtially concurrent ranges in several European localities.
Conclusions
1) Two palynoflorule datums have been established in Morocco.
These date the youngest Paleozoic rocks on the
Meseta (Khenifra Basin) as of Autunian age (StephanianSakmarian); and the Triassic rocks on the High Atlas (Ourika Valley) as of mid-Carnian age.
Based on the palynoflo-
rules these Moroccan strata are correlated with beds of equivalent age in many localities in both North America and Europe.
194
Fi0.5.
Stephanian - Autunian depositional domaines and paleogeography,
195
2 ) The Autunian palynoflorule from the Moroccan Meseta
closely resembles those from the upper members of the Pictou Group in Nova Scotia, as described by Barss and Hacquebard ( 1 9 6 7 ) .
This gives additional support to the
Bullard reconstruction of Pangaea (Bullard et al, 1 9 6 5 ) which places Nova Scotia opposite the Moroccan Meseta during pre-drift time. 3 ) The pollen data document a major unconformity in
Morocco that extends from:
(a) the Autunian-Sakmarian to
the overlying Sinemurian on the Moroccan Meseta; and (b) from the Stephanian-Autunian to the middle Carnian of the High Atlas. The hiatus indicates that marked uplift and erosion of this region may have been dominant processes for for over 75 million years.
This episode of extensive cru-
stal thinning is considered to be of great significance in that it led to isostatic uplift, rifting, volcanism and the ultimate fragmentation of Pangaea. 4) The great increase in gymnospermous pollen near
the Permo-Carboniferous boundary, which to a great extent replaced the dominant lower vascular plant spore assemblages so characteristic of the Carboniferous swamp lowlands, is connected with climatic changes that profoundly affected the distribution of Late Paleozoic plant communities.
Since paleomagnetic data indicate that the posi-
tion of the equator remained fairly stable through the Permo-Carboniferous interval, this climatic change is primarily correlated with orographic uplift. 5) Palynofloristic, paleobotanical, and lithostrati-
graphic records indicate that the degree of uplift and its duration in the Late Paleozoic-Triassic interval varied markedly at different geographic localities.
Rapid change
upwards from typically Carboniferous spore assemblages to
Fig.6. C i t e d s t r a t i g r a p h i c and g e o g r a p h i c o c c u r r e n c e s of a g e - d i a g n o s t i c C e n t r a l H i g h A t l a s o f Mo r o cco ( m o d i f i e d f r o m S c i e n c e Volume 1 9 1 , palynomorphs p.944). from t h e
,
ANlSlAhi
HUSCHELKALK
I
1
I
I
MII)I)LE
LADlNlAN
1
LOWER LOWER
I
1 -
NORIAN
- A RATRISE'ORITES
- -SPECIES --
A L l S P O b l l a S MINUTlSACCUS POTONlEl .CCUS .. ..-. ....
CARNIAN
MIDDLE
UPPER
UPPtR K E U P E R
T R I A S S I C
~
I
1 RHAET
ALPISI-
t N G L A N D 1131 S W I T Z E R L A N D 1101 E N G L A N D 1121 F W I T Z E R L A N D 19)
4 {
E N G L A N D 112, 131
ARIZONA
ENGLAPID (131
ENGLAND ( 1 3 ) S W I T Z E R L A N D (10)
ALGbRIA.TUhlSIA 12?1 NEW J E R S E Y 1 2 1 1
ARIZOYA (141
E N G L A N D ( 1 2 , 13) N O R T H S E A (81 AUSTRIA 1111 S W I T Z E R L A N D 19 10, F R A N C E 119)
{
1
I
. .-
bPIGLAND ( 1 2 1
NLW J t R S E Y (21) E N G L A N D (131 WORT11 S t A 18) AUSTRIA ( I l l S W l T Z t R L A N D 110)
ENGLAND I131 AUSTRIA ( II I S W I T Z E R L A h D 1'1. 101
E N G L A N D (131 S W I T Z E R L A N D I Y 101 AUSTRIA (REIFFLINGER KALKlj V A ,Y O R T H C A R O L I N A T E X . h MEXICO ARIZONA I NEW J E R S E Y ( 2 1 )
{ S W I T Z E R L A N D 110)
<
J
1
{ S W I T Z E R L A N D I101
1
TEX N MEXICO
ALGERIA-TUYISIA (221 VA , N O R T H C A R O L I N A
LNGLAND ( 1 2 . 131 N O R T H SLA 18)
VEW J E R S E Y 1211
1
~
A U S T R I A (II I N O R T H SEA 181
(11)
1
CITATIONS E N G L A N D 1121 C E R M A h Y I201 S W l T Z E R L A h V 1101 E N G L A N D IIII T E X A S 1161
I
1
LVR
IIIIOUP
StRItS
SYSTI 4 '
197
dominantly gymnospermous saccate pollen associations occurs in both the type Autunian Basin of France and the Pictou Group sediments of Nova Scotia.
Dunkard Group sedi-
ments of West Virginia are presumed to be of equivalent age based on paleobotanical and paleontological evidence; however, spore assemblages of Carboniferous aspect remain dominant throughout the Dunkard sequence, indicating the persistence of a marginal lowland basin in this region extending into the Lower Permian.
Re ferences Ballard, R.D. and Uchupi, E., 1975. Carboniferous and Triassic rifting: a preliminary outline of the tectonic history of the Gulf of Maine. Geol. SOC. America, Bull., 83: 2285-2302. Barlow, J.A., (editor) 1975. Proceedings of the First I. C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey, 352pp. Barss, M . S . , 1967. Carboniferous and Permian spores of Canada. Geol. Surv. Can. Pap. 67-11 , 17pp. Barss, M . S . , and Hacquebard,P.A., 1967. Age and the stratigraphy of the Pictou Group in the Maritime Provinces as revealed by fossil spores. Geol. Assoc. Can., Spec. Paper 4: 267-282. Barss, M.S., 1972. A problem in Pennsylvanian-Permian palynology of Yukon Territory. Geoscience and Man, 14: 67-71. Bode,H.H., 1975. The stratigraphic position of the Dunkard; In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey, pp. 143-154. Brown, R.H., 1974. The Argana Basin: A Triassic model for early rifting. M . S . thesis, Univ. of S. Carolina; Columbia, S.C. Bullard, E.C., Everett, J.C. and Smith, A.G., 1965. The fit of the continents around the Atlantic; In: A symposium on continental drift. Royal SOC. London Phil. Trans. Sec. A; 258: 41-51.
198
Cavaroc, V.V., Padgett, G., Stephens, D.G., Kanes, W.H., Boudda-Ahmed and Wollen, I.D., 1976. Late Paleozoic of the Tindouf Basin-North Africa. Jour. Sed. Pet. 46(1): 77-88. Chamot, G.A., 1965. Permian section at Apillapampa, Bolivia and its fossil content. Jour. Paleontology 39 (6), 1112-1124. Choubert, G. , and Faure-Muret, A., 1962. E(vo1ution du domaine Atlasique-Marocain depuis les temps Paleozoiques. In: Durand-Delga, M. ed. Livre la mgmoire du Prof. Paul Fallot: SOC. GQol. France, Mdm. 1: 447-527. Clariond, L. and Leca, F., 1934. 6tudes sur le Stephanien du versant nord de 1'Atlas de Marrakech: SOC. Ge/ol. France, Comptes Rendus Sommaire 10: 210-212. Clarke, R.F.A., 1965. Keuper miospores from Worcestershire, England. Palaeont. 8: 294-321. Clendening, J.A., 1965. Palynological evidence for a Pennsylvanian age assignment of the Dunkard Group in the Appalachian Basin. Part I. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey, pp. 195-221. Part 11. Coal Geology Bulletin 3. West Virginia Geological Survey. Cornet, B., Traverse, A., and McDonald, N.G., 1973. Fossil spores, pollen, and fishes from Connecticut indicate Early Jurassic Age for part of the Newark Group. Science 21: 1243-1247. Cornet, B., and Traverse, A., 1975. Palynological Contributions to the chronology and stratigraphy of the Hartford Basin in Connecticut and Massachusetts. Geoscience and Man 11: 1-33. Cousminer, H.L., 1965. Permian Spores from Apillapampa, Bolivia. Jour. Paleontology 39(6), 1097-1111. Cousminer, H.L. and Manspeizer, W., 1974. Late Triassic palynoflorules form Morocco: Comparison with eastern North America. 1974 Annual G.S.A. meeting, Miami Beach, Florida, p. 697. , and , 1976a. Triassic pollen date the incipient rifting o f Pangea as Middle Carnian. Science 191: 943-945. , and , 197613. Autunian and Carnian palynoflorules: Contribution to the chronology and tectonic history of the Moroccan Pre-Atlantic Borderland. In: Swain, F.M. (editor), Symposium on StratigraphTc Micropaleonto3ogy of Atlantic Basin and Borderlands, Proceedings Volume (in press) , Elsevier Scientific Publishing Company.
199
Cross, A.T., 1975. The Dunkard in perspective: Geology, sedimentation and life. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey, pp. 297-299. Darrah, W.C., 1975. Historical aspects of the Permian flora of Fontaine and White, Ibid, pp. 81-101. Daubinger , J. , 1974. Ztudes palynologiques dans 1 'Autunien. Rev. Palaeobot. Palynol., 17(1/2); 21-38. Defretin, S. and Fauvelet, E., 1951. Prgsence de phyllopodes triassiques dans la re'gion d'Argana-Bigoudine (Haut Atlas Occidental). Notes it M6m. Serv. Ge/ol. Maroc 5 (85), 129-134. Dewey, J.F., Pitman, W.C., Ryan, W.B.F., and Connin, J., 1973. Plate tectonics and the evolution of the Alpine system. Geol. SOC. Amer. Bull. 84: 3137-3180. Dunay, R.E., and Fisher, M.J., 1974. Late Triassic palynoflorules of North America and their European correlatives. Rev. Palaeobot. Palynol., 17(1/2): 179-186. Durden, C.J., 1975. Age of the Dunkard: Evidence of the insect fauna. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey, p. 295. Dutuit, J.M., 1964. Dkcouverte de gisements fossiliferes dans le Trias du couloir d'Argana (Atlas-occidental Marocain). Comptes Rendus Acad. Sci. Paris 285(4): 1285-1287. Eager, R.M.C., 1975. Some nonmarine bivalve faunas from the Dunkard Group and underlying measures. In: J.A. Barlow (editor). Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey. pp. 23-67. Feys, R., and Greber, C., 1963. Stephanien et 1'Autunien du Souss dans les Ida ou Zal (Haut Atlas OccidentalMaroc); Notes et M6m. Serv. Geol. Maroc 22(170): 19-35. Fisher, M.J., 1972. The Triassic palynofloral succession in England. Geoscience and Man 4: 101-109. Geologic Map of Morocco , 1954-56. Protectorate Rgp. Fr. Maroc; Div. Mines, Service Ggologique. Gillespie, W.H., Hennen, G.J., and Belasco, C., 1965. Plant megafossils from Dunkard strata in northwestern West Virginia, and southwestern Pennsylvania. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard'l. West Virtinia Geological and Economic Survey, pp. 223-248.
200
Greber, C., and Proust, F., 1958. Sur le Permien et le Trias dans le Haut Atlas Occidental. SOC. G'eol. France, Comptes Rendus Somaires, 1 0 : 210-212. Havlena, V., 1965. European Upper Paleozoic C a l l i p t e r i s c o n f e r t a , and the Permo-Carboniferous Boundary. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard" . West Virginia Geological and Economic Survey, pp. 7-22. Jansa, L.F., and Wade, J.A., 1975. Geology of the continental margin off Nova Scotia and Newfoundland, In: Offshore geology of eastern Canada. Geol. Surv. Can. paper 74-30, 2. Leschik, G., 1955. Die Keuperflora von Neuwelt bei Basel. Schweiz. Pal'ciont. Abh. (SeparatdrGcke) 72: 1-68 pp. Lorenz, J., 1974. Triassic sediments and basin structure of the Kerrouchen Basin, central Morocco. M.S. thesis, Univ. S . Carolina, Columbia, S . Carolina. Lund, R., 1975. Vertebrate fossil zonation and correlation of the Dunkard Basin. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey, pp. 171-182. Maedler, K., 1964. Die geologisch Verbreitung von Sporen und Pollen in der deutschen Trias-Geol. Jahrb., Beih., 65: 147 pp. Manspeizer, W., Puffer, J., and Cousminer, H.L., 1975. The Triassic record, a view from northwest Africa (Morocco). Symposium on Triassic Stratigraphy, Wesleyan College, Connecticut. (Oral Presentation only). I , and , 1976. Subduction, rifting, and sea floor spreading, Eastern Section G.S.A. meeting, Arlington, Virginia, March. I , and , 1976b. Opening of the North Atlantic Ocean: A Triassic-Jurassic record in Morocco and North America. Section of geological sciences, The New York Academy of Sciences, Monday, April 5, (Oral Presentation only). Mattis, A., 1975. Nonmarine Triassic sedimentation, central High Atlas Mountains, Morocco. Ph.D. dissertation, Rutgers University, New Brunswick, New Jersey. Olsen, W., and Leyden, R.J., 1973. North Atlantic rifting in relation to Permian and Triassic systems and their mutual boundary. Lamont-Daugherty Geol. Observ. Contr. #1810, pp. 720-732.
201
Olson, E.C., 1975. Vertebrates and the biostratigraphic position of the Dunkard. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey, pp. 155-165. Phillips, J.D., and Forsyth, D.W., 1972. Plate tectonics, paleomagnetism, and the opening of the Atlantic. Geol. SOC. America Bull. 83: 1579-1600. Pitman, W.C. 111, and Talwani, M., 1972. Sea €loor spreading in the North Atlantic. Geol SOC. America Bull. 83: 619-746. Remy, W., 1965. The floral changes at the CarboniferousPermian Boundary in Europe and North America. In: J. A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey. pp. 305-352. Robb, J.M., 1971. Structure of the continental margin between Cape Rhir and Cape Sim, Morocco, Northwest Africa. Am. Assoc. Petr. Geol. Bull. 5: 643-650. Schenk, P.E., 1971. Southeastern Atlantic Canada, northwestern Africa, and continental drift. Canadian Journ. Earth Sci., 8: 1218-1251. Scheuring, B., 1970. Palynologische und palynostratigraphische Untersuchungen des Keupers in Bolchen Tunnel (Solothurner Jura). Schweiz. Palaeont. Abh. 88: 1-119. Schultz, G., and Hope, R.C., 1973. Late Triassic flora from the Deep River Basin, North Carolina. Palaeontographica B. 141: 63-88. Sohn, I.G., 1975. Dunkard Ostracoda-An evaluation. In: J.A. Barlow (editor), Proceedings of the First West Virginia Geological and Economic Survey. pp. 265-278. Tasch, P., 1975. Dunkard estherids as environmental and age indicators. Ibid pp. 281-292. Van Houten, F.B., 1975. A precocious Atlantic reconstruction. Geology 3: 194-195. , 1976. Late Variscan nonmarine deposits, Northwestern Africa: Implications for pre-drift North Atlantic reconstructions. American Journal of Science, 276: 671-693. Van de Poll, H.W., 1973. Stratigraphy, sediment dispersal, and facies analysis of the Pennsylvanian Pictou Group in New Brunswick. Maritime Sediments, 9: 72-77. Yochelson, E.L., 1975. Monongahela and Dunkard nonmarine gastropods. In: J.A. Barlow (editor), Proceedings of the First I.C. White Symposium-"The Age of the Dunkard". West Virginia Geological and Economic Survey. pp. 249-263.
202
Acknowledgements This research has been supported by a National Science Foundation Grant (GF325104) comprising a special foreign currency grant from the Office of International Programs, and from the Earth Science Section of the Division of Environmental Science. This grant is administered by the University of South Carolina. The senior author acknowledges the use of office and laboratory facilities at York College of the City University of New York, where this research was begun. Thanks are also given to Professor Daniel Habib of Queens College, CUNY, for reviewing the manuscript.
203
Discussion Dr.
B . K . Holdsworth: Has your new work thrown any more l i g h t on t h e d a t i n g of t h e l a t e C a r b o n i f e r o u s - e a r l y Permian r o c k s a t h i g h e r l a t i t u d e s ? Can you s a y a n y t h i n g more a b o u t t h e p o s i t i o n of t h e Permo-Carboniferous boundary i n t h e Gondwana F o r m a t i o n s , f o r i n s t a n c e ?
Cousminer:
Concerning t h a t a r e a , I c a n ' t t h i n k of any d a t a .
Holdsworth: I t ' s s i m p l y t h a t I have been t r y i n g t o f i n d o u t the t r u e a g e of t h e E o a s i a n i t e s ( g o n i a t i t e ) l e v e l i n t h e I t a r a r e F o r m a t i o n of Uruguay which seems t o compare q u i t e n i c e l y w i t h p a r t of t h e S o u t h A f r i c a n s u c c e s s i o n . Some p e o p l e c o n s i d e r t h i s l e v e l i n t h e Gondwana a s l a t e Carboni f e r o u s , o t h e r s t a k e i t a s e a r l y Permian. Cousminer: A r e t h e r e no t e r r e s t r i a l b e d s a s s o c i a t e d ? d i d you s a y t h e s e a r e ? Holdsworth: T h e s e a r e i n t e r g l a c i a l s . South A f r i c a . Cousminer:
What
I t ' s t h e Dwyka i n
T h e r e i s some p o l l e n d a t a on t h e Dwyka S e r i e s .
Holdsworth: T h e r e i s i n d e e d , and I ' m s i m p l y wondering how you would a s s e s s t h i s ? Cousminer: Ill11 t a k e a l o o k a t t h a t . something.
You p u t u s o n t o
Holdsworth: W e l l , i t may b e t h a t t h e r e Ss s i m p l y no c o n n e c t ion a t a l l . I t ' s o b v i o u s l y much h i g h e r l a t i t u d e t h a n t h e s t u f f you a r e t a l k i n g a b o u t . Cousminer:
That's right.
I . G . Sohn: I assume t h a t you a t t r i b u t e t h e d i s t r i b u t i o n o f t h e s e t o wind p a t t e r n s . Is t h a t c o r r e c t ?
Dr.
Cousminer: I n p a r t , y e s . Sohn: W e l l , what c o n f u s e s m e i s t h a t you have a n o r t h - s o u t h d i s t r i b u t i o n no matter how you t u r n i t , which would s o r t of c o n t r a d i c t a wind d i s t r i b u t i o n f o r a l l t h e a r e a s you a r e t r y i n g t o c o r r e l a t e , o r am I wrong? Cousminer: T h e r e c o u l d have been a s e a s o n a l change i n wind d i r e c t i o n , d u e t o a l t e r n a t i o n of t h e t h e r m o c l i n e . Sohn:
Yes,
but f o r t h a t long a distance?
Cousminer: A l l I c a n d o i s p r e s e n t my d a t a . The o t h e r t h i n g i s , of c o u r s e , t h e s e c o n t i n e n t s were much c l o s e r t o g e t h e r t h a n t h e y a r e now. The Dunkard Group was v e r y c l o s e t o t h e Moroccan S a l i e n t and may have been c l o s e r t o a B u l l a r d f i t i f you d i s c o u n t t h e c o n t i n e n t a l s h e l f which may b e o f p o s t T r i a s s i c age.
204
D . Habib: Based on t h e r e c o n s t r u c t i o n of c o n t i n e n t s , your p a l y n o f l o r a l a s s e m b l a g e s were p r o b a b l y w i t h i n a n e s s e n t i a l l y s i n g l e phytogeographic zone. I f t h i s i s t h e c a s e , t h e n you d o n ' t have t o r e s o r t t o l o n g d i s t a n c e wind transportation.
Dr.
Cousrniner: I t h i n k t h a t a more f i r m c o n t r o l m i g h t be e l e v a t i o n , where you m i g h t g e t a lowland and a n u p l a n d d i s t r i b u t i o n of p l a n t s t h a t a r e c l o s e g e o g r a p h i c a l l y and m i g h t d i f f e r q u i t e a b i t . You see t h i s i n t h e Dunkard Group i n which you have what many p e o p l e s a y a r e Permian ( o r Lower Permian) o r A u t u n i a n e q u i v a l e n t s i n which t h e s p o r e s a r e of l a r g e l y C a r b o n i f e r o u s c h a r a c t e r . I t h i n k t h a t t h a t c a n be e x p l a i n e d by t h e f a c t t h a t t h e Dunkard b a s i n was s t i l l r a t h e r low, r e l a t i v e t o t h e s e o t h e r b a s i n s .
205
MESOZOIC FORAMINIFERA
-
WESTERN ATLANTIC
RICIIAFT, X. OLSSOK CEOLOGY 3EPARTMENT. RUTCERS UNIVERSITY, NEW BRUNSWICK, N. J. ABSTRACT Planktic foraminiferal zonal. control of Upper Cretaceous sequences is provided by species of the genera Globotruncana, Earginotruncana, Rotalipora, Hedberqella, Rugotruncana, Archeoglobigerina, Abathomphalus, and yentilabrella. Many of these species are tethyan so that recog:iition of tethyan biostratigraphic zones is apparent for the most part. Other species, however, are strictly boreal. Consequently, a zonal framework for this region reflects tethyan and boreal water as they influenced the region during specific time intervals of the Cretaceous. Due to preservational factors a planktic zonation for the Lower Cretaceous is at present incomplete. Benthic foraminifera1 assemblages are known from a few Lower Cretaceous and Upper Jurassic sections. They are sparse but apparently belong to shelf to abyssal facies. Upper Cretaceous assemblages occur in shelf, slope, and abyssal facies. Shelf assemblages include species of the genera Coryphost_om., Globulina, Guttulina, Gaudryina, and Gavelinella in addition to a number of laaenid species. Slope assemblaqes are characterized by species of the genera Gyroidinoides, Praebuliioina, Pullenla-, Eangularia, Gavelinella, Bolivinoidc-5. Dorothia and others. A number of these suecies also occur in abyssal planktic oozes. Biostratigraphic control is rather general being useful for the reconnition of state intervals. The Bolivinoides zonation is the one exception and it CO::;:J~~ments the planktic zonation in the Santonian, Campanian, and Maestr ichtian. ~~
INTRODUCTION The intent of this paper is to briefly summarize the available data on the Mesozoic foraminifera of the Western Atlantic exclusive of the Canadian Shelf (see papers by Ascole and Bujack, Gradstein and Williams in this volume). Most of the data comes from the Cretaceous but a little comes from the Jurassic. There are no Triassic foraminifera known from the Western Atlantic. Along the Atlantic margin of the U . S . sediments of Cretaceous age are exposed in the Atlantic Coastal Plain (fig. 1). The Cretaceous strata dip eastwardly beneath the coastal plain an?, continental shelf. Cretaceous strata also crop out along submarine canyons and the continental slope. In the adjacent Atlantic Basin a number of DSDP sites have penetrated Cretaceous deposits (fig. 1). Jurassic strata are not exposed in the coastal plain but they have been identified in a few coastal plain wells. Two DSDP drill sites have encountered Jurassic deposits (Sites 100 and 105).
206
The Upper Cretaceous of the Atlantic Coastal Plain consists of an alternating sequence of marine-marginal marine strata deposited during several cycles of transgression and resression. The Lower Cretaceous is entirely nonmarine in the coastal plain. These nonmarine sediments must extend some distance below the continental shelf for they are still encountered in coastal plain wells. Marginal marine Jurassic is suggested in some coastal plain wells. In the Atlantic Basin marine Lower and Upper Cretaceous and marine Jurassic were penetrated in the DSDP drill sites. Only beds of Campanian-Maestrichtian age have yielded foraminifera in coastal plain outcroppings (Olsson, 1960, 1964 and Mello et al., 1964). In the coastal plain subsurface more complete sequences of foraminifera which range in age from Cenomanian to Maestrichtian have been recovered (Maher and Applin, 1971; O'Grady, 1976; Olsson and O'Grady, 1976; Olsson and Petters, 1975; Perlmutter and Todd, 1965; and Petters, 1975). Foraminifera from DSDP sites have been reported on by Peterson et al.,(1970, Upper Cretaceous), Cita and Gartner (1971), Luterbacher (1972, Lower Cretaceous and Upper Jurassic), Hayes et al., (1972, Lower and Upper Cretaceous), and Lauqhton et al. (1972, Lower and Upper Cretaceous). Lower and Upper Cretaceous strata were penetrated in DSDP drill Sites 382, 384, 385, 386, 387, 390, 391, and 392 ( Tucholke et al., 1975; Eenson et al., 1975) but paleontologic reports are not yet available. The following discussion attempts to summarize the data from the above sources in terms of planktic foraminifera1 biostratigraphy, benthic forarniniferal biostratigraphy, and paleobathymetric distribution. DLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY A planktic forarniniferal biostratigraphic framework has recently been presented by Petters (1975, 1976) for the Upper Cretaceous Atlantic Coastal Plain. This zonation (fig. 2 ) is similar to that proposed for the western Sulf Coast (Pessagno, 1967) and also bears a number of similarities with the idealized zonation for the low latitudes given by Van Hinte (1976). This is undoubtedly due to warm water influences from southern areas in a smaller Atlantic Ocean. The utility of species of the genera RotaliapoE an?. S i n o t r u n z in CenomanianSantonian sections would seem to indicate a strong Tethyan influence in the Atlantic region during this time interval. On the other hand Campanian and Maestrichtian zones based on species of Archeoglobigerina and Rugotruncana, suggest diminished Tethyan influence in the Atlantic region. Influence of warmer water masses in the Atlantic during late Cretaceous time is apparently related to the development of major current systems during the opening of the Atlantic (Berggren and Hollister, 1974). Although many of the valuable biostratigraphic markers utilized in tethyan regions can be utilized also in the west-
207
10
Fig.
1
O u t l i n e m?.p of W e s t e r n A t . l a n t i c showing l o c a t i o n o f DSDP d r i l l i n g s i t e s w h e r e M e s o z o i c sect i o n s have been e n c o u n t e r e d . Dashed l i n e shows w e s t e r n l i m i t of A t l a n t i c C o a s t a l P l a i n a n d t h e d a r k e n e d a r e a s show where C r e t a c e o u s s e d i m e n t s outcrop. The numerous c o a s t a l p l a i n w e l l s t h a t p e n e t r a t e Mesozoic s e c t i o n s a r e n o t shown f o r l a c k of s p a c e .
208
P1,ANKT CC P O R A M I N I P E I U i L
BOLIVINOIDES
ZONATION
ZONA'T I O N
B 01, I V I NO I DE S DKACO
GLGBOTRUNCAIJA G A N S S E R I RUGOTRUNCANA SUBC JRCUIINODIFCR G.
B.
CALCARA'CA B.
G.
IIILARIS
DECORATUS
ELEVATA
V E N T I L A D R F L L A GL4BRATA Ai?CI!COGI,OBIGBRINA
G.
BLOVII
rORNICATA
3.
CULVEREIJSlS
3.
STRIGILLATU!
MARGINOTRUNCANA CONCAVATA
ROYAL I P0K A C U S IIP TAN I
Fiq" 2
P l a n k t i c f o r a m i n i f e r a 1 z o n a t i o n and B o l i v i n o i d e s z o n a t i o n for A t l a n t i c C o a s t a l Plain (after P e t t e r s , 1976).
209
ern Atlantic region, in general these elements are much rarer. Upper Cretaceous planktic foraminifera1 faunas of the Western Atlantic are dominated more bv species of alobiserine forms such as Hedbergella, Globigerineila, Rugogiobigerina, Archeoglobigerina and Rugotruncana than are Tethyan faunas. Faunal distributions in the late Cretaceous of Atlantic planktic forasinifera appear to be characterized by broad geographic range of many species with significant changes in abundances from one region to the next. Biogeography would appear to be a fruitful line of study for this fossil group. Lower Cretaceous planktic foraminifera have been identiin Deep Sea Drilling Sites 101, 105, 111, 143 and 144 but because of preservational factors a zonation is yet to be realized. Nevertheless, the presence of species such as Hedbergella trochoidea, E. infracretacea, E. hauterivica, E. globigerinelloides, Rotalipora appenninica, Globigerinel-loides eaglefordensis, G. ultramicrus, and Planomalina buxtorfi attest to the aiversity of species in the Atlantic during Early Cretaceous time and assure that in suitable sections planktonic foraminifera will provide a useful biostratigra2hic zonation. Deep Sea Drilling Sites 384, 386, 387, 390, 391, and 392 penetrated Lower Cretaceous sections in the Bermuda Rise, J. Anomaly, Blake Nose, and Blake-Bahama Basin but foraminifera1 reports are not yet available. Although planktic foraminifera offer little potential for zonation in the Jurassic where their origin lies, it is useful to note that Luterbacker (in Laughton et al., 1972) has identified the species "Globigerina" helvetico jurassica in the Tithonian-Kimmeridgian of site 105. BENTHIC FORAMINIFERAL BIOSTRAT IGRAPHY CANPANIAN
-
NAESTRICHTIAN:
The most useful group of benthic foraminifera for purposes of biostratiqraphy re species of Bolivinoides. Petters (1975) has shown that the Bolivinoides zonation (Earr, 1966, 1970) can be utilized in Atlantic Coas?:al Flain sections (fig. 2). This upper Santonian-hestrichtian zonation compliments the planktic foraminifera1 zonation. Although there is no comparable benthic zonation for older Cretaceous and Jurassic sections the presence of certain sFecies tends to characterize various stratigraphic levels. CONIACIAN-SANTONIAN : As noted above the evoluticn of Bolivinoides is seen in the upper Santonian with the advent of Bolivinoides strigj lI.at~.is. Eenthic species associated with the ConiacianSaiitonian include Citharina texana, C. wadei, C. simondsi, Tritaxia capitosa, ___- Planulina texana, and gaudryina ellisorae. Coniacian foraminifera are absent in updip sections of the Atlantic Coastal Plain due to an important and widely recognized disconformity.
210
CENOMANIAN - TURONIAN: Benthic assemblages are characterized by a rich diversity. In coastal plain wells these assemblages are associated with the well-known major transgression of the AlbianTuronian. In the Atlantic Coastal Plain this transgression covered the Early Cretaceous nonmarine sediments with marine sediments of Cenomanian-Turonian age. Deep Sea Drilling has recovered Cenomanian-Turonian sediments at sites 101, 105 (Luterbacker, 1972); 111 (Van Hinte in Laughton et al., 1972); and 143, 144 (Beckman, in Hayes et al., 1972). Site 386 also encountered sediments of this age (Tucholke et al., 1975). A number of benthic species characterize assemblages of this age. These include Vaginulina debilis, Marginulina siliquina, Citharina kochi, Saracenaria duckcreekensis, Lenticulina gaultina, Eponides moremani, Hoeglundina charlottae. Valvulineria ~ infreauens. Gavelinella _ V. lotterlie. _ dakotensis, G. plummerag, 5.' baltIca and Gavelinopsis Emanica. The latter species appears not to ranqe above the Cenomanian (Van Hinte, 1976) EARLY CRETACEOUS: Marine sediments of Early Cretaceous age have not been identified so far in the Atlantic Coastal Plain. Sediments of this age have been recovered at sites 100, 101, 105 (Luterbacker, 1972); 111 (Van Hinte in Laughton et al., 1972); and 143, 144 (Beckman, in Hayes et al., 1972) of the Deep Sea Drilling Project. Sites 384, 386, 387, 390, 391, and 392 have encountered early Cretaceous sediments (Tucholke et al., 1975; Benson et al., 1975) but paleontologic reports are not yet available. Early Cretaccous benthic assemblages are rather sparse and so far do cot offer good stratigraphic control. Dorothia praehauteriviana is reported by Luterbacker (1972) as a possible marker sp,.?ciesfor the Valanginian. Species identified include Reophax helveticus, Textularia washitensis, T. rioensis, Spiro-lectavmina alexanderi, Lituola subgoodlandensis, Lenticulina s i x E G r m , L. ex gr. -_ muensteri, L. ouachensis ouzhensis, L. ouachensis multicella, Astacolus incurvatus, Discorbis minutrssima, Lingulina nodosaria, and GaveTinella cf. barremiana. JURASSIC: Although strata of Jurassic age have been questionable identified in a few Atlantic Coastal Plain wells no definitive analysis of foraminifera has been published. Luterbacker (1972) has reported on the diverse microfaunas found in Upper Jurassic sediments in sites 100 and 105 of the Deep Sea Drilling Project. He subdivided the benthic foraminifera into three groups, "primitive foraminifera" which predominate and consist of the simply structured arenaceous foraminifera, the lagenids, and other foraminifera not contained in the first two groups and which are minor elements of the assemblages. Some of the more typical species include Brotzenia mosquensis,
211
Bi enerina jurassica, B. arcuata, Bolivinopsis helvetoju:assicus, Marssonella doneziana, Trocholina t-sarii, _Lenticulina _ _ ~ polonica, _____ garginalina minuta, and "Planularia" pseudoparallela. PALEOBATHYMETRIC DISTRIBUTION Benthic foraminifera are well known for their importance in delineating environments of deposition and bathymetry in Cenozoic deposits. Few studies have used foraminifera for such interpretations in pre-Cenozoic deposits because of the uncertainty of the environmental significance of extinct species and genera. Sliter and Baker (1972) are the first to propose a bathymetric model for Cretaceous benthic foraminifera that defines the generic composition of shelf and slope assemblages. Their model is based on a study of CampanianMaestrichtian shelf and slope deposits of California. It is based on criteria which includes comparison of depth restriction of genera as deduced from recent distributions, variations in faunal diversity and abundance, recurrent species asscciations, and homeomorphic comparisons of Cretaceous and Recent species. In the Western Atlantic region, Cretaceous nonmarine, marginal marine, and shallow-middle shelf deposits are exposed in the Atlantic Coastal Plain, outer shelf-bathyal deposits are encountered in some coastal plain wells (Olsson, 1975; Olsson and O'Grady, 1976), and bathyal-abyssal sections are encountered in the DSDP drill sites. Some sites such as 111 (Laughton et al., 1972) are drilled in sunken portions of the continental margin and thus contain shallower-water deposits than the depth from which the samples were taken. Other sites must represent locations where bathyal-abyssal deposits of Cretaceous age are sampled. The following presentation is a preliminary effort to summarize the known foraminifera1 distributions and assemblages as bathymetric models following the approach of Sliter and Baker (1972). CAMPAN IAN -MAESTRICH TIAN : Four bathymetric assemblages of Campanian-Maestrichtian foraminifera can be recognized in the Western Atlantic region; inner shelf, outer shelf, bathyal, and lower bathyal-abyssal (figs. 3 and 4). The inner shelf assemblage is characterized by low species diversity and high dominance. Planktic foraminifera occur rarely in this assemblage. Benthic species that tend to dominate in this assemblage are Gavelinella henbest& and Praebulimina carseyae. Outer shelf assemblages are Characterized by increased diversity of species, a greater variety of lagenid and rotaliid species. Species of planktic foraminifera become prominent associates of this assemblage. The more abundant species of this assemblage include Anomalinoides nelsoni, A_. pinguis, zlavulina trilaterus, Pulsiphonina prima, and Coryphostoma plaitum.
21 2
cc z w z
3
MILIOLIDS
GAVEL1 NELLA HENBESTI
LAGEN I D S
C I B I C I D E S HARPER1 GY R O I D I NO I D E S DEPRESSA VALVULI hER1A ALLOEORPH INO I D E S
GLOBUL I NA LACR IMA GIJTTbLINA NONIONELLA CRETACEA
GAUDRY I NA
P R A E 6 U L I K I N A ASPERA P P A E E U L I I I N A CARSEYAE U-I w
L U)
cc w +z 0
i
i F1.c;. 3
POLYKOP,PH I N I cs LAGEN I D S NEOFLABELLINA RETICULATA PRAEBULIrfINA CARSEYAE KYPHGPYXA CHiiISTNERI CORYPHOSTOXA P L A I T U M PULS IPHONI NA P R I r4A CLAVULINA TRI LATERUS NEOBULIMI NA S P I NOSA SIPHOGENERINOIDES PLUMWERAE
ANOMALINOIDES P I N G U I S AN OKA L IN0 ID ES 11E L S ON I GPVELINELLA N A C A T 0 C H E N S I S ANGULOGAVELI NELLA GRAC I? I S ALLOKORPHINA ALLORORPHINOIDES HOEGLUNDINA SUPRACRETACEA BOLIVINOIDES TEXTULAR IA R I PLEYENSI S
Tentative bathymetric model of Campanian ;Caestrichtian shelf assemblages based on distribution of foraminifera in Atlantic Coastal Plain outcrop and well sections.
213
STENS IOE IN A
RZEHAKINA EPIC-ONA
CORYPHOSTOMA INCRASSATA
DOPOTH I A OXY CONA
P R 1,E BU L I M INA ARK AD E L P H I AN A
DOROTHIA E L L I S O B A E
PRAEirULI M I N A K I CKAPOOEHSI S
DOROTH I A BULLETTA
ARAGONIA VELASCOENSI S
DOPOTHIP PETUSA
GY R O i D I NO I D E S N I T I D U S GY R C I D I I I O I D E S G I EARDANUS C H I LCSTO?IELLA T R I N I C A D E N S I S
A F El! 9 B U L I M I hA
F RP N K E I
AREUOBLiLIF11 NA SUBSPYAEPI CA GPUDRYI NA LAEV I GATA
CSANGULAR! A NAVARROANA
C L A V U L I V A T R I LATERUS
B 0 L I 1’!N C ICES
BATHYS I PHON
P U L L E N I A CRETACEA
S P I ROPLECTAWINA
STi LCSTOEELLA PYRAPiIDIJA SZAJNOCHAE PLECROSTOKELLA TORT!
Fig. 4
ARAGONI A VELASCOENSIS
BFiTHY S I PHON
P RAEBU L IM INA TAY LO REN S I S
P E L O S I N A COMPLANPTA
LAGENA OS All G U LAR IA
S P I ROPLECTANMI NA P E X I A E N S I S T R O C H A W I NO I D E S IRREGULARI S
D O R J T H I A OXYCONA
GLOP’iOSP IRA GORDI A L I S
RZEH AY I NA EP IGONA
VEPNEUI L I NA
Tentative bathymetric model of CampanianMaestrichtian bathyal-abyssal assemblages based on distribution of Foraminifera in Atla-ntic Coastal Plain wells and DSDP drill sites.
214
CITHARINA TEXANA
T P I T A X I A CAPITOSA
CITHARINA WADE1
CLAVULINA TRILATERUS
CITHARINA S IMONDSI
PLANULINA TEXANA
LAGEN I D S
GYPOIDI NOIDES UMBILICATA
POLYMORPHIN I D S
GYROIDINOIDES DEPRESSUS HOEGLUND I NA SUPRACRETACEA
NONIONELLA ROEUSTA PRAEBULI M I NA CUSHMAN I LENTICULINA PRAEBULIMI NA FAB IL I S
BEPTHELINELLA BPTHYSIPHON
CORYPHOSTOMA I NCRASSATA
VULVULINA
GYRO IDINOIDES N I T I D U S VALVUL INER IA PLUYMERAE
ARE WE!! L 1!I I NA DOPOTH I A
E L L IP S 0 GLAND U L IN A
TEXTULARI A
S T I LOSTOFELLA
CLAVULINA TRILATERUS
BATHY S IPHON
STENSIOEINA
REOPHAX
BANCYELLA
LITUOTUBA AMMGDISCUS Fig. 5
Tentative bathymetric model of Coniacian-Santonian shelf-abyssal assemblages based on distribution of foraminifera in Atlantic Coastal Plain wells and DSDP drill sites.
215
LAGEN IDS POLYRORPH I N IDS
HAPLOPH RAGMO I DES TROCHAMMINA
VAGINULINA D E B I L I S
GAVELINELLA DAKOTENSIS
MARGI NULINA
Fig. 6
SI L I Q U I N A
GAVEL I NELLA M I N IMA
NEOBUL I MINA ALBERTENS IS
GAVEL1 NELLA PLUMMERAE
PRAEBULIMI NA EXIQUA
CASS IDELLA TEGULATA
EPONIDES MOREMANI
CERATOBULI M I NA PARVA
QUI NQCELOCL’LINA
HOEGLUNDI NA CHARLOTTAE
LAGEN IDS
GAVELINELLA BALTICA
POLYMORPHINIDS
GAVELINELLA DAKOTENSIS
CITHARINA KOCHI
GAVELINELLA MONTERELENSIS
VAG I N E L 1 NA DEB IL I S
GAVEL1NELLA PLUMMERAE
MARG INUL I NA S I L I QUI NA
GAVELINOPSIS CENOMANICA
SARACENARIA BONONIENSIS
VALVULI NERI A I NFREQUENS
SARACENARI A DUCKCREEKENS I S
VALVULI NERI A LOTTERLI E
LENTICULINA GAULTINA
HOEGLUNDI PIA CHARLOTTAE
BU L I M I iiE LLA FAB I L I S
GAUDRY I NA
FURSENKOI NA CP.ONE ISI
SPIROPLECTAFMINA
NEOBULIEINP, ALRERTENSI S
QUINQUELOCULINA
Tentative bathymetric model of Turonian-Cenomanian shelf assemblages based on distribution of foraminifera in Atlantic Coastal Plain wells and DSDP drill sites.
216
Bathyal assemblages contain a qreater diversity of arenaceous species than do-snelf assemblages. Species of Dorotnia, Arenobulimina, Gaudryina, Clavulina, and S y s i p h o n are prominent. Typical calcareous genera include Stensioeina, bulimina, Gyroidinoides, Csangularia, Pullenia, Chilostomella, and Bolivinoides. Planktic foraminifera1 species are welldeveloped and diverse.
=-
Lower bathyal-abyssal assernhlages are distinguished by dominance of planktic foraminifera and low numbers and diversity of benthic species. Benthic species include Aragonia velascoensis, Rzehakina epigona, Pelosina complanata, and Glomospira gordialis. CONIACIAN-SANTONIAN: Little information is available on assemblages of this age in part due to the Coniacian disconformity. Shelf assemblages appear to be characterized by J. diversity of lagenids of which species of Citharina are prominent; bathyal assemblages contain the genera Praebulimina, Gyroidinoides, -Bathy_-siphon, Arenobulimina, Dorothia, and Clavulina, among others; and a few benthic genera are observed in bathyal-abyssal assemblages (fig. 5) TURONIAN-CENOMANIAN: Four bathymetric assemblages, an inner shelf, outer shelf, bathyal, and lower bath:ial-abyssal, can be recognized in this interval (figs. 6 and 7). Species which tend to dominate the low diversit-; inner shelf assemblaqe are Prae_____ bulimina exiqua, Gavelinella dakotensis and minima. lundina charlot,tae. is fairly common. Planktic species are rare.
G.
s-
Outer shelf assemblages are characterized by increased diversity of species, a variety of lagenid s2ecies of which a number are typical of the aqe, and prominent association of planktic species. Abundant-species-include Gavelinella plumerae, G. nonterelensis, Valvulineria lqtterlie, and ~ _ _ Buliminellz fabilis. The deeper bathymetric assemhlages are composed of species of Neobulimina, Gavelinella, Valvulineria, G y r o i d i n o i s , Bathysiphon, Dorothia, Ammobaculites, Ammodiscus, Spirillina, and Glomospira which in addition to a variety of other arenaceous taxa contrasts distinctly with the shelf assemblages. EARLY CRETACEOUS: Luterbacker's (1972) study of sites 100, 101 and 105 showed that the lower Cretaceous sections are characterized by sparse assemblages of foraminifera, the composition of which consists of a predominance of simply structured arenaceous foraminifera and rare lagenids. He compared these to similar assemblages that occur in Early Cretaceous strata of the Alpine-Mediterranean region which are interpreted as being le-
217
NEOBULIMINA M I N I M A
GAVEL1 NELLA MONTERELENS IS
NEOBULI PiINA ALBERTENS IS
V A L V UL I N EFI I A IN F PEQ UENS
BU L I M I NE L L A FAB I L I S E P I S T O M I N A LACUNOSA
VALVULINERIA LOTTERLIE HOEGLUNDI NA CHARLOTTAE
LAGENA SULCATA GAVEL I N EL L A
BATHY S I PHON
L E N T I C U L I NA NODOSARi I D S
REOPHAX H E L V E T I CUS
OS ANGU LA R I A
N EOB I,L I i4 I NA GY ROIDIQOIDES AFF, N I T I D U S
AMMOD I SCUS
GY90IDIW I D i S OCTACAMERATUS
SP I ROPLECTAMMI NA SP I R I LLI NA
L I NGULINA PSEUDONODOSARIA TUBEROSA PSEUDOTEXTULARI I D A E Fig. 7
DCROTH I A AMPOBACULITES TEXTGLARIA
TROCHAMPII NA GLOXOSPIRA
Tentative bathymetric model of Turonian-Cenomanian bathyal-abyssal assemblages based on distribution of foraminifera in Atlantic Coastal Plain wells and DSDP drill sites.
218
LAGEN IDS
GAVELINOPSIS CENOMANICA
POLYMORPH I N I D S
GAVELINELLA AMRONOIDES
VALVU L INE R I A
ARENOBULIMINA PRESSLI I
LENTICULINA SAXOCRETACEA
QUIMQUELOCULINA SABELLA SP I ROPLECTAW I N A ALEXANDER1
L I NGULIMA NODOSARI A D I SCOEB I S MI NUT1 SSIPIA PATELLINA SUBCRETACEA
TEXTULARIA RIOENSIS L I TUOLA SUBGOODLANDENS IS
LAGENIDS
HYPERAKMI NA
GAVEL1 NELLA
HAPLOPH RAGMO I DES
GY ROIDINOIDES
GLONOSPI RA
BATHYSIPHOI
RHIZAEYIINA
REOPHAX HELVETICUS
PROTEON I NA
DOROTHIA PRAEHAUTERIVIANA
LAGENAMMI NA
AMMOEACULITES
TOLY PAMMINA
TROCHAMKI NA
VERNEU I L I N O I D E S
AMMODISCUS
AMMOVERTELLA
SPI R ILLINA
HECHTINA
Fig. 8
Tentative bathymetric model of Lower Cretaceous shelf-abyssal assemblages based on distribution of foraminifera in DSDP drill sites.
219
posited in bathyal to abyssal water-depths (fig. 8). This view is strengthened also by analogy with the simplystructured arenaceous assemblages found at great deptli in the oceans today. The absence for the most part of calcareous foraminifera also suggests deposition below the carbonate compensation depth. The Lower Cretaceous in Site 111 penekrated shallow-water deposits (Van Hinte, in Laughton et. al., 1972) The assemblages in this section consist of lagenids, po'ymorphinids, and species of the genera Gavelinella, Gavelinopsis, Arenobulimina, Valvulineria. This compares well with the shelf assemblages of the Upper Cretaceous. The analysis of foraminiferal distributions in Sites 384, 380, 387, 390, 391 and 392 will add valuable data to the understanding of Early Cretaceous bathymetric assemblages. JURASSIC: The foraminifera1 faunas of Sites 100 and 105 are generally rich and consist mostly of simple arenaceous foraminifer2 and lagenids (Luterbacher, 1972). They are typical of Jurasic assemblages found in many other parts of the world. The bathymetric interpretation, as noted by Luterbacher, for these assemblages is difficult because of the much lower diversity than assemblages which succeed them in time. Jurassic foraminifera were probably very broadly niched. The Sites 100 and 105 assemblages conpare with shelf assemblages as defined by Gordon (1970) but the dominance of "Spirillina" in some sections compare favorably with similar faunas from deposits interpretcd as bathyal in origin from the Central Appennines. Obviously, more data is needed on the bathymetric composition of Jurassic foraminifera. ACKNOWLEDGEMENT "Acknowledgment is made to the Donors of the Petroleum Research Fund, administered by the American Chemical Society, for partial support of this research". REF E RENCES Barr, F.T., 1966. The foraminifera1 genus Bolivinoides from the Upper Cretaceous of the British Isles: Palaeontology, V. 9, p . 220-240. I 1970. The foraminifera1 genus Bolivinoides from the Upper Cretaceous of Libya: Jour. Palaeontology, v. 44, p. 642-654.
Benson, W. E., et al., 1975. Summary of Deep Sea Drilling Project: Leg XLIV. Unpublished. Berggren, W. A., and Hollister, C., 1974. Paleogeography, paleobiogeography and the history of circulation of the
220
Atlantic Ocean: 2 W. W. Hay (Editor), Studies in Oceanography. SOC. Econ. Paleontol. Mineral., Spec. Publ., v, p. 126-186. Cita, M. B., and Gartner, S. Jr., 1971. Deep Sea Upper Cretaceous from the Western North Atlantic: In Proceedings of the I1 Planktonic Conference, Roma 1970 p. Farinacc1,Ed.) p. 287-319. Gordon, W. A., 1970. Biogeography of Jurassic foraminifera: Bull. Geol. SOC. Am. v. 81, p. 1689-1704. Hayes, D. E., et al., 1972. Shipboard Site Reports: Hayes, E. E., Pimm, A. C., 1972, Initial Reports of the Deep Sea Drilling Project, Vol. XIV, Washington ( U. S. Government Printing Office) p. 1-338. Laughton, A. S., et al., 1972. Shipboard Site Reports, Site 111: 1 2 Lauqhton, A. S., Berggren et al., 1972, Initial Reports of the Deep Sea Drilling ProjecL, Vol. XII, I.iashington (U. S. Government Printing Office) p. 33-160. Luterbacher, H., 1972. Foraminifera from the Lower Cretaceous and Upper Jurassic of the northwestern Atlantic: Kn- Hollister, C. D., Ewing, J. I., et al., 1972, Initial Reports of the Deep Sea Drilling Project, Vol. XI. Washington (U. S. Government Printing Office) p. 561-594. Maher, J. C., and Applin, E. R., 1971. Geologic framewosk and petroleum potential of the Atlantic Coastal Plain and Continental shelf: U. s. Geol. Survey Prof. Paper 659, 98 p . Mello, J. F., Elinard, J. P., and Owens, J. P., 1964. Foraminifera from the Exogyra ponderosa zone of the Marshalltown Formation at Auburn, New Jersey: U. S. Geol. Survey Prof. Paper 501-B, p . 61-63. Olsson, R. X., 196C. Foraminifera of Latest Cretaceous and Earliest Tertiary Age in the New Jersey Coastal Plain: Jour. Paleontology, v. 34, p. 1-58.
, 1964. Late Cretaceous planktonic foraminifera from New Jersey and Delaware: Micropaleontology, v. 10, p. 157-188.
, 1975. Upper Cretaceous and Lower Tertiary Stratigraphy of New Jersey Coastal Plain: Second Annual Field Trip Guidebook, Petrol. Exploration SOC. N. Y., p. 1-49. Olsson, R. K., and O'Grady, M. D., 1976. Cretaceous and early Tertiary Paleobathymetric history of New Jersey Coastal Plain: Annual Meetings, New Orleans, Abstracts, Amer. Assoc. Petrol. Geol., Bull., v. 60, p. 704. Olsson, R. K., and Ulrich, B. C., 1976. Timing of Transgressions and Regressions in Cretaceous and Tertiary of New Jersey: Abstracts, Annual Meetings New Orleans, Amer.Assoc.
221
Petrol. Geol., Bull., v. 60, p. 704. Perlmutter, N. M., and Todd, R., 1965. Correlation and foraminifera of the Monmouth Group (Upper Cretaceous) Long Island, New York: U. S. Geol. Survey Prof. Paper 483-1, p. 124. Pessagno, E. A., Jr., 1967. Upper Cretaceous planktonic foraminifera from the Western Gulf Coastal Plain: Paleontographica Americana. v. 5, no. 37, p. 245-445. Peterson, M. N. A., et al., 1970. Shipboard Site Reports: In Peterson, M. N. A., et al., 1970, Initial Reports of the Deep Sea Drilling Project, Vol. 11. Washington (U. S. Government Printing Office) p. 1-306. Petters, S. W., 1975. Subsurface Upper Cretaceous stratigraphy and foraminifera1 biostratigraphy of the Atlantic Coastal Plain of New Jersey: Unpublished Ph. D. thesis, Rutgers University, New Brunswick, N.J., 258 p.
, 1976. Upper Cretaceous Subsurface stratigraphy of Atlantic Coastal Plain of New Jersey: Amer. Assoc. Petrol. Geol., Bull., v. 60, p. 87-107. Sliter, W. V., and Baker, R. A., 1972. Cretaceous bathymetric distribution of benthic foraminifers: Jour. Foram. Res., v. 2, no. 4, p. 167-183. Tucholke, B., et al., 1975. Summary of Deep Sea Drilling Project: Leg XLIII. Unpublished. Van Hinte, J. E., 1976. A Cretaceous time scale: Amer. Assoc. Petrol. Geol., Bull., v. 60, p. 498-516.
222
EXPLANATION OF PLATES All figured specimens are from the Cretaceous of New Jersey. Stage refers to the stratigraphic level from which the figured specimen was taken. Plate I A.
Dorothia ellisorae (Cushman), X100, Campanian.
B.
Dorothia stephensoni Cushman, X 110, Santonian.
C.
Dorothia retusa (Cushman), X 1 5 0 , Campanian.
D.
Clavulina clavata Cushman, X 35, Santonian.
E.
Tritaxia capitosa serrulata (Cushman), X 45, Santonian.
F.
Tritaxia capitosa (Cushman), X 6 5 , Maestrichtian.
G.
Clavulina trilatera- plummerae (Sandidge), X 3 5 , Campanian.
H.
-Clavulina trilatera Cushman, X 5 0 ,
I.
Meterostomella faveolata (Marsson), X 75, Campanian.
J.
Texularia ripleyensis W. Berry, X 100, Campanian.
X.
Arenobulimina subsphaerica (Reuss), X 110, Campanian.
L.
Eggerella trochoides (Reuss), X 1 6 0 , Campanian.
Campanian.
Plate I1 A.
Bolivinoides decoratus (Jones), X 120, Campanian.
B.
Chilostomella trinidadensis Cushman and Todd, X 9 0 , Maestrichtian.
C.
Pullenia cretacea Cushman, X 125, Campanian.
D.
Praebulimina kickapooensis (Cole) X 9 5 , Maestrichtian.
E.
Siphoqenerinoides plummerae (Cushman), X 90, Campanian.
F.
Praebulimina cushmani (Sandidqe), X 150, Campanian.
G.
Praebulimina carsayae (Plummer), X 1 2 5 , Campanian.
H.
Vaqinulina debilis (Berthelin), X 120, Cenomanian.
I.
Citharina kochii (Roemer), X 90, Turonian.
J.
Saracenaria duckcreekensis Tappan, X 9 0 r Cenomanian.
K.
Marginulina siliquina Eicher and Worstell, X 120, Cenomanian.
223
P l a t e I1 ( c o n t ' d ) Osangularia navarroana
L.
( C u s h m a n ) , X 1 5 0 , Carnpanian.
P l a t e I11 Hoeglundina c h a r l o t t a e ( V i e a u x ) , X 150, Turonian.
A.
B
-
Valvulineria l o t t e r l i e D - X 3 0 0 , Turonian.
D.
(Tappan) , B
E , F . G y r o i d i n o i d e s g l o b o s u s (Hagenow), E Campanian. G,H G l o b o r o t a l i t e s rnichelinianus
-
-
X 360, C
X 135, F
-
-
X 450,
X 185,
(D'Orbigny) , X 1 1 0 ,
Campanian. G a v e l i n e l l a plummerae ( T a p p a n ) , X 1 5 0 , T u r o n i a n .
I - IC.
B a t h y s i p h o n a l e x a n d e r i Cushman, X 3 0 , S a n t o n i a n .
L.
Plate I V A
-
C.
V a l v u l i n e r i a i n f r e q u e n s Morrow, A , B C - X 2 5 0 , Cenornanian.
D , E . G a v e l i n e l l a ammonoides ( R e u s s ) , D Carnpanian.
-
-
X 300,
X 120, E
-
X 140,
F,G.Anomnlinoides p i n g u i s ( J e n n i n g s ) , X 1 5 0 , M a e s t r i c h t i a n . H,I.=elinella
n o c h a t o c h e n s i s ( C u s h r n a n ) , X 1 5 0 , Carnpanian.
J,K.Anornalinoides
nelsoni
(W.
B e r r y ) , X 2 2 5 , Carnpanian.
224
PLATE I
225
PLATE I1
226
PLATE I11
227
PLATE IV
228
Discussion Dr. F. M. Gradstein: Do you have indication that the ratio of Globotruncaniids/Hedbergelliids and Heteroheliciids is related to distance from shore? The latter two groups would tend to, predominate the more shallow deposits, but are by no means confined to them. Globotruncaniids tend to favour more offshore conditions. Such a simple distribution has been brought forward earlier by Dr. W. Sliter and Dr. V. Scheibnerova. In my own experience this might be true also for the Upper Cretaceous of the Grand Banks and Scotian Shelf. Olsson: We know that Recent species of planktic Foraminifera float within a certain depth range in the water column; some are shallow floaters whereas others utilize a deeper water column. As the water column becomes shallower and shallower over the continental margin the ecologic column of planktic species is interfered with because it j.s intersecting the sea floor. And so those species that are deep floaters in the adult stage cannot occur in shallow waters. They are more strictly oceanic forms. Others which are shallow water floaters can occur farther in over the shelf, particularly if they are very abundant. As an example, rubers in the present seas and other globigerinid species show this relationship. Some of the Hedbergellas seem to have done that and the heterohelicids, in particular, occur in shallow water deposits. Matter of fact, there were a lot of questions on whether some species of heterohelicids were really true planktic forms because they are found in situations which were obviously shallow water. It was only until we began the DSDP where we know we are dealing with very deep water material that we could say for certain that some of them floated in the oceans. But there is this, you'll lose diversity of planktic forms in shallow water and in some cases vlanktic forms are absent. Such genera as the Archeoglobigerinas, the Rugoglobigerinas and others appear to have been very shallow water dwellers because they-occur in shallow wate; sediments. The globotruncanids apparently preferred open water conditions and perhaps were deep floaters in the adult stage because they are almost absent from shallow water deposits. Thus the criteria of diversity of benthic species, the diversity of planktic species, and the changes in the abundances of certain planktonic taxa such as the heterohelicids all go together in pointing towards either shallow water or deeper water deposition. Dr. H. Habib: Do the benthonic Foaminifera represent minimum depths, since they can be carried downslope posthumously? Can you use abundances that way? Olsson: Most transportation of Foraminifera that we are familiar with usually is with turbidites-turbidity current transported Foraminifera; and that becomes quite obvious in a change of sediment texture and in the occurrence of different assemblages. There are some fairly good examples of that kind of thing. But we really don't see much evidence of massive transportation of Foraminifera such that it destroys the whole profile so that you don't know what you're doing, You don't see that. Usually one can spot forams that are transported and out of place. They're usually not very abundant either. Mostly rare forms. Now of course in the Cretaceous we don't really know for sure what the bathymetric ranges of species were, but we can compare them with their
229
modern a n a l o g s , i n some c a s e s v e r y w e l l and i n o t h e r c a s e s n o t so w e l l . Morphotype a n a l y s i s i s a u s e f u l a p p r o a c h . I t becomes e v i d e n t when a g e n u s had a s h a l l o w e r r a n g e t h a n i t d o e s t o d a y ; f o r i n s t a n c e s p e c i e s o f O s a n g u l a r i a . Back i n t h e C r e t a c e o u s and P a l e o c e n e t i m e s some s p e c i e s of t h e genus o c c u r r e d i n s h a l l o w s h e l f d e p t h s b u t t h e y a r e v e r y r a r e and s m a l l i n s i z e . They were d w e l l e r s o f t h e g r e a t e r w a t e r d e p t h s , t o o and t o d a y t h e g e n u s i s c o n f i n e d t o t h e d e e p e r water d w e l l e r s . D r . J . E . Conkin: Can you g i v e any r e a s o n s f o r a g g l u t i n a t e d F o r a m i n i f e r a b e i n g found i n d e e p ( o r d e e p e r ) w a t e r i n t h e Mesozoic-Cenozoic whereas t h e y a r e found a b u n d a n t l y i n s h a l low w a t e r d u r i n g t h e P a l e o z o i c . Could t h i s be p a r t i a l l y comp e t i t i o n between a r e n a c e o u s and c a l c a r e o u s forms b e g i n n i n g i n m i d d l e Devonian and r e s u l t i n g i n t h e g r a d u a l d i m i n i s h i n g of a r e n a c e o u s forms from t h e e p i c o n t i n e n t a l n i c h e s , and t h u s dominance of t h e c a l c a r e o u s w a l l e d f o r m s t h e r e ? Olsson: W e l l t h e a r e n a c e o u s forams 0ccu-C from t h e l a g o o n s r i g h t o u t i n t o t h e deep sea Conkin: Though you u s u a l l y t h i n k o f them a s b e i n g d e e p w a t e r and t h e y a r e v e r y a b u n d a n t i n d e e p w a t e r . O l s s o n : T h e i r d i v e r s i t y i n s h a l l o w w a t e r i s v e r y much l e s s than it i s i n t h e deeper water, Conkin: And t h e abundance of them a s w e l l , i s n ' t t h i s t r u e ? Olsson: The abundance? W e l l it d e p e n d s , i n c e r t a i n s h a l l o w w a t e r s e d i m e n t s you c a n g e t v e r y h i g h dominance of T e x t u l a r i a , 9 9 % o r something l i k e t h a t . I t could be very abundant t h e r e . With t h e a d v e n t of t h e R o t a l i n i i d s , t h e c a l c a r e o u s f o r m s , t h a t t h e r e was a s i g n i f i c a n t r a d i a t i o n o n t o t h e cont i n e n t a l s h e l f . T h a t may i n t u r n have l e d p e r h a p s t o t h e e x t i n c t i o n o f many a r e n a c e o u s forms t h e r e o r t h e crowding o u t o f them. Conkin: Could t h e r e b e a n y c o m p e t i t i o n between t h e s e c a l c a r e o u s forms and t h e a r e n a c e o u s o n e s ? Olsson: Some d e e p e r w a t e r a s s e m b l a g e s a r e composed of mixt u r e s of c a l c a r e o u s and a r e n a c e o u s t y p e s , some a s s e m b l a g e s a r e a l m o s t e n t i r e l y a r e n a c e o u s and i n o t h e r s c a l c a r e o u s forms a r e dominant. Faunas below t h e c a r b o n a t e c o m p e n s a t i o n d e p t h a r e e n t i r e l y arenaceous. Conkin: I s i t n o t u n u s u a l t o f i n d a b u n d a n t a r e n a c e o u s forams i n beach d e p o s i t s t o d a y ? O l s s o n : T h a t d e p e n d s where you a r e b u t i n g e n e r a l p e r h a p s it is true. M r . M. Polugar: Did I g e t t h e i m p r e s s i o n awh:.le ago t h a t you were i m p l y i n g t h a t t h e r e was a d i r e c t c o n n e c t i o n between a k e e l e d form and d e p t h r a t h e r t h a n t e m p e r a t u r e ? You mentioned something a b o u t s i n g l e - k e e l e d G l o b o t r u n c a n a v s . d o u b l e keeled ones. Olsson: I n shallow water sediments Globotruncanas, s i n g l e o r d o u b l e - k e e l e d a r e e i t h e r a b s e n t o r e x t r e m e l y r a r e ; so t h e y l o o k a s i f t h e y p r e f e r r e d a more o c e a n i c h a b i t a t . Whether y o u ' r e up i n m i d - l a t i t u d e o r down i n t h e t r o p i c s t h e same t h i n g a p p l i e s . T h e r e may be d i f f e r e n t s p e c i e s b e c a u s e of t e m p e r a t u r e c o n t r o l , t h a t i s l a t i t u d i n a l c o n t r o l on t h e d i s t r i b u t i o n of s p e c i e s . P o l u g a r : I n C a l i f o r n i a , i n t h e Miocene, where w e have a v e r y d e e p w a t e r a b u n d a n t p l a n k t o n i c f a u n a , y o u ' l l f i n d no k e e l e d g l o b o r o t a l i i d s r e g a r d l e s s of how d e e p w a t e r you go i n t o . So you c a n ' t r e a l l y r e l a t e them t o d e p t h .
230
O l s s o n : W e l l , i t d e p e n d s , i n C a l i f o r n i a where t h e r e w a s i n f l u e n c e of c o l d w a t e r , y e s , you c o u l d n ' t do this. I n h i g h l a t i t u d e a r e a s , s a y A l a s k a o r t h e h i g h e r l a t i t u d e s of t h e A t l a n t i c Globotruncana can be expected t o b e a b s e n t . I d o n ' t t h i n k you c a n do t h e same t h i n g f o r GloboPolugar: I t h i n k s i n g l e - o r double-keeled Globotruntruncana e i t h e r . c a n a i s a f u n c t i o n of p o s s i b l e t e m p e r a t u r e and t i m e , a s you f i n d double-keeled Globotruncana as y o u . . . W e were Olsson: I d o n ' t t h i n k I s a i d anything about t h a t . t a l k i n g a b o u t t h e b a t h y m e t r i c p r o f i l e irnd how t h e d i s t r i b u t i o n of p l a n k t o n i c s i s r e l a t e d t o t h a t p r o f i l e .
...
231 JURASSIC OSTRACODA OF T H E ATLANTIC BASIK by
R.H. BATE B r i t i s h M u s e u m ( N a t u r a l H i s t o r y ) , L o n d o n SW7 5 B D ,
England
ABSTRACT Varine Jurassic sediments of t h e Atlantic Basin are widely developed in the eastern borderland region of the North Atlantic. Here. warm shallow seas supported a rich and varied ovtracod fauna over 200 species being found i n t h e English Bathonian alone. This region is regarded as being one of t h e major areas i n t h e development and evolution of Post-Palaeozoic Ostracoda. The e;istcrii borderlands may b c divided iiito a northerti European (epicontinctital sea) Province. ~
nd a southern North African province. Thc S o u t h Atlantic Basiii was not tlevelopccl a t t h a t time. On thc western borders of the North Atlantic Basin Jurassic sediments arc present along the cast coast of Grccnland and offshore along the K o r t h American continent ~. ostracods having affinities with the epicontincntal European f n u n a s are present off Canada, while d e e p water ostracotls, i n part cornp;irable with tethyan species f r o m Italy. are present off U.S.A.: shallower water faunas are recorded from t h e Gulf Coast region (American Province). Biostratigral?iiicaill~~ i m p o r t a n t ostracotl gcncra of t h e cpicontiiiental provinccq belong t o threc families: tlic Progonocq theridae, t h e S~hnlerideitlaeand the Protocytlicridac. Tliiq contrasts with the Jurassic of t h e Ilitiiali Ocecii Basin (East African Province) where on11 t h e Progotiocythcridae is of equal iiiiportaiice. Ostracod genera belonging t o o t h e r families play an iinportant b u t subsidiary role in the biostratigraphy of t h e North Atlantic Jurassic. Fresh t o 'srackish water ostracods become important in t h e Middle Jurassic b u t reach thcir biostrati:raphical acme in the continental deposits of the Uppcr Jurassic/Lower Cretaceouq when t h e genus Ci~pritlcaachieved a world-wide diqtribution. a iieeper u.ater T e t h y a n Proviti
INTRODUCTION The Atliiiitic Ocean was n o t in existence during Lower Jurassic times ;IS t h e scparation of t h e continents only coiiimcnced a b o u t t h e time o f Middle Jurastic. I n order, therefore. t o present the distribution o f t h e marine Jurassic sediments on a m a p of t h a t time a base m a p showing t h e situation during t h e Hauterivian has been used. This map [Text-fig. 1 I , t h e m o s t accurate o n e available. has been taken f r o m Owen (1976) a n d , because o f t h e tlow rate of displacement of the continents. shows a position o f t h e continents n o t t o o far removed from their position a t the close o f t h e Period. Before the separation o f t h e continents began. tnaritie conditions appear t o have been restricted to the north-eastern area o f t h e Atlantic: there is no evidence of marine sediments south of Nova Scotia until Upper Jurassic times. Indeed t h e southward transaression o f t h e Tethys appears t o have been dependent u p o n the opening of t h e Atlantic. Europe. on the o t h e r hand, was covered by a large epicontinental sea. interspersed with islands, t h a t e x t e n d e d
232
Y
Y
Tex 1-Fig. I
,
x
I
Outcroi> o f tn:irinc J u r x s i c sediiiieiits plotteJ oil :I base tiiw ? h o w i n c thc position of the continctits iliiritig thc Hautcrivi;in 11) from Owen 1976 I .
eastwards into Asia. In Middle t o Upper Jurassic times this linked northwards with thc epicontinental sea of north-western America. I t was through these large epicontinental seas that migration of the ostracods t o o k place. The east-west orientated Tethys appears t o have been a somewhet narrow h u t deeper water seaiocean t h a t acted more as a barrier than as a route for migration. This is especially noticeable when comparing the ostracod faunas of the north with those of the southern hemisphere. T h e study of the Jurassic ostracods from the Atlantic Basin indicates that four faunal provinces may be recognised: a European. a Tethyan, a North African and an American. T h e North African Province is perhaps rather tenuouc, a t this stage and may have t o be merged with the Tethyan Province at a later date -.
233 T h c ostracod fauna of thc East African Province is n o t directly relevant t o a stud) of the Atlantic Basin b u t is important in the context of being able t o identify a sotithern as distinct from a northern hemisphere fauna - it is in this way t h a t this province will be briefly involved. Althoiigh a t the present time the greatest diversity of faunas is gcnerally in the tropics and declines away from the equator this does not appear t o have been the case in the Jurassic where a more uniform climate existed. Circumstances a t the onset of the Mesozoic. folloLving o n after thc extinction of most of the Palaeozoic ostracods. were unique in that ecological conditions i i i certain parts of the world were ideal for speciation and subsequent ostracod evolution. Such conditions existed in Europe from the close of Lower Jurassic times onwards. T h u s by the cnd of the Lower Jurassic four important families were alrcad), established : thc Schuleridcidae. the Progonocythcridae. the Protocytheridac and the Cytheruridae. In the Middle Jurassic the ?-rachyleberididae and the Limnocytheridae appear while in thc Upper Jurassic the Ilyocypritlidae dominate the continental deposits. At the present time a zonation of the Jurassic is n o t available although some authors have initiated a zonal scheme for parts of the succession: Anderson ( 1 9 7 1 ) for the Purbeck: Bate (1965) in the Bajocian: Bate & Coleman ( 1 9 7 5 ) in the Toarcian and Michelsen (1975) f o r the Lower Jurassic. A zonal scheme for the British Jurassic is currently i n preparation by Bate. Kilenyi and Lord (see Bate & Robinton).
FAUNAL PROVINCES
The European Province. T h e lateral extent of this province r a n p from Nova Scotia in the west to the Ukraine in t h e east. Northwards the e x t e n t is n o t known as t h e Greenland faunas have n o t been described. southwards the epicontinental sea of the province merges with the deeper waters of the Tethys. . T h e provincial boundaries do n o t remain static through the entire period and it will be some time before precise boundaries can be drawn. Spain. for example. is considered t o have been a part of the European Province during the Lower Jurassic b u t with its anit-clockwise rotation through the Jurassic possibly moved into the Tethy an Province towards the close of the period. T h e maximum diversity of ostracod genera and species exists only a t the centre of the province extending through northern France. England, Germany and Poland. This region appears t o have heen covered by warm, shallow seas in which numerous islands cxisted. Thus not only are marine ostracods numerically a b u n d a n t b u t brackish water lagoonal and estuarine conditions existed f o r further speciation t o take place. FolIowing the extinction of most of the Palaeozoic lineages and a period when continental conditions prcdominatcd in the Trias, the marine transgression during the Jurassic entered a region here that was ideally suited t o ostracod colonisation and subsequent speciation. Because of this rapid evolution in the ostracods t h e European Province appears t o have been the birth-place of the majority of the important Mesozoic families. Indeed some families which evolved then are important a t the present time [c.g. Cytheruridae and Limnocytheridae] . Lower Jurassic - Hettangian t o Toarcian. Although sediments of Lower Jurassic age have been found off Newfoundland and Nova Scotia the ostracods have n o t y e t been described. I t does n o t appear t o b e unreasonable, however, t o tie this region in with the European Province particularly as Ascoli has noticed a relationship between the faunas for Middle and Upper Jurassic (per?. comm.).
234 In general the lower stages of the Lias are not as rich in ostracod species as is the case from the Sinemurian onwards. Initially the faunas are dominated by ogmoconchids and these have been used by Michelsen ( I 975) as the basis of his zonation of the Danish Lower Jurassic. The genus Kitikrlinellu first appears in the British Trias, evolves rapidly throughout the province and the many stratigraphically short ranging species make it an ideal ostracod for zonation (Bate & Coleman 1975). Indeed the two subgenera, Kinkeliriellu fliinkelinellul and Kinkelinellu (Ekt,vphocj,therel are the most important of the cytheracean ostracods I family Protocytheridae] developing at that time. The genus ~2licroprrci,mutoc~,there is first recorded from the Sincmurian [see Proci,therideu reticiilutu Klingler & Neuweiler 19591 and this is currently considered to be the earliest true record of the Progonocytheridae. This family replaces the Protocytheridae i n importance in the Middle and Upper Jurassic with the decline of the Kinkrlirielh-Ekt~'~Jhocj'there lineage, although the Protocytheridae still retain an important position in ostracod phylogeny and become importznt again with respect t o their use in stratigraphical correlation. Many new genera appear towards the close of the Lower Jurassic [ e.g. A phclocythere, Truchj,q.there, Xunuc,vthere & Proc>,thc.rirra] that are significantly important in correlation during the Toarcian and the Aalenian ~. there is no clear division between the Lower and the Middle Jurassic at this point with respect to the ostracod faunas. Also by this time the family Cytheruridae was well established [ the earliest record being of Cvtheropteron reticulutunz Michelsen (1 975) from the Lower Sinemuriaii] but the fourth important family, the Schulerideidae, was only just developing Prueschiilerideu pseudokinkelinella Bate & Coleman ( 1 975) from the Toarcian, the earliest representative, considered to be very close t o being the ancestor of the family. The Schulerideidae devclopcd too late to be important in the Lower Jurassic but achieved this role in the Middle and Upper Jurassic. Middle Jurassic - Aalenian to Callovian. The Aalenian has a fauna that links it closely with the Toarcian below hut at the same time contains ostracods having affinities with the Bajocian [e.g. Pnrrrmutoc.cthcre] . In particular the genns Prueschi/lerideu becomes important all over Europe where several spccies occur, all characteriscd by a coarsely pitted ornamentation (Malz 1966). Throughout thc other stages of the Middle Jurassic Prueschiilerideu remains important although its species become smooth-shelled [with a few exceptions - P. butei in the Callovian] and, through Eoschtilcridcu [see Bate 1967, p. 4 0 1 , gives rise to the Schirlerideu lineage sometime i n the early Bathonian. The Prue.whulerideu lineage dies out in the Callovian with P. hutci i i i Britain and P. uicciutu in southern France. that is so In the Bajocian the Progonocytheridae add the important genuy Glj~ptoc.i~there valuable for correlation all over the province. Indeed this genus is known to extend as far east as the Ukraine (Permjakova 1970) and, outside the province, as far east as Uzbekistan currently the (Masumov 1973, see ,I.lucrodcritinu usperu). The genus ;Microp~ieiirnutoc~~tliere, oldest of the family also develops rapidly through the Middle Jurassic where, together with Gl,i~proc,rthereit may be used in zonation. In fact a zonal scheme using species of these two genera is currently being prepared for publication (see Bate & Robinson). appears in the Bathonian of England and Poland more or less at The genus Progotroc~j~there about the Same time and once was thought to be restricted t o the Middle Jurassic. The many species subsecluently placed in this genus from the Upper Jurassic have proven t o belong more correctly to other genera, although one species, as yet undescribed, has bcen found in thc Kimmeridgian of Spain. Although giving its name to the family this genus has become
235 relatively unimportant stratigraphically. In the southern hemisphere, however, a very close relative of Progunoc~ytkere,the genus Mujiingudlu, becomes strikingly successful throughout the Middle to Upper Jurassic and through the Cretaceous. The geographical distribution of .Ilujiinguellu and the subgencra,'genera that developed from it is from Australia and Africa t o South America. This is quite a remarkable achievement considering that both genera certain11 arose from a common ancestor and became virtually restricted to their respectivc hemispheres [only one species of Prugo/iuc:i.there P. luec.i.rcz/lu in the Callovian t o Oxfordian of India is known from the south while no species of Muji/nguella are known from thc north]. Within the Progonocytheridae there are two morphological groups -- the first rather qtiadrate/oval in outline and represented by Progonocyfhere and Glj,ptucj,t/iereand the second having a much more elongate carapace as in Lophocj.tlierc and Firhrbergiellu. Both groups are charactcrised by having many short ranging species iniportant in the correlation of the Jurassic throughout this province. The Kirikelinellu Ekfj,p/ioc>.flierelineage; so important in the Lower and basal Middle Jurassic apparently dies out in the Bathonian to be continued in the Callovian by the genus Pseiitiokirtsoniu. Whatley ( 1970) rightly considers Pseirtiuhot,roiiia Weinholz (1967) to be congeneric with Buloit-c~lluWienholz ( I 967) both originally described from the Callovian of East Germany. Thus the rccord of Buloicvllu from thc Oxfordian of Canada (Brooke & Braun 1972) and of Pmdohiirsoniu from the Callovian and Oxfordian of Britain (Whatley 1970) extend7 the geographical range of this important lineage. Interestingly. lineages i n the northern hemisphere continue with many more changes at genus level than is the case i n the southern hemisphere. In this consideration of the faunal provinces the important ostracods have all been marine i n habit, the diversity of the faunas being measured by the fact that i n the Bathonian of England alone there are some 55 genera and over 100 species. Within this Province. however? the variety of ecological niches provides for the development of both brackish and of freshwater ostracods Bi.sirlc,oc.,i,/,ri.sand some of which are peculiar to the Middle Jurassic but some, Tlieriusj,noc.c,iii?i, Rlicumz, are also important i n later parts of the geological column. The gentis Davii.i/rirlu. extending through the Palaeozoic to the Recent is, of course, also represented, With thc exception of the last named, development of these new ostracods appears to have taken place within the European Province with subsequent migration effecting a world-wide distribution. This also appears to apply to Timiriuseviu and Lirnnucj'thcve although stratigraphically they are never as important as the first three. The record from Israel by Gerry & Oertli { 1967) of B i , ~ i i l m ( ~ i ~ p r i ~ from the Trias is probably misleading as the age of the beds concerned is now considered to be Jurassic (Gerry, pers. comm.). Upper Jurassic - Oxfordian to Tithoiiian. Upper Jurassic ostracods of the European Provincc continue the diversity of forms already encountered. The Progonocytheridae continue with Lophocj~tiiercs. 1. in the Oxfordian and with Macrudentiuu in the Kimmeridgian. Culliuc~c~therit/cu is characteristic of the Upper Oxfordian and Kimmeridgian of northern France (Oertli 1957) and of the Kimmeridgian of southern England (Kilenyi 1969) but is not well represented in the Upper Jurassic of Switzerland (Oertli 1959) close t o the Tethynn Province. It is in Switzerland, as well as i n southern France iDonze 1967,), on the margins of the Tethyan Province, that the true European representatives of Proc~i~flierit/c.u are found. Also within the Upper Jurassic the important Cretaceous genus Protuc,vfhere makes its appearance and is re~
~
corded by Oertli (1957) from the Kimmeridgian of the Paris Basin and by Pokorny i1973) from
236 the Tithonian of Czechoslovakia. Some early form ofProtocythere appears t o be present in the Bathonian of England but this still has to be verified. Otherwise the migration route of the genus would be from the east where it has been recorded from Uzbekistan (see Macrodentina aspera in Masumov 1973). Important amongst the late Jurassic marine ostracods is Paranotacythere - species of which are stratigraphically short ranging but geographically widely dispersed throughout the epicontinental sea area of the province (Bassiouni 1974). Large areas of the province became emergent during the final phase of the Jurassic and continental deposits with fresh and brackish-water ostrdcods stretch from Denmark through England and France and northern Germany into northern Spain and Portugal. Indeed such conditions existed over large areas of Asia, North and West Africa and South America during this time. Freshwater ostracods first appear in the Upper Jurassic of Portugal where they have been utilised by Helmdach (1971) to subdivide both the Oxfordian and the Kimmeridgian. It is only in the Upper Kimmeridgian that Cypridea begins to dominate the freshwater faunas both in Portugal and in the offshore Kimmeridgian of southern Ireland. Henceforth Cvprideu becomes established world-wide in the Jurassic - Cretaceous interval. Indeed: the use of ostracods in zonation was first employed in the Purbeck by Forbes as early as 185 1. Subsequently refinements have been made and the zonal importance of this genus in the British sequence owesmuch t o the work of F. W. Anderson. The Tethyan Province. In Lower Jurassic times this province extended as far west as the Canadian coast (Nova Scotia and Newfoundland) but expanded in the Upper Jurassic into the southern Atlantic Basin. The north-south extent of the Province takes in western North Africa and the southern part of Europe. Lower Jurassic - Hettangian t o Toarcian. As the Lower Jurassic marine transgression spread north and west over Europe the ancestral stock of the new cytheracean lineages was introduced. The resultant diversity that ensued developed away from the Tethys which was a deeper water environment characterised in the main by smooth-shelled ostrdcods. In the Lower Jurassic the genera Ogmocoxcha and Ogmoconchdla dominate the Tethyan faunas, especially in the Pliensbachian where highly ornate species of Ogmoconcha occur in southern Germany (Lord & Moorley 1974c and Malz 1975), Sicily (Barbieri 1964a) and in the Djebel Zaghouan section in Tunisia. Material collected from the Schwabische Alb [SW-Germany] exhibits both a preponderance of ogmoconchids and some of the diversification more commonly associated with the European Province; as such this region is considered to have been situated on the boundary between the two at that time. The fauna described by Drexler (1958) from Bavaria. south of the Schwabische Alb, is of a deeper water fauna more typical of the Tethys. Similarly the Hettangian fauna in the A r d k h e [S. France1 described by D o m e (1966) is also of this deeper water province and is characterised by species of Ogmoconchella, Bairdiac,vpri.~?, Cytherclla and Ci,therelloidea. The ogmoconchids appear to have migrated west through the Tethys and t o have moved northwards into the European Province where they survived until the basal Toarcian when possibly they were unable t o compete with the rapidly evolving cytheraceans. A parallel with the ornate tethyan ogmoconchids is the ornate group of hairdiids that were common in the Trias of this province. For example, Ptychobairdiu schaubergeri is found both in the Kleckenmergel of Austria (Kollmann 1963) and in the Pliensbachian of Djebel Zaghouan,
237 Tunisia while Lord & Moorley (1974a & b) record highly ornate bairdiids: B. hahni and B. a.telfingencnsi.s from the Pliensbachian of SW-Germany. Rarely do these bairdiids occur outside the Tethyan Province although fragments of Ptj.chobairdia sp. have been recorded from the Toarcian of England (Bate & Coleman 1975). Barbieri (1964b) describes the only ostracods t o come from the centre of the province and this Sicilian fauna is composed entirely of smooth-shelled ostracods. All the available cvidence indicates, therefore, that the evolution of the cytheracean ostracods was initiated in the warmer epicontinental seas bordering the Tethys. Middle Jurassic - Aalenian to Callovian. A rather poor fauna is recorded by Barbieri ( 1 964b) from the Bajocian of Sicily and by Donze (1962) from the Bathonian - Callovian of southern France where only three species: Schuleridra caidata, Proc.vtheridea martini and Oligoc.vtherei.7 gauthieri are recorded. The French fauna is not particularly representative of deep water conditions and is considered t o be marginal between the European and Tethyan Provinces. If this is so it would indicate that the northern boundary of the Tethys had receded since the Lower Jurassic. Eastwards through Europe the Middle Jurassic faunas described all lie outside the Tethyan Province - this is true as far east as the Dnieper-Don Depression where Permjakova 11969. 1970) describes a fauna containing at least 5 species of Gl.yptoc,i.thcre. Further east in Uzbekistan the faunas described byMasumov (1973) belong to an epicontinental sea province that must have had continuity with the European Province and at the same time lay t o the north of the Tethys. Perhaps the Middle t o Upper Jurassic faunas of the Middle East should more accurately belong t o the Tethyan Province - in which case the North African Province could be dispensed with entirely. This will become more apparent as research on these faunas continues. Upp-r Jurassic - Oxfordiaii to Tithonian. In the Upper Jurassic the Tethys moved into the opening Atlantic Basin and appears t o have taken with it the important ostracod family. the Schulerideidae. The migration route would not be through the deeper water regions b u t through the more shallow waters of the margins. The genus Procj~tlzeritleasensustricto, first described from the Callovian of western North America (Peterson 1954), is know to be present in the Callovian of Tanzania (Bate 1975), the Callovian and Oxfordian of southern France (Donze 1962) and in the Oxfordian of Switzerland (Oertli 1959) - as well as in western North America and Canada. Because of this wide distribution pattern a simple westward migration by way of the Tethys appears to have been extremely unlikely. Two ostracod faunas described by Oertli (1972) from DSDP wells in the Atlantic [site 100 off the Bahamas :uid site 105 off Cape Hatteras] have been compared with the deeper water Tethyan faunas described by Oertli (1967) from Italy and with the Middle to Upper Jurassic ostracods described from SE. France by Donze (1 962). Only Rairdia (Akidobairdiai farinacciae correlates the Upper Jurassic of Italy directly with DSDP site 100 but the faunas are, nevertheless clearly representative of a deeper water Tethyan Province. Similarities between species of Ac,roc,v?hrre figured by Oertli (1972) and by Pokorny (1973) are not conclusive and in any case the Czechoslovakian fauna is considered to belong more correctly with the European Province. The close proximity of DSDP sites 100 & 105 to the American Province is remarkable for the complete contrast between the faunas. This contrast is considered t o be due entirely t o differences in water depth - a clear indication that Schiileridea could not have migrated westwards in the deeper water regions of the Tethys.
238
Ostracoils such ;is Mucrotlc~ritiriuand Gulliuccj thc~ridcu,so typical of Upper Jurassic sediments of tlie European Province are absent from deeper waters of the Tethys where Donze ( 1 962) describes a fauna. from southern France. dominated by either smooth-shelled or alate species. \Vliercus a zonal scheme for the epicontinental sea facies of Europe and North America is possible it will be some time before this will be so for the deeper water Tethys where smootlishelleti ostracody prcdoniinate and where there is a considerable decline in numbers. The North African Province. the area covered by this province is considered to extend over that part of North Africa lying south of Tunisia and taking i n Egypt arid the Middle East. There is no published information available and studies on the ostracod faunas are at a very preliminary stage. As a result it may be necessary to revise the decision taken here to separately identify this province. Lower Jurassic Hettanpian to Toarcian. No Lower Jurassic ostracods have been described as those ohtainecl from Djebel Zaghoudn are considered to belong to the Tethyan Province. Middle to Upper Jurassic Aalenian to Tithonian. This i n t e n d is grouped together simply hccausc the Jurassic matcrial examined has been rather loose13 dated as Callovian/Oxfordian. I n Jordan. ‘4rnic i~tlic,rit/c.uand Afi.ocj~thcriticaand in southern Israel, Afi.vc>~thcrit/eu. are represeiitell amongst an ostracoti fauna typical of a shallow marine environment. Both 4rnic~~thc~riticu and ilfi.oc.i.thc.uii1c.a were originally described from the Callovian of Tanzania (Bate 1975) and have since been observed i n the Jurassic of Kenya aiid Somalia. Thus these two genera extend northwards as far as North Africa but do not appear to cross the barrier of the Tethys. It is bccnusc of this that a decision was taken to separately identify the North African fauna as being ~
~
distinct from either the Tethyan to the north or the East African to thesouth. but havingconnections with both. The East African Province. The Southern hemisphere ostracod faunas during the Jurassic and. indeed for the Mesozoic in general, are qiiite distinct from those of the northern hemisphere. Ccrtninly many ostracod genera are cosmopolitan in their distribution but the essential composition of the faunas is different. Whereas i n the European Province three families, the Progonocytheridae, the Protocytherih e a n d the Schulerideitlac. play an important role in stratigraphical correlation. here only the Progoiiocytheridae assumes any importance the Schulerideiilae and Protocytheridae are reprcsentccl but i n a subsidiary role. In the East African Province the most important penus is .Maiirngat~/luwhich is very closely related t o Progonocj,thcrc of the European Province. The latter plays a very insignificant part i n Jurassic stratigraph) whereas Majungaellu may be used for correlation between East Africa. India, Madagascar and South Africa. Indeed, five zonal species of ,Vlaj/ingaellahave been recognised in the Jurassic (Bate 1975): IM. mund~rla Callovian; .21. o.yfort/iariu Oxfortlian: .M. kirnriieritlgiana - Lower Kimmeridge: .M. pracperfiirutu ~
~~
~
Middle or Upper Kimmeridge (precise age uncertain) and ‘44. perforatu Tithonian. Additionranges u p into the Cretaceous and is known from Africa, South America [see ally :llujiir~guc~/lu .Yoivcj~thcrrRossi tle Garcia 19721 and Australia. Thus the presence of this genus. or of one of the very closely related subgenera, readily identifies a southern hemisphere fauna as distinct from one from the north. The American Province. The buried Upper Jurassic of the Gulf Coast and of the SE-Atlantic coast of the USA together form the American Province which in faunal terms is characterised by
239
the dominance of Schukridea, Paraschuleridea and Hutsonia. This is a unique, shallow water marine fauna that is more easily correlated with the western North American Jurassic than with the faunas figured by Oertli (1972) from the DSDP well sites in the Atlantic. Lower Jurassic - Hettangian to Toarcian. Absent. Middle Jurassic - Aalenian to Callovian. Absent. Upper Jurassic - Oxfordian to Tithonian. The age dating of this sequence is not precisely known and probably not all stages are present. The characteristic feature of this province is the absolute importance of the the Schulerideidae. Of the large number of rapidly evolving genera that appeared in Europe during the Jurassic only the genus Schulcridea in association with Asciocj,there and Parascliirli~rideaappears to have developed here. European genera such as Fuhrbrrgiella and Aeudohutsorzia present i n the Upper Jurassic of western North America must have arrived there via Greenland and one could postt~latc that the southward extension of the Tethys with the opening of the Atlantic was initially a deep water transgression that was unsuited to the migration of the European genera. I n this context a detailed study of the eastern Canada Jurassic faunas should be very interesting as it would appear that thcre was little or no continuity between there and the American Province further south. The absence of Lower and Middle Jurassic marine sediments in the southern North Atlantic is taken t o indicate that marine conditions were totally dependent upon the opening of the Atlantic and the introduction of the Tethys. The marine Upper Jurassic ostracods of Louisiana and Arkansas [Swain 1946, and Swartz & Swain 19461 and from North Carolina [Swain 1952 and Swain & Brown 19721 are considered to represent a rather restricted, shallow water marine environment that has little direct correlation with the marine Provinces of Europe and North Africa. Unfortunately we have n o information concerning the Jurassic ostracods of Mexico but it is possible that some connection west with a Pacific Province might have been in existence during the Upper Jurassic even though the essential marine transgression of the southern North Atlantic came from the North.
CONCLUSIONS The distribution of the Jurassic Ostracoda was examined in the context of a re-positioning of the continents as they were in late Jurassic times. With the Atlantic closed during the Lower Jurassic, marine conditions entering from the east reached only as far west as eastern Canada. The central part of this transgression - the Tethys - has been identified as a deeper water Tethyan Province characterised by a rather restricted fauna of smooth-shelled ostracods. The primary role of the Tethys appears to have been t o act as a north-south barrier to ostracod migration. Possibly this was because the newly evolving ostracods were unable to tolerate the deeper [cooler?] waters of the Tethys. Bordering the Tethyan Province epicontinental seas spread north over previously barren Triassic landmasses and, following on from the extinction of most of the Palaeozoic lineages, a unique situation arose that led t o rapid speciation and subsequent evolution of many new lineages. Although there is some evidence that similar conditions existed eastwards into Asia, the European Province is considered t o have been one of the major sites of ostracod evolution
240 and certainly the most important in terms of the present study. Elements of the European fauna migrated north to reach western North America during the Upper Jurassic and likewise migrated east into Asia. I n the Lower Jurassic of the European Province the important ostracods are the ogmoconcliids and the Kiii/wliti?lIa Ekti.phocj.therc, lineage of the Protocytheridae. The cytherdcean ostracods develop rapidly towards the end of the Lower Jurassic and through the Middle and the Upper Jurassic the stratigraphically important ostracods belong to the Progonocytheridae and to the Schulerideitlae. Fresh to brackish-water ostracods of the Limnocytheridae become important in the continental deposits at the close of the Jurassic where the genus Cypridea achieves a worldwide distribution. The development of many short ranging species of this genus led to its recognition as a zonal fossil h y the English geologist Forbes as early as 185 1. The American Province does not appear to have had an epicontinental sea connection with the European Province and as the Tethys was not an effective migratory route only a very restricted fnuna developed of which the most important is the Schulerideidae. Indeed the common occurence of species of Schiileridca, Pamschirlcridea and of Asciocj,there are a characteristic feature of this province. The North African Province has been little studied to date and is presently recognised as containing elements of the East African fauna associated with the cosmopolitan genus C.vthercllu. It is too early i n the investigation of this province to indicate any positive trends in ostracod dcvclopment. The East African Province is introduced into this study solely to draw attention t o the diffcrcnces that exist between the northern and the southern hemisphere faunas and t o the restricting effect 011 migration of the Tethys. Although inany ostracods such as Cj.thcvellu and Fairtliu have ii cosmopolitan distribution those ostracods that had their origins in the Jurassic were essentially restricted to a migration route within their respective hemisphere. AnzicFtherideu and A froci,thcritiea for example extend north into the North African Province but d o not extend further. The genus .Ilajitrigaelh from East Africa appears to achieve a world-wide distribution in thc southern hemisphere hut is not known in the north. The Tethys did, however, permit some migration Proc,vtheridea present in North America and East Africa appears t o be associated in Europe with the marginal facies of the Tethys and the Lower Jurassic ogmoconchids were certainly brought into the European Province by the Tethys. Finally. three Lower Jurassic genera of the European Province: Proc.vthcrura, Enc.vthc,rirru and C),theropteroii appear much later in the Middle t o Upper Jurassic of East Africa (Bate 1975) indicating that some north-south migration has been possible. ~
241 REFERENCES .ASDERSOU, F. U , 1971. In ANDERSON. F. W. & BAZLEY, R. ,A. B. The Purheck Beds of the W'edld ( E n g l a n d ) . Rirll. Geul. Sirrl'. G , B. no. 3 4 , I 7 4 p p , 23 pls. BARBIERI, F . 1964a. lliiizgurcliu l i r ' b l i ~ an u o v o Ostracode del Domeriano. /.'Arcnru Purnzensc, vol. 35. supl. 1 , p. 3-7. .___.._ 1964h. _ _ . hlicropnleontologia del Lias e Dogper del por2o Ragusa 1 (Sicilia). R ~ Y/to/. . Paiconi., vol. 70, p. 709-830, pli. 5 1-66. r e 8. B.ISSIOUNI. M . E. A . A , 1974. P a r a r i u t u c ~ ~ i h c 11. iOstracodd) a u s dem Zzi!raurn Oherjura bis Unterkreide (Kimmeridgium his Albium) von N'cstcuropa. Gcul. Jb.. vol. 1 7 , p . 3-1 I I , I 3 pls. B.\TE. R. H. 1965. Middle Jurassic Ostracoda from the Grey Limestone Series, Yorkshire. Bill/. Rr. .llus. nut. Hirt. geoi., voi. I I , p. 73.133, pls. 1-21. . - .- 1 9 6 7 . The Bathonian Upper Estuarine Series of Eastern England. Pt. I : Ostracoda. Bull Br. .Mus. t i U I . / f i s t . geul., vol. 1 4 , p . 21-66, 22 PIS. . . _ _ .1975. ___ Ostracods __ trom Callovian to Tithonian sediment\ o f Tdnzania. East ,Africa. Biiii. Br. .Zlu\. noi. /list. pevl., vol. 2 6 , p. 161.223, pls. 1-14. . . _ _ .& _COLEMAN. ___ B. t. 1975. Cpper Lias Ostraciida from Rutland and Huntingdonshire. Bidi. G i d S u r b . G,B. no. 55. p. 1-42, 1 5 pls. _ _ _ - _ _ . . & R O B I N S O NE.(eds.). , A Srratigruphicai I n d e x o f British Osrruc~tdu. See1 House Press (in prepn.). BROOKE, \l. M.& B R 4 U U , W. K. 1972. Biostratigraphy a n t i microfaunan of the Jurassic System of Saskatchewan. D e p t . Miti. Res. Sushatchr~waii Report., 1 6 1 , p. 1-83, 26 pls. DOUZE, P. 1962. Contribution ii ]'etude pal&mtologiique d e I'Oxfcirtlien Supt.rieur d e Trept ( I s h e ) .
___
pls. 9-1 1. ..... ....1966. Ortracodes de I'Hettangien entre ,,\tibends et Privas iArd6che). Tvuv. /.ah. Gc%i/. /,,l'UJl, .\'. s., 110. 13, p. 1 2 l - l 3 9 , p 1 ? , 5 - 7 . DRFXLER, I . 1958. Foraminiferen iind Ostracciden aus dern Liasoc von Siebeldingen/Pfdk. Geol. Jh., vol. 75. p. 475.554. pls. 20-27.
GERRY, E . 8, OERTLI, H. J . 1967. Bisuicoc.vpris ' triarsica n . sp. (Crust., Ostrac.) from Israel. Biiil. C e n t r e R c t h . Puii-S!VPA.. vol. 1 , p. 375-381, 1 pl. HklhlDACH, F. F. 1971. Z u r Gliederung limnischhrackischcr sedimcnte des portirgiesischen Ohcrjura ( o h . Callovien - Kimmeridge) m i t Hilfe y o n Ostrakoden. .V. J b . Geol. Paluoi!:. .Iiir.. vo1. 1 I , p , 645-662. KILENYI, T. 1. 1969. The Ortrac(ido cif the Dorsel Kimmeridge Clay. P i i i u e o ~ i t ~ ~vi d~ .~ 1p2~, ~p ,. 1121 GO, PIS. 23-3 1. KLINGLER, W.& UEUWEILER, F. 1959. Lcitende Ostracodcn airs den1 deutscticn LiasA. G e o L J h . . vol. 76. p. 373-410, 6 pls. KOLLLIAUN, K. 1963. 0rtr:icoden :ius tlcr alpinen l r i a r 2. Weiterc Bairtliid;iz. Jti. (;col., v o l . 106, 1). 121-203, pls. 1-11.
LORD, :A. & XfOORLEY, A . 1 9 7 4 a On Biiirdin iiiiiiiii Lord & \loorley sp. n o v . Stcrrw .Atlri.! Osrrac~idSiiell,r, v o l . 2 . p. 1-4. ____ 1974h. _._. On _Boiriliaaselringenensis _ Lord & M o o r l c y S ~ I n, o v . Stereo .1tius O.rtr(iL,oiI S l i e i l i , vol. 2, p , 5-8. _ _ _1_ 9 7. 4. ~ ._ On _ 0 , ~_ n i o c o n c h aiinihii Lord & 4loorlzy s p . nov. .S'rcrro .4 tlar Osirai,iid Siiellr. vol. 2 , p. 9-16. MALZ. H. 1966. Zur Kenntni.. einigcr 0str;icoricii: W e n der Gattunpcn Kiiihclincllu u n d Priic~ ( I i i i i c r i d e u .Sciick. l L ? t l i , , vol. 4 7 , p , 385-404, 111s. 48-49. MASUh4OV. A . S. 1973. Jiirassic iJstrac;idisof Lrhekistaii. 157 p p , , 14 pls. Tarhkcnt [in Russian], MICHELSEN. 0.1975. Lowcr Jiirassic hiostratigraphy and ostracods of the Danish Embaynient. D~rziriarl~, gcol. iirid?rs., n u 1 0 4 , 287 pp., 4 1 pls. OERTLI, H. J. 1957. Ostracodes tlu Jurassiqtie SupCrieur d u Bdssin clc Paris ( S o n d a g c Vernon I ) . R e v . I i i s t . Frurlc. PCtrole, v o l . 1 2 , p . 6476 9 5 , pls. 1-7. __ _._1959. _ . Malrn-Ostrakodcn aus tleni rchweiierischcn Juragchirgc. .ilfini. Soc. Hriv. S r i . .\ur., vol. 8 3 , p. 1-44, pls. 1-8. . -1972. 22 Jurassic ostracode? of DSDP Lcg I I (nitcs 1 0 0 antl 105) - Preliminary account. p. 645-6. pls. 1-5. I n , HOLLISTER, C. D., FWING. J. I., e t al., Iriiliai R e p o r t s o f t h c D c c p Sea Drilliiiy Projecl, vol. 1 I , O W E S , 11. G . 1976. Continental ilisplaccment and expansion of thc Earth diiring the hlecoznic antl Cenoioic. Phil. Trails. roi'. S ~ J C Scr. .. A, vol. 281, p. 223-291. PFRMJAKOVA, M.N, 1969. New specicc of Ostrac i i d a from the Bajocian Dcpmits o f the Diiieper-Don Depression. Puleorit. Shorilik., no. 6 , p. 34-38, I 111. [ i n Russian]. 1970. OFtracoda of the genus G/,sptocythere from the Midrllc Jurassic Deposit? o f the Dniepcr-Don Depression. Paieont. S b o r n i k . , no. 7 , p . 61-67, 1 pl. lin Russian]. POKORNY, V. 1973. T h e Ostracoda of t h e Klentnice Formation (Tithonian?) Czechoslovakia. A c d . riahiad. Ces. A k u d . ved. YO]. 4 0 , P. 1-107, ?o plS. ROSS1 DE GARCIA, F. 1372. Ostracoda. In: MALMUMIAN, N., MASIUK, V. X ROSS1 D E GARCIA, E. Micrtifbsilcs del CretPcico superior d e 13 pcrforacibn SC-I, provincia ric Santa Cruz, Argentina. Revia. Asoc. @iil. Arcyenr., vol. 2 7 , p . 265-272, I pl. SWAIN, F . M. 1946. Upper Jurassic Octracoila from the Cotton Valley G r o u p in Northern Louisiana: the genus Hutsonia. J . Paleont., YOI. 20, p. 119-129, PIS. 20-21. .___ 1952. __ Ostracoda _ _ _ from wells in North Carolina: Part 2 . Mesozoic Ostracoda. U.S. Geol. Suri'. Prof. Pup., 234-B. p. 59-93. pls. R,9.
__
_____
________
242 REFERENCES SWAIN & BROWN, P. M. 1972. Lower Cretaceous, Jurassic ( ? ) and Triassic Ostracoda f r o m the Atlantic Coastal Region. U. S. Geol. Surv. Prof. Pup, 7 9 5 , p . 1-55, 9 pls. SWARTZ, F. M , & SWAIK, F. M. 1946. Ostracoda f r o m the Upper Jurassic Cotton Valley Group of LouiTiana and Arkansas. J. Puleont., vol. 20, p. 362-373, pls. 5 2 , 53.
WHATLEY, R . C. 1970. Scottish Callovian and Oxfordian Ostracoda. Buii. Br. Mus. n a f . H i s i . geui., vol. 1 9 , p. 297-358, 9 pls. WfENHOLZ, E. 1967. Neue Ostracodcn aus den1 n
243
Discussion Dr. W. Manspeizer: Our (Manspeizer, Cousminer, and Puffer) work over the past few years disagrees, in part, with the reconstruction presented in this paper. Our studies and those of Prof. Ager, Chris Kendell, Ian Evans, Bill Ryan, etc. generally show that the High Atlas contains a marine fauna, and was a deep-water basin that became detached from the African Platform in the early Jurassic. Similar marine conditions prevailed throughout most of the middle and Late Jurassic. Our studies in easternmost Morocco show that the Muschelkalk Sea emanating from the Tethys Sea to the north and east, transgressed the region near the Algerian border in the middle Jurassic and probably extended into the Rif Province of western Morocco by Late Triassic and Jurassic time. These waters from Tethys may well have served as the source for the salt in the Argo Salt (early Jurassic) on the Scotian Shelf, and for the marine transgression associated with the acritarchs and tasminids of the Waterstone Formation of the Central Midlands of England. In short, stratigraphic data from Morocco indicates that a marine Tethys transgression extended into the Atlantic region by the late Triassic. Bate: Dr. Manspeizer quite rightly indicates that Morocco belonged to a Tethyan Province during the Jurassic-this agrees with my paper as I have placed Tunisia firmly in this province for the Jurassic. What must have confused Dr. Manspeizer was the use of the term North African Province. This term was used for that part of North Africa extending from Libya to the Middle East. Even so I have suggested on the second page of my paper that the eastablishment of this province”is rather tenuous and may have to be merged with the Tethyan Province at a later date”. With respect to Dr. Mansseizer’s comment concerning the Upper Trias in Morocco I see nothing in my paper that would disagree with this. All I would suggest is that Morocco was situated on the most easterly extension of the deeper-water Tethyan transgression with shallower marine conditions extending farther east. Dr. F. M. Swain: There may be a little too much emphasis on provincialism in this discussion and not enough on the possibilities of local environmental variations and possible endemism. The Upper Jurassic ostracodes from the mid-Atlantic and Gulf of Mexico regions have certain similarities, particularly in the Schulerideas, Hutsonias, and Fabanellas. In the Scotian Shelf area there are also similarities to the preceding, the three areas showing gradational relations. But a difference occurs between these late Jurassic ostracode faunas and those of the Sundance Sea of the western interior U. s. There we find ostracodes that are more like those from western Europe than they are like Gulf and Atlan tic. Bate: I would say that some relationship between the Gulf Coast, Mid-Atlantic, and Scotian areas is to be expected through migration via shallow epicontinental seas. The deeper water Tethys appears to have been less helpful in this respect. Dr. I. G. Sohn: I believe I found the same ornate bairdiids (ie. Ptychobairdia) in the Triassic of Nevada, and Kolmann’s from Austria are Triassic also.
244
B a t e : T h e i n t i m a t i o n from D r . Sohn t h a t h e h a s found o r n a t e b a i r d i i d s from t h e T r i a s of Nevada s i m i l a r t o forms d e s c r i bed by Kolmann from A u s t r i a i s of i n t e r e s t b e c a u s e it p a r a l l e l s t h e s o r t o f d i s t r i b u t i o n I have mentioned f o r t h e J u r a s s i c genus P r o c y t h e r i d e a sensu s t r i c t o .
245 Cretaceous Ostracoda of t h e North A t l a n t i c Basin JOHN I V .
NEALE
Department o f Geology, The University, Hull, Yorkshire, England
Abstract F o r m o s t o f Lower C r e t a c e o u s t i m e , N o r t h A t l a n t i c o s t r a c o d f a u n a s are o n l y well-developed knowledge i s o n e - s i d e d .
i n t h e E a s t e r n area and o u r
Varine t r a n s g r e s s i o n and t h e
e x t e n s i o n o f d e p o s i t i o n i n t h e A l b i a n and Upper C r e t a c e o u s gave e x c e l l e n t f a u n a s on b o t h s i d e s o f t h e A t l a n t i c and t h e Basin appears 'two-sided'
.
Besides being stratigraphically
u s e f u l , t h e o s t r a c o d s p r o v i d e ecological i n f o r m a t i o n on f a c t o r s such as s a l i n i t y , t e r p e r a t u r e and m i g r a t i o n . Introduction D u r i n g C r e t a c e o u s t i m e s t h e N o r t h A t l a n t i c B a s i n was r e l a , t i v e l y s m a l l , t h e S o u t h A t l a n t i c R i f t w a s o n l y j u s t commencing a n d m o s t o f o u r i n f o r m a t i o n is d e r i v e d f r o m l a n d a r e a s s u r r o u n d i n g the Basin.
Lower a n d U p p e r C r e t a c e o u s b o t h h a v e d i v e r s e
o s t r a c o d f a u n a s which are s t r a t i p r a p h i c a l l y u s e f u l .
They a l s o
o f t e n p r o v i d e m u c h - a d d i t i o n a l i n f o r m a t i o n s u c h as c h a n g e s from non-marine deltaic/.mudf.lat t o m a r i n e e n v i r o n m e n t s which can hav'e i m p o r t a n t a p p l i c a t i o n s i n o i l e x p l o r a t i o n .
Besides
s a l i n i t y , t h e s e s m a l l c r u s t a c e a n s may a l s o g i v e i n s i g h t s i n t o o t h e r p a r a m e t e r s s u c h as t e m p e r a t u r e , geography.
r e l a t i v e d e p t h and bio-
I n t h e North A t l a n t i c area t h e development of
o s t r a c o d - b e a r i n g s e d i m e n t s was n o t i c e a b l y o n e - s i d e d i n t h e Lower C r e t a c e o u s .
With t h e A l b i a n a n d Upper C r e t a c e o u s , a b i g
marine expansion produced a two-sided
B a s i n and f a u n a s o f t h e s e
a g e s a r e w e l l known i n W e s t e r n a r e a s .
This review d e a l s f i r s t
w i t h Lower C r e t a c e o u s a n d t h e n w i t h t h e U p p e r C r e t a c e o u s w o r k i n g f r o m e a s t t o west i n e a c h c a s e . Lower C r e t a c e o u s N o n - M a r i n e F a u n a s In l a t e J u r a s s i c - E a r l y C r e t a c e o u s times non-marine d e p o s i t i o n was a n o t a b l e f e a t u r e i n many a r e a s .
Fuch ' f e a l d e n ' f a c i e s i s
c h a r a c t e r i s e d b y r a p i d l a t e r a l a n d v e r t i c a l v a r i a t i o n s o f muds
246
1Land
0Sea
Lower Cretaceous Cypridea Faunas Fig.1 EARLY CRETACEOUS - Distribution of Land, Sea and Non-marine Ostracod Faunas
247
and s a n d s and is o f t e n of c o n s i d e r a b l e i n t e r e s t t o t h e p e t r o leum g e o l o g i s t an d r e s e r v o i r e n g i n e e r .
Formerly regarded as
d e l t a i c s e d i m e n t a t i o n , t h e most r e c e n t t h e o r y r e g a r d s t h e
c l a s s i c Wealden e n v i r o n m e n t as a v a r i a b l e s a l i n i t y mudplain w i t h o v e r l o a d e d streams p e r i o d i c a l l y p r o v i d i n g a b r a i d e d s a n d u r env ironrnent . The d e p o s i t s y i e l d e x c e l l e n t o s t r a c o d f a u n a s , e s p e c i a l l y t h e g e n u s C y p r i d e a , and t h e f u l l e s t development o c c u r s i n s o u t h e r n B r i t a i n where t h e b e d s r a n g e from L a t e J u r a s s i c up t o and i n c l u d i n g t h e Barremian.
Based on t h e s e
' b e a k e d ' s p e c i e s , Anderson (1973) i n a m a s t e r l y r e v i e w h a s been a b l e t o r e c o g n i s e two a s s e m b l a g e s o f J u r a s s i c a g e and e i g h t o f C r e t a c e o u s a g e w h i c h e n a b l e s c o r r e l a t i o n s t o b e made i n t h e NW European area ( F i g . 2 ) .
I n N . Germany t h e ' P ' e a l d e n '
Beds o n l y
i n c l u d e t h e f i r s t two C r e t a c e o u s a s s e m b l a g e s a n d a r e o v e r l a i n by m a r i n e M i d d l e V a l a n g i n i a n .
similar.
The s i t u a t i o n i n H o l l a n d is
I n t h e Swiss J u r a t h e f i r s t Cretaceous assemblage is
p r e s e n t b u t t h e b u l k o f t h e non-marine b e d s are J u r a s s i c ,
as i n
t h e case o f t h e F r e n c h J u r a a n d t h e P a r i s B a s i n w h e r e C r e t a c e o u s non-marine
faunas have not been a t t e s t e d .
The D a n i s h
I s l a n d o f Bornholm y i e l d s b o t h J u r a s s i c and C r e t a c e o u s assemb-
lages of which t h e h i g h e s t from t h e J y d e g a a r d Formation sugg e s t s t h e second Cretaceous assemblage.
Outside Britain the
m o s t c o m p l e t e d e v e l o p m e n t s a p p e a r t o b e t h o s e of t h e I b e r i a n p e n i n s u l a where t h e e a r l i e r J u r a s s i c and C r e t a c e o u s non-marine a s s e m b l a g e s h a v e b e e n f o u n d a t a number o f p l a c e s .
West o f
Logrona, n o r t h o f Burgos and also near Cuenca, t h e h i g h e s t
C r e t a c e o u s a s s e m b l a g e has b e e n f o u n d . The g e n u s C y p r i d e a is s u r p r i s i n g l y w i d e s p r e a d and Anderson ( 1 9 7 3 ) h a s s u g g e s t e d t h a t i t was c l i m a t i c a l l y c o n t r o l l e d a n d o c c u p i e d a z o n e on e i t h e r s i d e o f t h e C r e t a c e o u s e q u a t o r whose
c l i m a t e was s i m i l a r t o t h a t o f t h e p r e s e n t d a y M e d i t e r r a n e a n . T h e m e a n s o f d i s p e r s a l o f t h i s g r o u p of o s t r a c o d s , w h i c h o c c u r i n d i s j u n c t b a s i n s , h a s so f a r n o t b e e n s a t i s f a c t o r i l y explained.
A l t h o u g h t h e f o r m s show a g e n e r a l s i m i l a r i t y , d o u b t -
l e s s d u e t o t h e l i m i t e d n u m b e r of u s a b l e t a x o n o m i c c h a r a c t e r s , indigenous s p e c i e s are developed.
Thus w h i l e c o r r e l a t i o n can
b e made b e t w e e n Gabon a n d NE B r a z i l , a n d e m p h a s i s e s t h e c l o s e j u x t a p o s i t i o n of t h o s e t w o areas i n e a r l y C r e t a c e o u s times ( s e e
B e r t e l s , t h i s symposium), t h e r e is n o t a s i n g l e s p e c i e s i n
248
ENGLAND
SPAIN
__
WEALD CLAY
1
Lower Barremian
Hauterivian
", '
lL 5
Lower
WEALD CLAY
Hauteriviar
TUNBRIDGE WELLS SAND
Upper
a
Valanginian
WADHURST CLAY
1
ij
I,
I'
WADHURST Va la n g in ia n
CLAY
I
1
Lower
L volanginior
Fig 2
Europe - Lower Cretaceous Non-Marine Ostracod Faunas (Anderson 1973 [pars] redrawn)
P l a t e 1. P a i r e d s t e r e o s c o p i c p h o t o g r a p h s of n o n - m a r i n e L o w e r C r e t a c e o u s o s t r a c o d s from s o u t h e r n England. 1. C y p r i d e a g r a n u l a t a f a s c i c u l a t a J o n e s . LV x 3 5 . 2 . C y p r i d e a s e t i n a A n d e r s o n . RV x n r i d e a p a u l s g r o v e n s i s ( A n d e r s o n ) . LV x 5 2 . 4 . C y p r i d e a a c u l e a t a J o n e s . LV x 5 0 . 5 . C y p r i d e a d o r s i s p i n a t a ( A n d e r s o n ) . LV x 6 0 . 6 . C y p r i d e a c l a v a t a A n d e r s o n . LV x 5 2 . 7 . C y p r i d e a s p i n i g e r a ( J . d e C . S o w e r b y ) . LV x 5 7 . 8. Cypridea t e n u i s A n d e r s o n . C a r a p a c e from r i g h t x 5 0 .
249
PLATE 1
250
common w i t h t h e E u r o p e a n a r e a . I n America, s p e c i e s o f C y p r i d e a and H u t s o n i a . have been found i n t h e h l i d d l e A t l a n t i c S t a t e s i n U n i t H o f S w a i n a n d Brown 0972) which is r e g a r d e d as p o s s i b l y J u r a s s i c a t t h e b a s e r a n g i n g u p
t o e a r l y Comanchean (
Aptian).
Marine s p e c i e s a l s o o c c u r i n
U n i t H, w h i c h i s p r e d o m i n a n t l y m a r i n e i n e a s t e r n N o r t h C a r o l i n a , becoming i n c r e a s i n g l y non-marine except i n t h e extreme east.
i n V i r g i n i a and New J e r s e y
I n t h e same a r e a U n i t G a l s o c o n -
t a i n s C y p r i d e a b u t i s e s s e n t i a l l y m a r i n e ( s e e u n d e r ALBIAN). C y p r i d e a f a u n a s are a l s o found i n Oklahoma, U t a h , I d a h o , Kyoming, M o n t a n a , S o u t h D a k o t a a n d A l b e r t a b u t a r e t o o f a r west t o be legitimately considered i n t h i s contribution. From t i m e t o t i m e , i n t e r c a l a t i o n s o c c u r o f o s t r a c o d s s u g g e s t i v e of h i g h e r s a l i n i t i e s t h a n t h o s e of t h e normal C y p r i d e a faunas.
A n d e r s o n c a l l s t h e s e 'S' P h a s e a n d t h e y h a v e b e e n
variously c a l l e d marine or quasi-marine bands.
The close
a s s o c i a t i o n o f s p e c i e s s u c h as F a b a n e l l a b o l o n i e n s i s , F . a n s a t a and M a n t e l l i a n a p u r b e c k e n s i s w i t h e v a p o r i t e d e p o s i t s h a s l e d t o t h e s u g g e s t i o n t h a t t h e y may h a v e f l o u r i s h e d i n h y p e r s a l i n e
waters.
K i l e n y i a n d A l l e n ( 1 9 6 8 ) who made a d e t a i l e d s t u d y o f
b a n d s i n t h e l o w e r p a r t o f t h e Weald C l a y o f S u s s e x a n d S u r r e y concluded on t h e b a s i s o f S c h u l e r i d e a ( E o s c h u l e r i d e a ) wealden-
e, Hutsonia
c a p e l e n s i s , Ammobaculites and C i r r i p e d e s t h a t t h e
s a l i n i t y i n t h a t s e d i m e n t was p o l y h a l i n e / e u h a l i n e o n o c c a s i o n . The s p e c i a l i s e d f a u n a s o f t h e s e ' S ' p h a s e b a n d s h a s s o f a r proved of l i t t l e u s e i n c o r r e l a t i o n , Lower C r e t a c e o u s M a r i n e F a u n a s Marine o s t r a c o d f a u n a s i n n o r t h e r n Europe have r e c e n t l y been r e v i e w e d by Neale ( 1 9 7 3 ) who g i v e s a f u l l b i b l i o g r a p h y . 1 . BERRIASIAN
P r o v i n c i a l i s m w a s most m a r k e d d u r i n g t h i s s t a g e w i t h c l e a r s e p a r a t i o n o f t h e c o l d e r , n o r t h e r l y f a u n a s o f B r i t a i n a n d Denm a r k a n d t h o s e o f t h e warmer a r e a s f u r t h e r south.
The B r i t i s h
f a u n a is c h a r a c t e r i s e d by G a l l i a e c y t h e r i d e a t e r e s ( u s e d as t h e s t a g e index f o s s i l by Christensen 1074), Mandelstamia s e x t i , S c h u l e r i d e a j u d d i , P a r a c y p r i s caerulea, Cytheropterina t r i e b e l i and P o n t o c y p r i s f e l i x .
Notable absentees are representatives
251
of t h e warm water genus C y t h e r e l l o i d e a and C y t h e r e l l a , P r o t o c y t h e r e and C y t h e r e i s , w h i l s t G a l l i a e c y t h e r i d e a and V a n d e l s t a -
mia l i n g e r
on from t h e J u r a s s i c .
I n t h e V o c o n t i a n T r o u g h o f S . F r a n c e some 10'
further south,
w h i c h i n c l u d e s t h e t y p e a r e a o f B e r r i a s o n t h e west, B e r r i a s i a n faunas have a very d i f f e r e n t a s p e c t w i t h C y t h e r e l l a , CytherelI-
loidea
P r o t o c y t h e r e , C y t h e r e i s and A c r o c y t h e r e a l r e a d y e s t a b -
l i s h e d , d i f f e r e n t s p e c i e s o f O r t h o n o t a c y t h e r e , P a r a c y p r i s and S c h u l e r i d e a , and i n d i g e n o u s g e n e r a s u c h a s K e n t r o d i c t y o c y t h e r e and Raymoorea.
Donze ( 1 9 7 1 ) h a s b e e n a b l e t o r e c o g n i s e
dif-
f e r e n c e s i n p a s s i n g from t h e m a r g i n s ( f a c i e s s e m i - e m e r s i f ) towards t h e deeper p a r t s of t h e b a s i n ( f a c i e s n e r i t i q u e ) .
The l a t t e r is c h a r a c t e r i s e d b y t h e d i s a p p e a r a n c e o f g e n e r a s u c h a s E u r y i t y c y t h e r e , Exophthalmocythere, Orthonotacythere, Quasiherrnanites, Kentrodictyocythere,
Schuleridea. 'Clithrocytheridea'
and o t h e r s , a marked i n c r e a s e i n t h e p r o p o r t i o n o f P a r a c y p r i s , C y t h e r e l l o i d e a , C y t h e r e l l a and B a i r d i a , and t h e p r e s e n c e o f Cypridina, Polycope, Cardobairdia, Annosacytherc, Hemicytherura.
Raymoorea and
The m a r g i n a l f a c i e s i n c l u d e s occasional
C y p r i d e a , F a b a n e l l a , Limnocythere and S c a b r i c u l o c y p r i s .
In
Portugal t h e Berriasian starts with a r e g r e s s i v e phase containi n g b o t h non-marine gressive phase.
and m a r i n e e l e m e n t s , f o l l o w e d b y a t r a n s -
The m a r i n e o s t r a c o d s s u g g e s t a melange o f
b o t h n o r t h e r n and s o u t h e r n European forms. N o r t h A f r i c a , a f t e r s e p a r a t i o n f o r a l o n g p e r i o d o f Mesozoic t i m e , shows l i n k s w i t h Europe t o t h e n o r t h d u r i n g t h e Berrias i a n and m i g r a t i o n of t h e fauna northwards, probably due t o s h a l l o w i n g o f T e t h y s i n between (Donze 1 9 7 5 ) .
I n North
T u n i s i a , t h e Djebel Ouest s e c t i o n s p r o v i d e o s t r a c o d s from t h e Tithonian through t h e Berriasian.
The B e r r i a s i a n g e n e r a
i n c l u d e Amphicythere, Eucytherura, Hemicytherura, P a r a c y p r i s and P o n t o c y p r e l l a .
T e t h y s i a . which a p p e a r s e a r l i e r i n N .
A f r i c a , a l s o o c c u r s i n t h e B e r r i a s i a n of b o t h t h e r e a n d t h e Vocontian Trough and c o n f i r m s t h e a f f i n i t i e s between t h e t w o
areas. I n A l g e r i a P r o t o c y t h e r e m a z e n o t i , g. p a q u i e r i a n d 2. c f . p. r e v i l i a l s o e m p h a s i s e t h e c l o s e l i n k s w i t h t h e s o u t h of F r a n c e i n B e r r i a s i a n times.
252
2 . VALANGINIAN
M a r i n e c o n d i t i o n s b e c o m e much m o r e w i d e l y e s t a b l i s h e d a n d i n t h e B r i t i s h area t h e r e l i c t J u r a s s i c g e n e r a d i s a p p e a r and C y t h e r e l l a a n d P r o t o c y t h e r e make t h e i r a p p e a r a n c e .
F a u n a s of
t h i s a g e a p p e a r i n Germany a b o v e n o n - m a r i n e d e p o s i t s a n d a r e w i d e l y known f r o m H e l i g o l a n d a n d t h e n o r t h German o i l f i e l d P a r t i c u l a r l y c h a r a c t e r i s t i c are Protocythe-
across t o Poland.
re hannoverana, S t r a v i a crossata, D o l o c y t h e r i d e a w o l b u r g i and S c h u leridea praethoerenensis. N o r t h e r l y m i g r a t i o n from T e t h y s c a n a g a i n be s e e n (Donze 1 9 7 3 ) and by t h e end o f V a l a n g i n i a n
times E u r y i t y c y t h e r e a n d W e x o p h t h a l m o c y t h e r e h a d a l r e a d y r e a c h e d NW Germany a n d t h e l a t t e r g e n u s a n d Kentrodictyocythere lhoving s o u t h w a r d i n t o warmer s e a s , d i f f e-
had r e a c h e d P o l a n d .
r e n t s p e c i e s o f many g e n e r a a r e p r e s e n t a s c l e a r l y s e e n i n t h e
area o f Neuch2tel and t h e A l p e s d e Haute Provence where P r o t o c y t h e r e i s r e p r e s e n t e d b y p. p r a e t r i p l i c a t a , p. d i v i s a , 2. h e l v e t i c a and
p.
reicheli.
3 . HAUTERIVIAN In t h e Hauterivian t h e s h a r p d i s t i n c t i o n s between n o r t h e r n and southern European faunas d i s a p p e a r .
Cytherelloidea, Euryity-
c y t h e r e , P a r e x o p h t h a l m o c y t h e r e and R e h a c y t h e r e i s r e a c h t h e B r i t i s h a r e a a n d t h e o s t r a c o d s a r e a b u n d a n t and w i d e s p r e a d . I n B r i t a i n and N .
Germany, P a r a n o t a c y t h e r e d i g l y p t a , P r o t o -
c y t h e r e h e c h t i and R e h a c y t h e r e i s s e n c k e n b e r g i are c h a r a c t e r i s -
t i c a s well a s e l e m e n t s o f t h e c o n t i n u o u s l y e v o l v i n g P r o t o c y t h e r e t r i p l i c a t a and C y t h e r e l l o i d e a o v a t a l i n e a g e s . important genera include r u r a and S c h u l e r i d e a .
m, Dolocytheridea,
Other Eucythe-
I n t h e Lower S a x o n y B a s i n , Kemper ( 1 9 7 1 )
r e c o g n i s e d t h r e e o s t r a c o d f a c i e s i n t h e Upper V a l a n g i n i a n a n d Lower H a u t e r i v i a n ,
Associated with a typical suspension
f e e d e r c o m m u n i t y was a f a u n a r i c h i n s p e c i e s b u t p o o r i n i n d i v i d u a l s w i t h P r o t o c y t h e r e f r a n k e i , B a i r d i a , P o n t o c y p r e l l a and many c y t h e r u r i d s p e c i e s .
The s h a l l o w n e r i t i c t o s u b - l i t t o r a l
c r i n o i d f a c i e s p r o v i d e d t h e r i c h e s t f a u n a s which i n c l u d e d H a p l o c y t h e r i d e a kummi, P r o t o c y t h e r e t r i p l i c a t a , R e h a c y t h e r e i s senckenbergi, W l e r i d e a thoerenensis, Dolocytheridea s p p . , a n d many o t h e r s . b a t h y a l areas
E.
In t h e deeper p a r t s t r a n s i t i o n a l t o t h e kummi a n d P r o t o c y t h e r e h e c h t i m i g h t b e p r e s e n t .
C y t h e r e l l o i d e a was c o n s i d e r e d i n d i c a t i v e o f warm, s h a l l o w w a t e r .
253 F u r t h e r s o u t h i n t h e P a r i s B a s i n t h e g e n e r a are similar as
w e l l a s some o f t h e s p e c i e s b u t new s p e c i e s s u c h a s P r o t o c y t h -
e r e p u m i l a , 2. 4.
c a n c e l l a t a and S c h u l e r i d e a e x t r a n e a appear.
--
BARREMI A N
I n t h e B r i t i s h area c h a r a c t e r i s t i c s p e c i e s s u c h as C y t h e r e l l o i -
-dea
e,
d a l b y e n s i s , Amphicytherura b a r t e n s t e i n i , Eucytherura
Paranotacythere blanda,
g.
i n v e r s a i n v e r s a and o t h e r s are p r e s -
e n t a l t h o u g h t h e g e n e r a r e m a i n much t h e same a s b e f o r e .
d o b y t h o c y t h e r e is r e c o r d e d f o r t h e f i r s t t i m e i n B r i t a i n , P l a t y c y t h e r e i s and M e t a c y t h e r o p t e r o n o c c u r i n F r a n c e and SchulapaIn t h e P a r i s Basin a c h a r a c t e r i s t i c
c y t h e r e i n Eomania.
Protocythere fauna includes
g.
s t r i g o s a and
Paranotacythere damottae damottae occurs, i n v e r s a , S c h u l e r i d e a v i r g i n i s and
2.
g. v i l l i e r e n s i s .
a s well a s
bernouilensis.
2.
inversa
In t h e
S o u t h o f F r a n c e , Donze ( 1 9 7 1 ) h a s a g a i n drawn a t t e n t i o n t o t h e d i f f e r e n c e b e t w e e n t h e p a r g i n s of t h e V o c o n t i a n T r o u g h w h e r e 2 5 g e n e r a o c c u r r e d i n o n e s a p p l e a n d t h e d e e p e r p a r t of t h e
Trough where o n l y t h e f o u r g e n e r a C y t h e r e i s , P r o t o c y t h e r e , C y t h e r e l l a a n d P o n t o c y p r e l l a were p r e s e n t i n t h e G i s e m e n t d'Angles (Basses-Alpes). 5 . APTIAN A p t i a n is well-known
and y i e l d s a g r e a t v a r i e t y of o s t r a c o d s .
In t h e general B r i t i s h , N.
German a n d P a r i s B a s i n area t y p i c a l
s p e c i e s are Centrocythere b o r d e t i , C y t h e r e i s bekumensis, geometrica, Dolocythere
s, Dolocytheridea
intermedia
C.
m-
theropteron stchepinskyi, Paranotacythere inversa tuberculata, Protocythere croutesensis, Veenia f l o r e n t i n e n s i s .
2.
d e r o o i , S c h u l e r i d e a d e r o o i and
Developments of P r o t o c y t h e r e g i v e rise
t o t h e g e n e r a S a x o c y t h e r e a n d B a t a v o c y t h e r e o f Kemper ( 1 9 7 1 ) w h i c h make t h e i r a p p e a r a n c e i n t h i s s t a g e a n d S a x o c y t h e r e
x-
costata tricostata
In
i s f o u n d i n B r i t a i n a n d N . Germany.
N . Germany B a t a v o c y t h e r e h i l t e r m a n n i a p p e a r s i n t h e U p p e r
Aptian.
In S. France, Moullade (1963) found P r o t o c y t h e r e
oulensis,
2.
oertlitand
go alexanderi
bed-
c h a r a c t e r i s t i c of t h e
B e d o u l i a n ( L . A p t i a n ) a l t h o u g h t h e f i r s t t w o r a n g e down i n t o t h e Barremian and Gargasian.
2. b e d o u l e n s i s
i s a l s o f o u n d i n t h e Lower
I n t h e G a r g a s i a n (Upper A p t i a n ) a t t h e t y p e loca-
l i t y o f A p t , O e r t l i ( 1 9 5 8 ) r e c o r d e d amongst o t h e r s , D o l o c y t h e -
254
r i d e a intermedia (found also i n t h e n o r t h ) , S c h u l e r i d e a - j o n e s i a n a (which d o e s n o t a p p a r e n t l y r e a c h t h e P a r i s B a s i n and nort h e r n E u r o p e u n t i l t h e A l b i a n ) , a n d a number o f n e w s p e c i e s including Cythereis bartensteini,
C.
b u c h l e r a e , N e o c y t h e r e E-
t e n s i and P l a t y c y t h e r e i s r e c t a n g u l a r i s . 6 . ALBIAN
H i t h e r t o , o u r information h a s been confined t o t h e e a s t e r n s i d e of t h e North A t l a n t i c Basin,
W i t h t h e A l b i a n came a s t r i k i n g
e x t e n s i o n of t h e s h a l l o w s h e l f s e a s i n t o t h e e a s t e r n a n d s o u t h e r n U n i t e d S t a t e s s o t h a t d e p o s i t i o n i n t h e A t l a n t i c B a s i n was The f a u n a s o f t h e s e b e d s are c o m p a r a t i -
no longer one-sided. v e l y well-known.
I n E u r o p e , t y p i c a l Albizn f o r m s i n c l u d e B a t a v o c y t h e r e gaultina, Isocythereis fissicostatus, vanveeni, Protocythere mertensi, Saxocythere dividera, Veenia h a r r i s i a n a .
S.
I.f o r t i n o d i s , p.
nodigera,
2.
Neocythere speetonensis,
n o t e r a , S c h u l e r i d e a j o n e s i a n a and
Dolocytheridea intermedia intermedia gets
i n t o t h e Lower A l b i a n a n d
2.
b o s q u e t i a n a is c h a r a c t e r i s t i c o f
t h e Middle and Upper A l b i a n and j u s t g e t s i n t o t h e Cenomanian. I n a d d i t i o n t h e r e is a w i d e v a r i e t y o f s p e c i e s o f A c r o c y t h e r e , Alatacythere, Argilloecia,. Clithrocytheridea,
Cythereis,
m-
e r e l l o i d e a , Dolocytheridea, Eucythere, Eucytherura, Habrocyth-
ere,
Hemicytherura,
Isocythereis, Krausella, Macrocypris,
s-
cythere, P l a t y c y t h e r e i s , Protocythere, Paxocythere, Schuleridea, and Veenia. occur.
In
The g e n e r a B a i r d i a , Conchoecia and P o l y c o p e a l s o p he
P a r i s Basin Matronella appears i n t h e Albian
and h a s been r e c o r d e d from t h e N o r t h e r n I r e l a n d Cenomanian. A l t h o u g h i n t e r m e d i a t e s a r e n o t known i t i s t h o u g h t t o b e t h e
P a i r e d s t e r e o s c o p i c p h o t o g r a p h s o f E u r o p e a n Lower Plate 2. Cretaceous marine Ostracoda. A l l specimens from England except where s t a t e d . 1 . G a l l i a e c y t h e r i d e a t e r e s ( N e a l e ) , RV x 5 2 , B e r r i a s i a n ; 2 . E l o m e d e s d o n z e i Neale, 7 c a r a p a c e x 35, B a s a l V a l a n g i n i a n , F r a n c e ; 3 . M a n d e l s t a m i a s e x t i N e a l e , LV x 7 0 , B e r r i a s i a n ; 4 , P r o t o c y t h e r e h a n n o v e r a n a B a r t e n s t e i n a n d B r a n d , LV x 6 4 ; V a l a n g i n i a n ; 5 . P a r a n o t a c y t h e r e d i g l y p t a ( T r i e b e l ) , LV x 9 9 , H a u t e r i v i a n ; 6 . C y t h e r e i s a c u t i c o s t a t a T r i e b e l , LV x 6 5 , B a r r e m i a n ; 7 . w l e r i d e a jonesiana (Bosquet), LV x 4 7 , A l b i a n - C e n o m a n i a n ; 8. P l a t y c y t h e r e i s g a u l t i n a ( J o n e s ) , LV x 8 8 , A l b i a n .
255
PLATE 2
256
a n c e s t o r of S p i n o l e b e r i s i n t h e Campanian.
Also very charac-
t e r i s t i c are C o r n i c y t h e r e i s , C y t h e r e i s , I s o c y t h e r e i s and P l a t y cythereis.
The g e n u s P a r a n o t a c y t h e r e a p p e a r s r e s t r i c t e d t o
B r i t a i n i n t h e Albian where t h e s i n g l e s p e c i e s occurs (Bassiouni 1974).
'.
fordensis
Chapmanicythereis which f i r s t
a p p e a r e d i n t h e A p t i a n a n d c o n t i n u e d on t o t h e T u r o n i a n i s a l s o i m p o r t a n t and i s s e p a r a t e d from t h e a l l i e d P l a t y c y t h e r e i s by t h e p r e s e n c e o f an e y e t u b e r c l e and l o n g i t u d i n a l r i b b i n g .
The
European f a u n a is f a i r l y c o s m o p o l i t a n d u r i n g t h i s (and subseque n t s t a g e s ) a l t h o u g h some r e g i o n a l d i f f e r e n c e s i n s p e c i e s a r e apparent. Sivain r e c o r d e d s p e c i e s o f C y t h e r e l l a , A s c i o c y t h e r e , S c h u l e r i d e a ? , C y t h e r e i s and P r o t o c y t h e r e from a core t a k e n i n t h e E a s t e r n A t l a n t i c a t 2 5 0 5 5 . 5 3 ' N , 27003.64'11' a p p r o x i m a t e l y 1000
kms west o f Cap B l a n c o n t h e A f r i c a n c o a s t a n d c o n c l u d e d t h a t t h e a g e o f t h e f a u n a was p r o b a b l y A l b i a n .
A Kestern Atlantic
c o r e f r o m 0 9 0 2 7 . 2 3 ' K , 5 4 O 2 0 , 5 2 ' \ Y , some 4 0 0 kms n o r t h of t h e Guianas y i e l d e d a l a r g e r fauna of 25 s p e c i e s i n c l u d i n g t h r e e s p e c i e s found i n t h e c o r e above and Swain a g a i n c o n c l u d e d t h a t a n A l b i a n a g e was most l i k e l y . I n f o r m a t i o n on p r e - A l b i a n Scarce.
beds i n t h e United S t a t e s is
S w a i n a n d Brown ( 1 9 7 2 ) s u g g e s t t h a t some o f t h e
m a t e r i a l f r o m t h e A t l a n t i c C o a s t a l P l a i n ( U n i t H) i s e a r l y Cretaceous i n age but t h a t t h e b a s e of t h e u n i t m a y b e J u r a s s i c . Vanderpool ( 1 9 2 8 , 1 9 3 3 ) d e s c r i b e d forms from t h e Upper T r i n i t y G r o u p of Sli A r k a n s a s , SE O k l a h o m a , T.J T e x a s a n d K11' L o u i s i a n a which i n c l u d e d A s c i o c y t h e r e p e r f o r a t a ,
A.
r o t u n d a , Eocytherop-
t e r o n t r i n i t i e n s i s , P a r a c y p r i s n e a t h e r f o r d e n s i s and S c h u l e r i d e a dorsoventrus. S w a i n a n d Brown ( 1 9 6 4 ) a d d e d t h e new name Cythe r e i s p r a e o r n a t a and C y p r i d e a d e q u e e n e n s i s and two o t h e r C y p r i deas C
> -. d i m i n u t a a n d
C. w y o m i n g e n s i s
occur.
The age is u s u a l -
ly a s s u m e d t o b e l a t e A p t i a n ? b u t more p r o b a b l y e a r l y A l b i a n i n t h e c a s e o f t h e Upper T r i n i t y m a t e r i a l .
S w a i n a n d Brown (1964)
d e s c r i b e a n e a r s h o r e A l b i a n f a u n a i n C e n t r a l a n d NE N o r t h C a r o l i n a o f 1 2 s p e c i e s ( a l m o s t a l l new) w h i c h were a s s i g n e d t o
E-
throcytheridea, Cythereis, ?Dolocytheridex, Eocytheropteron, Eucythere, Eucytheroides, Fossocytheridea, Haplocytheridea, t h o n o t a c y t h e r e ar.d P e r i s s o c y t h e r i d e a .
The l l i d d l e and Upper
T r i n i t y f a u n a Prom t h e A t l a n t i c C o a s t a l P l a i n of S w a i n a n d
e-
257
Brown ( 1 9 7 2 ) h a s n o s p e c i e s i n common w i t h t h e p r e v i o u s f a u n a a n d i n c l u d e s m a i n l y s p e c i e s of A s c i o c y t h e r e , D o l o c y t h e r i d e a , S c h u l e r i d e a , P a r a s c h u l e r i d e a and H u t s o n i a . T h e h i g h e r Lower a n d ! , l i d d l e A l b i a n ( F r e d e r i c k s b u r g G r o u p ) a n d U p p e r A l b i a n ( W a s h i t a G r o u p ) h a v e b e e n c o v e r e d i n d e t a i l by Alexander (1929, 1932-34)
f o r N . T e x a s a n d i s v a i n a n d Brown
( 1 9 7 2 ) c o r r e l a t e t h e i r U n i t F of t h e A t l a n t i c C o a s t a l P l a i n w i t h t h e s e two g r o u p s .
A l e x a n d e r d e s c r i b e s many s p e c i e s a s s i -
gned t o B a i r d i a , B y t h o c y p r i s , C y t h e r e l l a , C y t h e r e l l o i d e a ,
w-
e r i d e a , Cytheropteron, Cytherura, Eocytheropteron, Paracypris and o t h e r s whose g e n e r i c taxonomy c o u l d do w i t h c o n s i d e r a b l e updating.
Swain and Brown's ( 1 9 7 2 ) U n i t F o f t h e A t l a n t i c
C o a s t a l P l a i n c o n t a i n s n o t h i n g i n common w i t h t h e T e x a s f a u n a and c o n s i s t s o f nen s p e c i e s , t o g e t h e r w i t h p r i n c i p a l l y s p e c i e s
of Swain and B r o m ( 1 9 6 4 ) and Swain ( 1 9 5 2 ) a s s i g n e d t o Asciocyt h e r e , Dolocytheridea, Cythereis, Eocytheropteron, Clithrocythe r i d e a , Eucythere, Fossocytheridea, Haplocytheridea, Orthonotac y t h e r e and P e r i s s o c y t h e r i d e a . Cpper C r e t a c e o u s i d e n i n g A t 1ant i c R i f t , c h a l k f a c i e s became w i d e l y e s t a b l i s h e d i n t h e n o r t h e r n h e m i s p h e r e as time p r o g r e s s e d . Only i n t h e e a r l i e s t and
h' i t h t h e c o n t i n u i n g C r e t ac e o u s t r a n s g r e ss i o n a n d
l a t e s t s t a g e s a r e b r a c k i s h w a t e r o s t r a c o d s known.
Vany Lower
C r e t a c e o u s g e n e r a s u c h as A p a t o c y t h e r e , C y t h e r o p t e r i n a , Dolocyt h e r i d e a , H a b r o c y t h e r e , P a r a n o t a c y t h e r e , and P r o t o c y t h e r e d i e d o u t or o n l y s u r v i v e d a s h o r t d i s t a n c e a b o v e t h e b a s e o f t h e Cenomanian.
Chapmanicythereis r e p l a c e d P l a t y c y t h e r e i s and
d u r i n g t h e U p p e r C r e t a c e o u s t h e r e was a g r e a t e x p a n s i o n a n d p r o l i f e r a t i o n of t r a c h y l e b e r i d s .
The P l a t y c o p a , B a i r d i a ,
K r i t h e , P a r a c y p r i s a n d many o t h e r g e n e r a c o n t i n u e a n d t h e f a u n a s b e g i n t o t a k e o n a much m o r e m o d e r n a s p e c t .
.
1
CENORIAN I AN
I n t h e E a s t e r n A t l a n t i c , f a u n a s a r e known f r o m N o r t h e r n I r e l a n d t o t h e Atlas Mountains.
E,
In t h e north Dolocytheridea bosqueti-
Neoeythere vanveeni and S c h u l e r i d e a ~-
j o n e s i a n a l i n g e r on
f r o m t h e A l b i a n a n d M a t r o n e l l a m a t r o n a e a l s o c r o s s e s t h e boundary.
Veenia r e p l a c e s P r o t o c y t h e r e a s t h e dominant p r o t o c y t h -
258
2.
S c h u l e r i d e a c o n t i n u e s as
erinid.
tumescens i n t h e Dordogne
and T o u r a i n e and D o l o c y t h e r i d e a i n b o t h t h e Dordogne and Toura i n e and A l g e r i a as
2.
crassa and
2.
atlasica respectively.
I n t h e P a r i s B a s i n t h e new g e n e r a D o r d o n i e l l a a n d F i s a l t i n a appear.
Many new t r a c h y l e b e r i d s make t h e i r a p p e a r a n c e , s u c h
as Dumontina, M a u r i t s i n a , O e r t l i e l l a and p r o b a b l y Limburgina a l t h o u g h t h e l a t t e r i s b e t t e r known f r o m t h e C a m p a n i a n a n d Maastrichtian.
In southern France, Amphicytherura, Annosocythere, and B a i r d i a are
C u r f s i n a , Metacytheropteron, Opimocythere important,
P a r t i c u l a r l y n o t a b l e i s t h e l a r g e number a n d v a r -
i e t y o f s p e c i e s of Z y t h e r e i s i n a l l a r e a s .
C y t h e r e l l a is ubi-
q u i t o u s but C y t h e r e l l o i d e a is rarely recorded.
Only i n t h e
D o r d o g n e (SW F r a n c e ) h a s a b r a c k i s h water f a u n a o f M e t a c y p r i s and Theriosynoecum been n o t e d ( C o l i n 1 9 7 4 ) . I n t h e A t l a n t i c Coastal P l a i n comparable faunas have been d e s c r i b e d b y S w a i n a n d Brown ( 1 9 7 2 ) i n U n i t E f r o m N . C a r o l i n a a n d V i r g i n i a a n d S w a i n a n d Brown ( 1 9 6 4 ) f r o m t h e Lower A t k i n s o n A w i d e v a r i e t y o f Q-
o f S . Alabama, N . F l o r i d a and G e o r g i a .
T h e more n o r t h e r l y area w i t h D o l o c y t h e r i d e a and S c h u l e r i d e a ? r e c a l l s t h e p o s i t i o n i n Europe. T h e more s o u t h e r l y a r e a w i t h C y t h e r e l l o i d e a s u g g -
t h e r e i s with Cytherella occurs throughout.
e s t s p e r h a p s some t e m p e r a t u r e z o n i n g . Both areas y i e l d t h e w e l l - k n o w n C y t h e r e i s e a g l e f o r d e n s i s , r e g a r d e d as a r e l i a h l e Upp e r C e n o m a n i a n i n d e x f u r t h e r west b y H a z e l ( 1 9 6 9 ) .
The s o u t -
h e r n a r e a s show a n a p p r o a c h t o t h e G u l f C r e t a c e o u s d e s c r i b e d b y Alexander (1929 e t a l . ) w i t h t h e o c c u r r e n c e o f C y t h e r e l l a Et i n e n s i s , B a i r d i a comanchensis, P a r a c y p r i s washitaensis.
rn a n d
Schuleridea
T h e W o o d b i n e of N . T e x a s y i e l d s C y t h e r e i s b u r l e -
s o n e n s i s , C. roanokensis,
C. w o r t h e n s i s ,
e n s i s , and P o n t o c y p r e l l a a l e x a n d e r i ,
P a r a c y p r i d e i s grayson-
I n t h e Hammond W e l l , Mary
l a n d (Swain 1 9 4 8 ) , Leguminocythereis? p u s t u l o s a * f r o m 1588-98'
came f r o m b e d s t h o u g h t t o b e o f a p p r o x i m a t e l y t h i s a g e . 2 . TURONIAN
The T u r o n i a n is e s s e n t i a l l y a c o n t i n u a t i o n and c o n s o l i d a t i o n o f t h e p o s i t i o n i n t h e Cenomanian.
E a s t e r n A t l a n t i c f a u n a s are
d o m i n a t e d b y t h e t r a c h y l e b e r i d s a n d i m p o r t a n t new s p e c i e s of C y t h e r e i s , C u r f s i n a , P t e r y g o c y t h e r e i s and S p i n o l e b e r i s , t o g e t -
~-
h e r w i t h A s c i o c y t h e r e , D o r d o n i e l l a , P t e r y g o c y t h e r e and o t h e r genera. I n S . F r a n c e , Dumontina, M a u r i t s i n a and T r a c h y l e b e r i * T h i s s pecie s i s t y p e of E u c y t h e r o i d e s S w a i n and Brown1964(ed.)
259
dea occur -
amongst o t h e r s .
S a i d a is e s t a b l i s h e d i n m i d d l e Eur-
ope and produced a s t r a t i g r a p h i c a l l y u s e f u l s e q u e n c e of s p e c i e s up t h r o u g h i n t o t h e T e r t i a r y ( H e r r i g 1 9 6 8 ) .
In Algeria t h e
t y p i c a l l y African Ovocytheridea and Brachycythere s p . g r .
ekpo
s u g g e s t l i n k s w i t h t h e m a r i n e C r e t a c e o u s of t h e Gulf of Guinea. I n t h e W.
A t l a n t i c t h e Upper A t k i n s o n o f S . Alabama, N . Flo-
r i d a a n d Georgia h a s y i e l d e d o n l y f o u r s p e c i e s b e l o n g i n g t o t h e genera Brachycythere, C y t h e r e l l a , C y t h e r e i s and Haplocytheridea. The d e v e l o p m e n t o f o s t r a c o d f a u n a s seems e q u a l l y r e s t r i c t e d i n N . Texas where t h e Eagle Ford S h a l e y i e l d e d o n l y C y t h e r e l l a
' m u n s t e r i ' , B a i r d i a a l e x a n d r i n a , P t e r y g o c y t h e r e s a r a t o g a n a and Cythereis eaglefordensis.
How f a r t h e E a g l e F o r d i n T e x a s
s h o u l d b e r e g a r d e d a s T u r o n i a n i n age a n d how f a r a s C e n o m a n i a n is a r g u a b l e . 3. CONIACIAN
Least w e l l - k n o w n o f t h e s t a g e s a s r e g a r d s i t s o s t r a c o d f a u n a , i n E u r o p e P o k o r n y ( 1 9 6 4 ) h a s shown e v o l u t i o n i n t h e s u b s p e c i e s o f C y t h e r e i s m a r s s o n i f r o m t h e U. T u r o n i a n t o t h e C o n i a c i a n . The s p e c i e s i s n o t known f r o m t h e S a n t o n i a n . t i o n is seen i n t h e s u b s p e c i e s of
A similar evolu-
C. o r n a t i s s i m a .
There is a
wide v a r i e t y o f C y t h e r e i s and o t h e r g e n e r a i n c l u d e C y t h e r e l l o i -
dea
Eucythere, I d i o c y t h e r e . K a r s t e n e i s , O e r t l i e l l a , Phacorbah-
d o t u s , P t e r y g o c y t h e r e i s , S p i n i c y t h e r e i s and S p i n o l e b e r i s . A l g e r i a O v o c y t h e r i d e a p r o d u c t a and
0.
In
b r e v i s a l o n g w i t h Brachy-
c y t h e r e s p . g r . ekpo, Veenia, Mauritsina?,
and P r o t o b u n t o n i a
a g a i n i n d i c a t e l i n k s w i t h t h e West A f r i c a n a r e a . I n t h e W . A t l a n t i c t h e C o n i a c i a n i n c l u d e s t h e Tokio Formation of Arkansas and N . L o u i s i a n a and t h e A u s t i n Chalk of Texas b u t t h e u p p e r p a r t o f t h e s e i s almost c e r t a i n l y S a n t o n i a n a n d t h e f a u n a s are b e s t r e g a r d e d a s 'Lower S e n o n i a n ' . b i c o r n i s is a v e r y t y p i c a l s p e c i e s . t h e f a u n a from 1470-80'
Cythereis
I n Hammond Well, l l a r y l a n d
of C y t h e r e i s c f . b i c o r n i s ,
C. p a r a u s t i -
n e n s i s and Haplocytheridea p a r v a s u l c a t a probably belongs i n this stage.
I n t h e G u l f , Phacorhabdotus pokornyi, Schuleridea
t r a v i s e n s i s and Veenia r e t i c u l a t a are r e c o r d e d i n E. T e x a s (Hazel & Paulson 1964). ded C y t h e r e i s b i c o r n i s ,
I n Arkansas t h e Tokio Formation y i e l -
C.
h a n n a i , P t e r y g o c y t h e r e i s t o k i a n a and
Cytheropteron s p . ( I s r a e l s k y 1929).
From t h e A u s t i n F o r m a t i o n
o f T e x a s come B a i r d i p p i l a t a r o t u n d a , B r a c h y c y t h e r e s p h e n o i d e s ,
260 I
/
FIG. 3
UPPER CRETACEOUS - DISTRIBUTION
OF LAND
am
SEA
P l a t e 3 . P a i r e d s t e r e o s c o o i c o h o t o a r a o h s of American C r e t a c e o u s Marine O s t r a c o d a . 1 . E o c y t h e r b p t e r o n tumidum ( A l e x a n d e r ) LV x 76. 2. R e h a c y t h e r e i s f r e d e r i c k s b u r g e n s i s ( A l e x a n d e r ) . LV x 5 7 . 3.=l o c y t h e r i d e a ? g l o b o s a ( A l e x a n d e r ) . LV x 63. 4. loc theridea? 5 . B r a d l e y a h:zar:i (Israelcf . e l u m m e r i ( A l e x a n d e r ) 8 RV x 7 0 . 7. sky) m x 2 . C y t h e r e l l a t u b e r c u l i f e r a A l e x a n d e r .LV x 5 2 . S p h a e r o l e h e r i s p s e u d o c o n c e n t r i c a ( B u t l e r & J o n e s ) RV x 1 3 9 . 8 . C u r f s i n a communis ( I s r a e l s k y ) . RV x 7 3 . 1,2. Alexander S t a t i o n 3-6. A l e x a n d e r S t a t i o n 6 0 , 5 , Texas. Kiamichi Fm., U . A l h i a n . 7 , 8 . C a . 2 m i l e s S . of Lucky, Texas. Navarro Fm., M a a s t r i c h t i a n . Louisiana. B u t l e r & J o n e s L o c a l i t y 1 9 5 7 . S a r a t o g a Fm., Campanian,
.
261
PLATE 3
262
B.
C.
t a y l o r e n s i s , Cythere cornuta v a r . g u l f e n s i s ,
Cythereis austinensis,
C.
bicornis,
C.
foersteriana,
dallasensis, Cytherella
a u s t i n e n s i s , K r i t h e c u s h r a n i , P a r a c y p r i s t e n u i c u l a and Veenia ozanana. 4 . SANTONIAN A l a r g e number o f s p e c i e s is known f r o m t h i s s t a g e i n n o r t h e r n
and c e n t r a l Europe.
i t a go
Imhotepia? s i m i l i s and I d i o c y t h e r e d e f i n -
r i g h t t h r o u g h t h e s t a g e and a l i t t l e above t h e b a s e o f
t h e Campanian i n t h e B a l t i c I s l a n d o f Rugen.
H e r r i g (1967 et
a l . ) , G r u n d e l ( 1 9 6 8 e t a l . ) a n d Ohmert ( 1 9 7 3 ) h a v e d e s c r i b e d f a u n a s f r o m Rugen a n d n o r t h E u r o p e a n d t h e a p p e a r a n c e o f Golcoc y t h e r e , M o s a e l e b e r i s and K i k l i o c y t h e r e is n o t a b l e i n t h e s e beds, together with various species of Costaveenia, Cytherella, Cytherelloidea, Cytheropteron, Eucytherura, Trachyleberidea, Veenia etc.
O h m e r t d e s c r i b e d t h e new g e n e r a D e r o o c y t h e r e a n d
K i k l i o p t e r y g i o n which o c c u r i n S a n t o n i a n b e d s .
Phacorhabdotus
s e m i p l i c a t u s seems f a i r l y w i d e s p r e a d a n d t h i s , or a c l o s e l y rel a t e d s p e c i e s , is f o u n d i n t h e A l p e s M a r i t i m e s o f F r a n c e w h e r e Donze and P o r t h a u l t ( 1 9 7 0 ) h a v e a l s o n o t e d s p e c i e s o f C y t h e r e i s , K a r s t e n e i s , O e r t l i e l l a , P t e r y g o c y t h e r e i s and T r a c h y l e b e r i d e a . I n t h e W. A t l a n t i c t h e Austin Chalk fauna h a s been considered for convenience with t h e Coniacian.
Similarly, t h e Taylor
f a u n a w h i c h may b e i n p a r t U . S a n t o n i a n is c o n s i d e r e d u n d e r Campanian.
H a z e l a n d P a u l s o n ( 1 9 6 4 ) c o n s i d e r t h e Brownstown
Marl i n w h i c h I s r a e l s k y ( 1 9 2 9 ) f o u n d V e e n i a s p o o r i t o b e L . Campanian. 5
.
CAMPAN I AN
A number o f J o n e s a n d H i n d e ’ s c l a s s i c s p e c i e s s u c h a s P a r a c y p r i s
s i l i q u a , P a r i c e r a t i n a t r i c u s p i d a t a , C u r f s i n a &,
and Cyther-
e l l a o b l i q u i r u g a t a come f r o m d e p o s i t s o f t h i s a g e i n B r i t a i n , b u t more r e c e n t w o r k o n t h e I s l a n d o f Rugen h a s b e t t e r s t r a t i g r a p h i c c o n t r o l ( H e r r i g 1967 e t a l . ) .
e, Amphicytherura dia denticulata
Here I d i o c y t h e r e r e p l i -
chelodon, A r g i l l o e c i a d e c u s s a t a and
w-
a l l make t h e i r a p p e a r a n c e a t t h e b a s e of t h e
Campanian, t h e l a t t e r t h r e e c o n t i n u i n g i n t o t h e M a a s t r i c h t i a n .
=,
G e n e r a l l y i n Europe C u r f s i n a , Dumontina, Limburgina, MosaelebeP l a n i l e b e r i s and S p i n o l e b e r i s are i m p o r t a n t g e n e r a t o g e t -
h e r w i t h Amphicytherura, Phacorhabdotus, Physocythere and Saida
263
a s w e l l a s many o t h e r s a n d t h e u b i q u i t o u s P l a t y c o p a .
In the
A q u i t a i n e B a s i n o f SW F r a n c e t h e f a u n a h a s c o n s i d e r a b l e a f f i n i -
ties w i t h t h a t of t h e M a a s t r i c h t i a n of Holland. Deroo ( 1 9 6 6 ) h a s s u g g e s t e d m i g r a t i o n from s o u t h t o n o r t h b u t C o l i n ( 1 9 7 3 ) f a v o u r s an e c o l o g i c a l e x p l a n a t i o n . The f u l l s i g n i f i c a n c e of t h e s i m i l a r i t i e s h a s s t i l l t o b e worked o u t . A f a u n a from A l geria includes t h e genera Acanthocythereis, Bradleya, Cythereis, C y t h e r o p t e r o n , V e e n i a a n d V e e n i d e a b u t p r o b a b l y t h e most i n t e r e s t i n g g e n u s is B u n t o n i a w h i c h i s s o w e l l d e v e l o p e d i n t h e N i g erian Cretaceous. I n t h e west t h e Matawan Group o f M a r y l a n d , Delaware a n d V i r g i n i a h a s y i e l d e d 27 s p e c i e s (Schmidt 1 9 4 8 ) .
Genera represen-
t e d include Bairdoppilata, Brachycythere, Cythereis, Cytherelloidea
->
K r i t h e and -
Xestoleberis.
T h e g e n e r i c s t a t u s o f many o f
t h e s p e c i e s r e f e r r e d t o C y t h e r e i s n e e d s r e a s s e s s m e n t i n terms of modern t ax o n o my .
B e d s a s s i g n e d t o t h e same G r o u p i n Ham-
mond Well, M a r y l a n d ( S w a i n 1 9 4 8 ) y i e l d e d a f a u n a o f B a i r d i a , C y t h e r e i s , Haplocytheridea and P a r a c y p r i s .
I t is f r o m t h e
G u l f C o a s t a r e a , h o w e v e r , t h a t t h e f a u n a i s b e s t known.
Acc-
ording t o t h e d i s t r i b u t i o n t a b l e s i n Alexander (1929), Cyther-
e i s a u s t i n e n s i s , g. b i c o r n i s
and
C. d a l l a s e n s i s
d i e o u t between
t h e A u s t i n and T a y l o r Formations w h i l e i n t h e l a t t e r sima, Haplocytheridea? grangerensis, a n g u s t a make t h e i r a p p e a r a n c e .
-Veenia time.
ozanana and
H? p l u m m e r i
C. r u g o s i s -
and P a r a c y p r i s
Cytherella navarroensis,
l. p a r a t r i p l i c a t a
b e c o m e common f o r t h e f i r s t
D e t a i l e d s t u d i e s o f t h e C a m p a n i a n h a v e a l s o b e e n made
by I s r a e l s k y ( 1 9 2 9 ) a n d B e n s o n a n d T a t r o ( 1 9 6 4 ) i n A r k a n s a s a n d B u t l e r and J o n e s (1957) i n Louisiana.
Important genera inc-
lude A l a t a c y t h e r e , Amphicytherura, B a i r d o p p i l a t a , Brachycythere, Bythocypris, Cytherella, Cytherelloidea, Haplocytheridea?, K r i t h e , Loxoconcha, E o n o t a c y t h e r e , P a r a c y p r i s , P t e r y g o c y t h e -
E, V e e n i a , a n d X e s t o l e b e r i s .
From E . T e x a s H a z e l a n d P a u l s o n
d e s c r i b e d a f a u n a r e g a r d e d a s of t h i s a g e w h i c h i n c l u d e d s p e -
c i e s of B r a c h y c y t h e r e , B r a d l e y a , P t e r y g o c y t h e r e i s and Veenia ( N i g e r i a ) b u t was c h i e f l y n o t a b l e for S c h u l e r i d e a t r a v i s e n s i s . T h i s i s t h e h i g h e s t C r e t a c e o u s r e c o r d of S c h u l e r i d e a ( a l s o n o t e d by them i n t h e C o n i a c i a n ) and i s s e p a r a t e d from o t h e r r e c o r d s by a c o n s i d e r a b l e i n t e r v a l .
They a l s o t h o u g h t t h a t
B r a c h y c y t h e r e d u r h a m i , which o c c u r s i n b o t h t h e A u s t i n Group
264
a n d t h e G o b e r C h a l k , was f a c i e s c o n t r o l l e d . 6 . MAASTRICHTIAN I n t h e t y p e area t h e f a u n a is knownin c o n s i d e r a b l e d e t a i l from t h e w o r k o f D e r o o ( 1 9 6 6 ) who d e s c r i b e d 1 6 3 I l a a s t r j . c h t i a n s p e -
cies,
F u r t h e r , h e was a b l e t o r e c o g n i s e s e v e n d i v i s i o n s i n
t h e s t a g e and s e p a r a t e it s a t i s f a c t o r i l y from t h e u n d e r l y i n g Campanian and o v e r l y i n g D a n i a n .
Species of Alatacythere,
As-
c i o c y t h e r e , B r a c h y c y t h e r e , C u r f s i n a , Dumontina, E u c y t n e r u r a , Hemicytherura, K i k l i o c y t h e r e , Kingmaina, Limburgina, h l a u r i t s i n a , Mosaeleberis, Netrocytheridea,
Phacorhabdotus, Sphaeroleberis,
V e e n i a , Veenidea and X e s t o l e b e r i s are i m p o r t a n t .
Acuticythe-
r e t t a , G l o b o l e b e r i s , F r o t o j o n e s i a and T u m i d o l e b e r i s are recorded f o r t h e f i r s t t i m e .
Bythoceratina
A t least t e n s p e c i e s are r e f e r r e d t o
a n d a c c o r d i n g t o Deroo t h e T e r t i a r y g e n e r a
a y i n a and P a l e o m o n s m i r a b i l i a and t h e T e r t i a r y and Recent
n i n a are
ww-
also p r e s e n t , T h e RGgen f a u n a ( H e r r i g 1 9 6 5 e t a l . ) c o n t a i n s a number o f s p e c i e s i n common w i t h H o l l a n d a s well a s
new s p e c i e s o f A r g i l l o e c i a , C y t h e r u r a , F o l y c o p e a n d t w o new species of Saida.
I n SW F r a n c e , B l a n c & C o l i n ( 1 9 7 5 ) h a v e
d e s c r i b e d a n Upper M a a s t r i c h t i a n f a u n a which i n c l u d e s Asciocythere
-9
B a i r d i a , E u c y t h e r e , K r i t h e , Limburgina?, Paleomonsmira-
b i l i a , a n d X e s t o l e b e r i s as well a s r a t h e r d o u b t f u l P a r a c y p r i s , Centrocytheridea and Cytheromorpha.
I n A l g e r i a i n r o c k s of
age B e l l i o n et a l . (1973) noted Proto-
p o s s i b l e l.!aastrichtian
buntonia numidica ( r a n g i n g t o t h e local b a s e of t h e Senonian), Brachycythere s p . g r .
sina sp.
a, Acanthocythereis
sp., and M a u r i t -
Evidence o f b r a c k i s h and l a g o o n a l c o n d i t i o n s is f o r t h - ;
coming f o r t h e f i r s t t i m e s i n c e t h e Cenornanian.
Damotte and
F o u r c a d e ( 1 9 7 1 ) f o u n d N e o c y p r i d e i s m u r c i e n s i s i n SE S p a i n a n d i n t e r p r e t e d t h e c o n d i t i o n s as a lagoon o r marginal environment with variable s a l i n i t y .
Also i n S p a i n i n t h e S. P y r e n e e s ,
L i e b a u ( 1 9 7 1 ) was a b l e t o r e c o g n i s e f o u r d i f f e r e n t s a l i n i t y controlled facies.
T h e t w o more b r a c k i s h f a c i e s c o n t a i n e d
B i s u l c o c y p r i s , C y p r i d e i s , C y p r i d o p s i s ? , C y t h e r o m o r p h a ? , Neocyp r i d e i s , P a r a k r i t h e ? and P a h m o n s m i r a b i l i a . I n t h e west o u r k n o w l e d g e i s b a s e d o n f a u n a s f r o m t h e N a v a r r o F o r m a t i o n d e s c r i b e d by A l e x a n d e r ( 1 9 2 9 ) a n d t h e f a u n a f r o m t h e A r k a d e l p h i a Marl d e s c r i b e d b y I s r a e l s k y ( 1 9 2 9 ) c o n s i s t i n g of B r a c h y c y t h e r e l e d a f o r m a ,
g.
sphenoides, Cythereis costatana,
265
Bradleya h a z a r d i , Phacorhabdotus t r i d e n t a t u s , C u r f s i n a
ivii i s
a n d two o t h e r s p e c i e s o f w h i c h C y t h e r e i s
S c h m i d t ' s f a u n a f r o m t h e Monmouth Group o f Rlary-
arachoides.
l a n d i s p o s s i b l y i n l a r g e p a r t Campanian.
I t c o n t a i n s Brachy-
cythere rhomboidalis, Haplocytheridea? fabaformis,
lira, H?
communis
probably Veenia
macropora,
H?
H?
f.multi-
a m y g d a l o i d e s b r e v i s , H? p l u n i m e r i ,
H?
u l r i c h i , C y t h e r e i s p i d g e o n i , C y t h e r o p t e r o n c o r y e l l i , Loxoconcha c r e t a c e a , P a r a c y p r i s monmouthensis and V e e n i a a r a c h o i d e s . Conclusions D u r i n g t h e C r e t a c e o u s , O s t r a c o d a a r e w e l l d e v e l o p e d on t h e E a s -
tern s d e of t h e A t l a n t i c i n t h e B e r r i a s i a n t o A p t i a n S t a g e s , and on b o t h s i d e s o f t h e A t l a n t i c from t h e A l b i a n onwards. Genera
f a u n a l s t u d i e s h a v e shown t h a t t h e y a r e u s e f u l i n re-
c o g n i s i n g t h e S t a g e s a n d sometimes f i n e r d i v i s i o n s .
Phylo-
g e n e t i c s t u d i e s of s i n g l e genera such as t h o s e of Bassiouni ( 1 9 7 4 ) , B e t t e n s t a e d t ( 1 9 5 8 ) , H e r r i g ( 1 9 6 8 ) , Neale ( 1 9 7 3 ) a n d o t h e r s hold out p o s s i b i l i t i e s of f i n e r d i s c r i m i n a t i o n .
Much
r e m a i n s t o b e done i n t h i s f i e l d and a l o t of t h e o l d e r work needs r e v i s i n g and u p d a t i n g t a x o n o m i c a l l y .
Often, t h e ostra-
cods can g i v e a l o t of u s e f u l information about t h e n a t u r e of t h e environment a t t h e time they l i v e d . Acknowledgements
I wish t o t h a n k D r . R. H . Bate, D r . R . G. Clements and D r . T . I . K i l e n y i who k i n d l y l o a n e d m e s p e c i m e n s o f C y p r i d e a . References Alexander, C.
I . 1929. Ostracoda of t h e Cretaceous of North
Texas, Alexander, C .
U n i v . T e x a s B u l l . 2 9 0 7 , 1 3 7 p p . , 10 p l s .
I . 1 9 3 4 . O s t r a c o d a o f t h e g e n e r a V o n o c e r a t i n a and
O r t h o n o t a c y t h e r e f r o m t h e C r e t a c e o u s of T e x a s . P a l e o n t . v . 8 , pp. 57-67,
J.
p l . 8.
Anderson, F . W. 1 9 7 3 . The J u r a s s i c - C r e t a c e o u s non-marine o s t r a c o d f a u n a s .
transition:
the
Geol. J . S p e c i a l I s s u e
No. 5 , p p . 101-111, 1 t e x t - f i g . Bassiouni,
El.
el A . A . 1974.
Paranotacythere n.g.
(Ostracoda
a u s dem Z e i t r a u m O b e r j u r a b i s u n t e r k r e i d e ( K i m m e r i d -
266 gium b i s Albium) von W e s t e u r o p a . pp. 1-112, Bellion, Y . ,
G e o l . J a h r b . v.17A,
p l s . 1-13, 1 t a b l e , 5 t e x t - f i g s .
Donze, P , and G u i r a u d , R .
1973.
Repartition Stra-
tigraphique des principaux ostracodes (Cytheracea) d a n s l e Cre/tac& S u p g r i e u r du Fud-Ouest
Constantinois Publ.
( C o n f i n s Hodna - A u r e s , Alge/rie du N o r d ) . S e r v . ge/ol. A l g g r i e ( N . S . )
No. 44, p p . 7-44, p l s .
1-7, 2 t a b l e s , 2 t e x t - f i g s . Benson, R . H . , and T a t r o , J . 0. 1964.
F a u n a l d e s r - L p t i o n of
O s t r a c o d a of t h e M a r l b r o o k Marl ( C a m p a n i a n ) , Arkan-
sas. 1-6,
Univ. Kansas P a l e o n t . C o n t r . 7 , p p . 1-32, p l s .
15 t e x t - f i g s . P h y l o g e n e t i s c h e Beobachtungen i n d e r
B e t t e n s t a e d t , F . 1958.
Mikropalgontologie.
Palaont. Z.
v . 32, p p . 115-140.
0
B l a n c , P . L., and C o l i n , J . P . 1975.
Etude Micropale/ontologi-
q u e e t Pal&ooe/cologique du V a a s t r i c h t i e n d e CgzanLavardens (Gers, S . O .
France).
Palaeontographica,
v . 148A, p p . 109-131, p l s . 1-4, 13 t e x t - f i g s . Butler, E . A.,
and J o n e s , D . E . 1957.
Cretaceous Ostracoda of
P r o t h r o and Rayburns S a l t Domes B i e n v i l l e P a r i s h , Louisiana.
La. G e o l . S u r v . B u l l . No. 32, p p . 1 - 6 5 ,
p l s . 1-6, 5 t e x t - f i g s . C h r i s t e n s e n , 0 . B . 1974.
Marine Communications t h r o u g h t h e
D a n i s h Embayment d u r i n g Uppermost J u r a s s i c and Lowermost C r e t a c e o u s .
G e o s c i e n c e and Van, v . 6 , p p . 99-
1 1 5 , 1 pl., 5 t e x t - f i g s . , C o l i n , J . P . 1973.
5 tables.
Nouvelle Contribution a l'e/tude des o s t r a -
c o d e s du Cre/tac& s u p g r i e u r d e Dordogne (S.O. F r a n c e ) . P a l a e o n t o g r a p h i c a , v . 143, p p . 1-38, p l s . 1 - 6 , 3 text-figs. C o l i n , J . P . 1974.
N o u v e l l e s esp\eces d e s g e n r e s E ' e t a c y p r i s e t
Theriosynoecum ( O s t r a c o d e s l a c u s t r e s ) d a n s l e Ce'nomanien d e Dordogne ( 8 . 0 . F r a n c e ) .
Rev. E s p a n . Mic-
r o p a l . v . 6 , p p . 183-189, 1 p l . D a m o t t e , R , and F o u r c a d e , E .
1971.
Neocyprideis murciensis n .
s p . , O s t r a c o d e nouveau du M a e s t r i c h t i e n d e l a p r o d e 1 ' E s p a g n e ) . B u l l . Soc. v . 13, p p . 169-173, p l . 6 , 1 t e x t - f i g .
v i n c e d e l h r c i e (Sud-Est geol. France.
267
Deroo, G . 1966.
C y t h e r a c e a ( O s t r a c o d e s ) du M a a s t r i c h t i e n de
Maastricht (Pays-Bas) e t d e s r g g i o n s v o i s i n e s ;
re's-
u l t a t s stratigraphiques et palgontologiques de l e u r ktude.
hfed. G e o l . S t i c h t . C . V .
2 , pp.
1 - 1 9 7 , pls.
1 - 2 7 , 9 t a b l e s , 20 t e x t - f i g s . Rapports e n t r e les f a c i b s et l a r g p a r t i t i o n
Donze, P . 1 9 7 1 .
ggne'rique d e s o s t r a c o d e s d a n s q u a t r e g i s e m e n t s - t y p e s , deux
% deux s y n c h r o n i q u e s , du B e r r i a s i e n e t du Rar-
rgmien du s u d - e s t
de l a France,
B u l l . C e n t r e Rech.
Pau - SNPA, v . 5 s u p p l . , p p . 6 5 1 - 6 6 1 , 2 t a b l e s , 2 text-figs. Donze, P
1973.
O s t r a c o d m i g r a t i o n s from t h e Mesogean t o Bor-
e a l P r o v i n c e s i n t h e E u r o p e a n Lower C r e t a c e o u s . G e o l . J . S p e c . I s s u e , L i v e r p o o l , v . 5 , p p . 155-160, 4 text-figs.
Donze, P
1975.
Pal6obiogebgraphie des populations d 'ostracodes
d e p a r t e t d ' a u t r e d e l a T e t h y s ( A f r i q u e du Rord e t E u r o p e O c c i d e n t a l e ) , a u J u r a s s i q u e s u p g r i e u r e t au Cr&tace/ i n f g r i e u r .
B u l l . S O C . Ge/ol. F r a n c e . ( 7 )
v . 2 7 , p p . 843-849,
3 text-figs. Thomel, G . , and V i l l o u t r e y s , 0 . d e . Le S g n o n i e n i n f 4 r i e u r d e P u g e t - T h g n i e r s
Donze, P . , P o r t h a u l t , B . , 1970.
(Alpes-Maritimes) et s a microfaune. p p . 41-106, Grundel, J . 1968.
p l s . 8-13,
text-figs.
Geobios v . 3 ,
1-4.
Neue O s t r a c o d e n a u s d e r S a l z b e r g m e r g e l -
F a z i e s ( S a n t o n ) i m w e s t l i c h e n T e i l d e r Deutschen D e m o k r a t i s c h e n R e p u b l i k . G e o l o g i e , 1 7 , p p . 947-963, p l s . 1-3
Hazel, J . E . 1969.
C y t h e r e i s e a g l e f o r d e n s i s A l e x a n d e r 1929
-
a g u i d e f o s s i l f o r d e p o s i t s o f l a t e s t Cenomanian a g e i n t h e w e s t e r n i n t e r i o r and Gulf C o a s t r e g i o n s of t h e United S t a t e s .
G e o l . S u r v . Res. 1 9 6 9 . D155-
D158, 2 f i g s . Hazel, J . E . , and Eaulson, O . L . ,
Jr. 1964.
Some new o s t r a c o d e
s p e c i e s from t h e A u s t i n i a n and T a y l o r a n ( C o n i a c i a n and Campanian) r o c k s of t h e E a s t T e x a s Embayment. J . P a l e o n t . v . 3 8 , p p . 1047-1064,
text-figs.
pls. 157-159, 2
268
1965.
Herrig, E
C y t h e r e i s r e t i c u l a t a v a r i a s s p . n . , e i n e neue
Ostracoden-Unterart
a u s d e r Rzgener S c h r e i b k r e i d e
(Unter-Maastricht).
B e r . g e o l . G e s . DDR. v . 1 0 ,
p p . 403-419, 1967.
Herrig, E
p l s . 1-4,
5 text-figs.
Moglichkeiten e i n e r F e i n s t r a t i g r a p h i e d e r
hgheren Oberkreide i n Nordostdeutschland m i t H i l f e von O s t r a c o d e n . Ber. d e u t s c h . G e s . Wiss. A . G e o l . P a l a o n t . v . 1 2 , p p . 557-574, 1968.
Herrig, E
p l s . 1,2, 8 t e x t - f i g s .
Zur G a t t u n g S a i d a H o r n i b r o o k ( O s t r a c o d a ,
Crustacea) i n der Oberkreide. Israelsky
,
M.C.
Geologie, 1 7 , pp.
p l . 1, 7 t e x t - f i g s .
964-981,
1929.
Upper C r e t a c e o u s O s t r a c o d a o f A r k a n s a s .
Ark. G e o l . S u r v . B u l l . No. 2 , p p . 1 - 2 9 , p l s . 1 - 4 ,
1 table. Kemper, E
1971a.
D i e Pal%oEkologische V e r b r e i t u n g d e r Ostra-
koden i m O b e r v a l a n g i n i u m und U n t e r h a u t e r i v i u m d e s Bull.
N i e d e r s a c h s i s c h e n Beckens (NW-Deutschland),
C e n t r e Rech. Pau -SNPA, v . 5 s u p p l . , p p . 631-649, pls. 1,2, 2 text-figs. Kemper, E . 1 9 7 1 b .
B a t a v o c y t h e r e und S a x o c y t h e r e , zwei n e u e
Protocytherinae-Gattungen ( O s t r a c o d a d e r U n t e r k r e i d e ) . S e n c k . L e t h . v . 5 2 , p p . 385-431,
p l s . 1-8,
1 text-
fig. K i l e n y i , T . I . , and A l l e n , N . W .
1968.
Marine b r a c k i s h bands
and t h e i r m i c r o f a u n a f r o m t h e l o w e r p a r t of t h e Weald C l a y of S u r r e y and S u s s e x .
Palaeontology,
v . 1 1 , p p . 1 4 1 - 1 6 2 , pls. 2 9 , 3 0 , 9 t e x t - f i g s . Liebau, A . 1971.
D i e Ableitung der Palzkologischen systematik
O b e r k r e t a z i s c h e n Lagune.
B u l l . C e n t r e Rech. Pau
SNPA. v . 5 s u p p l . , p p . 577-599,
-
2 pls, 2 tables, 1
text-f i g . Moullade, M .
1963.
P r i n c i p a u x r e p r g s e n t a n t s du g e n r e P r o t o c y -
t h e r e ( O s t r a c o d e s ) d a n s l e Cretace i n f g r i e u r du Sud-
E s t de l a France.
F e v . M i c r o p a l . , v . 6 , p p . 102-108,
pls. 1 , 2 , 2 tables.
N e a l e , J . W. 1973.
O s t r a c o d a a s means of c o r r e l a t i o n i n The
B o r e a l Lower C r e t a c e o u s w i t h s p e c i a l r e f e r e n c e t o t h e B r i t i s h marine Ostracoda. L i v e r p o o l , v . 5 , p p . 169-184,
Geol. J . Spec. I s s u e , 2 text-figs.
269
Les Ostracodes de 1'Aptien-Albien d'Apt.
O e r t l i , H . J . 1968.
Revue I n s t . f r . P g t r o l e 1 3 , p p . 1499-1537, p l s . 1 - 9 , 1 table, 3 text-figs.
Ohmert, W .
1973.
O s t r a c o d e n a u s dem S a n t o n d e r G e h r d e n e r B e r g e .
B e r . N a t u r h i s t . G e s . v . 1 1 7 , p p . 163-194, p l s . 15-18. Pokorn$, V . 1 9 6 4 .
The P h y l o g e n e t i c l i n e s of C y t h e r e i s n l a r s s o n i
Bonnema. 1 9 4 1 ( O s t r a c o d a , C r u s t a c e a ) i n t h e Upper C r e t a c e o u s o f Bohemia, C z e c h o s l o v a k i a . C a r o l . S e o l . No. 3 , p p . 255-274, figs
A c t a Univ.
p l s . 1 , 2 , 16 t e x t -
.
Schmidt, R . A . M .
1948.
O s t r a c o d a from t h e Upper C r e t a c e o u s
and Lower Eocene of M a r y l a n d , D e l a w a r e , and V i r g i n i a . J . P a l e o n t . v . 2 2 , p p . 389-431,
Swain, F. M .
1948.
p l s . 61-64.
O s t r a c o d a from t h e Hammond Well.
Maryland
Board N a t . R e s . , C r e t a c e o u s and T e r t i a r y S u b s u r f a c e Geology, p p . 1 8 7 - 2 1 3 , p l s . 1 2 - 1 4 . Sivain, F . M . 1 9 5 2 , O s t r a c o d a from w e l l s i n N o r t h C a r o l i n a . Part 2.
Mesozoic O s t r a c o d a .
U.S.G.S. P r o f . Pap.
243-B. Swain, F . M . , a n d Brown, P . I f . 1 9 7 2 . (?),
Lower C r e t a c e o u s J u r a s s i c
a n d T r i a s s i c O s t r a c o d a from t h e A t l a n t i c
C o a s t a l Region. p p . 1-55,
P r o f . Pap. U . S .
g e o l . Surv. 795,
10 p l s . , 32 f i g s .
Vanderpool, H . C . 1928.
F o s s i l s from t h e T r i n i t y Group (Lower
Comanchean), J . P a l e o n t . , v . 2 , p p . 95-107,
pls.
12-14, Vanderpool, H . C . 1933.
e r n Oklahoma. 49.
Upper T r i n i t y M i k r o f o s s i l s from s o u t h J . P a l e o n t . , v . 7 , p p . 406-411,
pl.
270
Discussion M . C . Keen: Is i t p o s s i b l e t o r e l a t e t h e C y p r i d e a z o n e s o f t h e N o r t h A t l a n t i c w i t h t h o s e o f West A f r i c a and S o u t h America? Neale: The d i f f e r e n c e i n t h e f a u n a s o f t h e two a r e a s i s s o g r e a t t h a t any d i r e c t c o r r e l a t i o n o f t h e z o n e s i s i m p o s s i b l e . Attempts t o r e l a t e t h e zones i n t h e r e s p e c t i v e areas t o t h e s t a n d a r d s t a g e s a r e s p e c u l a t i v e and c a n o n l y s u g g e s t v e r y t e n t a t i v e broad e q u i v a l e n c e s . D r . B . K . Holdsworth: Can o n e r e c o g n i s e b a t h y a l a s s e m b l a g e s i n t h e Cretaceous y e t ? Neale: Depth i n i t s e l f , a s d i s t i n c t from i t s a f f e c t o n tempe r a t u r e , l i g h t i n t e n s i t y a n d t o some e x t e n t s u b s t r a t e , i s unimpoktant a s a n e c o l o g i c a l f a c t o r and v e r y d i f f i c u l t t o assess a c c u r a t e l y . I n c o n s e q u e n c e it d e p e n d s how a c c u r a t e l y you want t o d e f i n e b a t h y a l . C e r t a i n l y it i s p o s s i b l e t o r e c o g n i s e i n s h o r e and o f f s h o r e f a u n a s a s s e e n i n t h e work o f Kemper i n t h e N o r t h German B a s i n and Donze i n V o c o n t i a n Trough. U n f o r t u n a t e l y , i n f o r m a t i o n o b t a i n e d by o i l e x p l o r a t i o n i n t h e European s h e l f areas h a s remained c o n f i d e n t i a l b u t I understand t h a t t h e s h e l f faunas p a s s outwards i n t o f a u n a s t h a t c o n t a i n a h i g h p r o p o r t i o n o f B a i r d i a w h i c h may i n d i c a t e an approach t o a t r u b a t h y a l environment. Keen: Do you t h i n k o s t r a c o d s w i l l b e o f a n y h e l p i n d e t e r mining t h e d e p t h of d e p o s i t i o n of t h e Chalk ( i n c l u d i n g chalk m a r l s , glauconitic layers chalk rocks)? I ask t h i s b e c a u s e Hancock h a s r e c e n t l y t h r o w n d o u b t o n some o f t h e d e p t h z o n a t i o n s d e t e r m i n e d by f o r a m s ( P r o c . G e o l . A S S O C . , 1975) Neale: B e a r i n g i n mind my p r e v i o u s r e m a r k s , I t h i n k t h a t e v e n t u a l l y i t w i l l b e p o s s i b l e t o g i v e some g e n e r a l i n d i c a t i o n o f d e p t h s i n r e l a t i v e terms s u c h a s i n n e r n e r i t i c , outer n e r i t i c , bathyal, etc. Dr.
CRETACEOUS OSTRACODA
-
271
SOUTH ATLANTIC
Alwine B e r t e l s Facultad de Ciencias Exactas y Naturales, U n i v e r s i d a d d e Buenos A i r e s , R e p 6 b l i c a A r g e n t i n a Consejo N a c i o n a l d e I n v e s t i g a c i o n e s C i e n t i f i c a s y T h i c a s , RepAblica A r g e n t i n a Abstract En e l p r e s e n t e t r a b a j o s e compendian 10s a p o r t e s c i e n t i f i c o s e f e c t u a d o s p o r d i v e r s o s a u t o r e s r e l a c i o n a d o s con l a s a s o c i acionec d e ostrAcodos que p r e v a l e c i e r o n d u r a n t e e l CretAcico en Areas d e l A t l & i t i c o S u r . S e a n a l i z a n l a s c u e n c a s s e d i m e n t a r i a s e n donde t u v i e r o n
lugar depositaciones durante e l C r e t h i c o . Hacia f i n e s d e l J u r A s i c o y comienzos d e l C r e t i c i c o t u v i e r o n l u g a r 10s p r i m e r o s movimientos d e o r i g e n t e c t 6 n i c o q u e p r o d u j e r o n d i v e r s o s t i p o s d e e s t r u c t u r a s e n 10s a c t u a l e s m&genes d e 10s c o n t i n e n t e s s u d a m e r i c a n o y A f r i c a n 0 y q u e c o n t r o l a r o n l a s Areas d e s e d i m e n t a c i d n . D u r a n t e e l Neocomiano, con e x c l u s i 6 n d e l a s r e g i o n e s a u s t r a l e s d e ambos c o n t i n e n t e s
,
l a s e d i m e n t a c i c h es n e t a m e n t e con-
t i n e n t a l . Las f a u n a s d e o s t r A c o d o s s o n e n l a m a y o r i a d e 10s c a s o s comunes a ambas A r e a s . Durante e l Aptiano-Albiano se producen l a s primeras i n gresiones marinas que originan dep6sitos de t i p o evaporitico. A p a r t i r d e e s t e i n t e r v a l 0 predomina l a s e d i m e n t a c i 6 n mar-
i n a q u e s e r e g i s t r a e n d i v e r s a s c u e n c a s aunque no s i m u l t i n e a mente e n t o d a s e l l a s . E l c i c l o C r e t A c i c o c u l m i n a con l a t r a n s g r e s i 6 n d e l MaasI
t r i c h t i a n o q u e ocupa v a s t a s a r e a s e n ambos c o n t i n e n t e s .
Las
a s o c i a c i o n e s q u e s e conocen h a s t a e l p r e s e n t e , s i b i e n p r e s e n t a n s i m i l i t u d e s , no e x h i b e n e s p e c i e s i d A n t i c a s , l o c u a l l l e v a a l a conclusi6n que d u r a n t e e l M a a s t r i c h t i a n o e l A t l i n t i c o Sur e s t a b a netamente d e l i n e a d o . Summary I n t h e p r e s e n t work t h e s c i e n t i f i c c o n t r i b u t i o n s made by several authors a r e synthesized r e l a t i v e t o the ostracodal assemblages which p r e v a i l e d d u r i n g t h e C r e t a c e o u s i n S o u t h A t l a n -
tic areas. A t t h e end of t h e J u r a s s i c and e a r l y C r e t a c e o u s t h e f i r s t t e c t o n i c e v e n t s produced s e v e r a l s t r u c t u r e s which c o n t r o l l e d
272
t h e sedimentary a r e a s . D u r i n g t h e Neocomian, e x c l u d i n g t h e s o u t h e r n m o s t r e g i o n s i n b o t h t h e S o u t h American and A f r i c a n c o n t i n e n t s , non-marine sedimentation prevailed. During Aptian-Albian
t i m e s t h e f i r s t marine transgression
t o o k p l a c e which formed e v a p o r i t i c b e d s . From t h i s t i m e f o r w a r d , m a r i n e s e d i m e n t a t i o n was predomin a n t i n t h e C r e t a c e o u s a l t h o u g h i t was n o t c o n t e m p o r a n e o u s i n a l l basins. The C r e t a c e o u s c y c l e c u l m i n a t e s w i t h t h e M a a s t r i c h t i a n t r a n s g r e s s i o n which o c c u p i e d v a s t a r e a s i n b o t h t h e S o u t h Ameri c a n and A f r i c a n c o n t i n e n t s .
The known o s t r a c o d a l a s s e m b l a g e s
show s i m i l a r i t i e s which would i n d i c a t e t h a t d u r i n g t h e Maast r i c h t i a n t h e A t l a n t i c Ocean was d e f i n i t i v e l y d e l i n e a t e d . Introduction The p u r p o s e of t h e p r e s e n t work i s t o s y n t h e s i z e t h e most d i a g n o s t i c o s t r a c o d a l a s s e m b l a g e s which p r e v a i l e d d u r i n g C r e t a c e o u s t i m e i n t h e p r e s e n t A f r i c a n and S o u t h American c o n t i nents. The s e d i m e n t a r y b a s i n s i n which C r e t a c e o u s d e p o s i t s o c c u r a r e analyzed i n both c o n t i n e n t s ; t h e f a u n a l assemblages a r e l i s t e d f o l l o w i n g a l a t i t u d i n a l o r d e r a c c o r d i n g t o t h e sediment a r y b a s i n s from w h i c h t h e y were r e c o r d e d . Acknowledgments The a u t h o r w i s h e s t o e x p r e s s g r a t i t u d e t o t h e A r g e n t i n a N a t i o n a l C o u n c i l f o r S c i e n t i f i c and T e c h n i c a l R e s e a r c h , f o r t h e e c o n o m i c a l a i d i n t h e r e a l i z a t i o n of t h e p r e s e n t w o r k , and who g e n e r o u s l y p r o v i d e d t h e u s e of t h e J e o l c o JSM-U3 s c a n n i n g e l e c t r o n m i c r o s c o p e w i t h w h i c h some of t h e m i c r o g r a p h s were t a k e n . The w r i t e r i s a l s o g r a t e f u l t o t h e Buenos A i r e s U n i v e r s i t y , D e p a r t m e n t of Geology f o r p r o v i d i n g f a c i l i t i e s t o a c h i e v e t h e c o m p l e t i o n of t h i s work. The a u t h o r i s i n d e b t e d t o D r . K a r l Kr6nunelbein, K i e l Univ e r s i t y , who k i n d l y s u p p l i e d t h e o s t r a c o d e m a t e r i a l and t o D r . John N e a l e , H u l l U n i v e r s i t y who g e n e r o u s l y p h o t o g r a p h e d t h e mat e r i a l and s e n t t h e s t e r e o g r a p h i c p a i r e d p h o t o g r a p h s of t h e non-marine o s t r a c o d e s . P a r t i c u l a r thanks a r e a l s o expressed t o D r . Carlos A. R i n a l d i and t o M r . P a t r i c i o G a n d u g l i a , who g e n e r o u s l y h e l p e d i n
273
t h e c o n s t r u c t i o n of t h e s t r a t i g r a p h i c c h a r t s and r a n g e c h a r t s which i l l u s t r a t e t h i s p a p e r . The Lower C r e t a c e o u s non-marine o s t r a c o d e s a r e d e p o s i t e d a t t h e c o l l e c t i o n o f D r . K a r l Krsmmelbein a t t h e U n i v e r s i t y o f Kiel.
The i l l u s t r a t e d m a r i n e m a t e r i a l i s d e p o s i t e d i n t h e Labora t o r y of M i c r o p a l e o n t o l o g y , F a c u l t a d d e C i e n c i a s E x a c t a s y Natu r a l e s , Buenos A i r e s
,
University. S o u t h America
Lower C r e t a c e o u s I n t h e %ipe/Alagoas
B a s i n t h e B a i z o Sao F r a n c i s c o and
t h e S e r g i p e g r o u p s a r e d i s t i n g u i s h e d i n which s e v e r a l s u b g r o u p s and f o r m a t i o n s a r e r e c o g n i z e d by S c h a l l e r ( 1 9 6 9 ) ; some format i o n s a r e o n l y r e c o g n i z e d i n Alagoas o r i n S e r g i p e , a l t h o u g h some f o r m a t i o n s e x t e n d s o v e r bo.th a r e a s T e x t F i g . 1. W i t h i n t h e C o r u r i p e Subgroup s e v e r a l lower C r e t a c e o u s sequences a r e r e c o g n i z e d ; t h e C o r u r i p e Subgroup c o m p r i s e s t h e following Formations:
B a r r a d e I t i u b a , Rio P i t a n g a and i t s
e q u i v a l e n t s i n A l a g o a s : t h e Penedo and Morro Chavez F o r m a t i o n s , Coqueiro S e c o , s u b d i v i d e d i n s e v e r a l members, and P o n t a Verde. The C o r u r i p e Subgroup i s c o r r e l a t e d w i t h t h e B a h i a Supergroup of t h e Reconcavo/Tucano b a s i n , The B a r r a d e I t i u b a F o r m a t i o n i s composed of o l i v e and b l a c k c l a y s t o n e s w i t h f i n e i n t e r c a l a t i o n s of s a n d s t o n e s , s i l t s t o n e s and c a l c a r e o u s r o c k s . I n t h e B a r r a d e I t i u b a Formation S c h a l l e r ( 1 9 6 9 ) recogn i z e d t h e zones o f : Cypridea (Morininoides) c a n d e i e n s i s K r o m e l b e i n
= (003)
P a r a c y p r i d e a b r a s i l i e n s i s Krommelbein
= (004)
P a r a c y p r i d e a o b o v a t a o b o v a t a (Swain)
= (005)
C y p r i d e a ( M o r i n i n a ? ) b i b u l l a t a Wicher
=
(006)
P e t r o b r a s i a m a r f i n e n s i s Krommelbein and Cypridea ( S e b a s t i a n i t e s )
fida
Krommelbein
= (007)
S c h a l l e r ( 1 9 6 9 ) c o r r e l a t e s t h e Barra d e I t i u b a Formation w i t h t h e C a n d e i a s , M a r f i n and P o j u c a F o r m a t i o n s of t h e Reconcavo/Tucano B a s i n . The Penedo F o r m a t i o n i s composed of p o o r l y s o r t e d s a n d s t o n e s w i t h c l a y e y and s i ’ l t y i n t e r c a l a t i o n s , m a i n l y w h i t e c o l o u r e d ; i t c o n t a i n s o s t r a c o d e s of t h e zones ( 0 0 7 ) ,
( 0 0 6 ) and
274
( 0 0 5 ) ; i t i s c o r r e l a t e d by S c h a l l e r (%.
&.)
with t h e Marfin,
P o j u c o and Sao S e b a s t i a o F o r m a t i o n s of t h e Reconcavo/Tucano Basin. The Rio P i t a n g a F o r m a t i o n i s c o n g l o m e r a t i c w i t h c l a y e y and s i l t y i n t e r c a l a t i o n s ; i t c o m p r i s e s t h e o s t r a c o d e zones ( 0 0 7 ) , (006) and (005) o f S c h a l l e r ( 1 9 6 9 ) . The Morro d e Chaves F o r m a t i o n i s composed of l i g h t g r a y c a l c a r e o u s s t r a t a and c o q u i n o i d m a r l s w i t h brown c l a y s t o n e s i n tercalated.
I t i s c o r r e l a t e d w i t h t h e Zones ( 0 0 7 ) and ( 0 0 8 )
a n d , i n t h e P i a c a b u c u a r e a w i t h t h e zones ( 0 0 9 ) and ( 0 1 0 / 0 1 1 ) . The C o q u e i r o S e c o F o r m a t i o n i s m a i n l y composed of a l t e r n a t i n g of s a n d s t o n e s , c l a y s t o n e s , and s i l t s t o n e s ; i t c o n t a i n s t h e (008)
zone of S c h a l l e r ( 1 9 6 9 ) .
The o v e r l y i n g P o n t a Verde For-
mation d i d n o t y i e l d o s t r a c o d e s . I n t h e S e r g i p e / A l a g o a s B a s i n t h e non-marine s e q u e n c e s w e r e s u c c e e d e d by a n A t l a n t i c m a r i n e t r a n s g r e s s i o n which f o l l o w e d t h e p r i n c i p a l t e c t o n i c s t r u c t u r e s of t h e b a s i n ; t h e s e sequences began w i t h t h e t r a n s i t i o n a l Muribeca F o r m a t i o n which c o n t a i n s e v a p o r i t i c b e d s . The b r a c k i s h - m a r i n e f a c i e s of t h i s f o r m a t i o n a r e of A p t i a n a g e and younger p e r s i s t i n g t h e m a r i n e f a c i e s i n a c o n s e c u t i v e s t r a t i g r a p h i c s e q u e n c e i n t o Lower Eocene times. The Muribeca F o r m a t i o n o v e r l i e s t h e Sao F r a n c i s c o Group. I n t h e e n t i r e s e c t i o n b r a c k i s h water-marine o s t r a c o d e s of t h e genus C y t h e r i d e a ? and t h o s e r e c o r d e d by S c h a l l e r ( 1 9 6 9 ) o f Zones (010/011) o c c u r .
F o r t h e u p p e r Member S c h a l l e r
(z.
-
c i t . ) i n f e r r e d an Aptian age. The t r a n s i t i o n a l Muribeca F o r m a t i o n i s o v e r l a i d by t h e m a i n l y m a r i n e S e r g i p e Group. The S e r g i p e g r o u p i s s u b d i v i d e d i n t o s e v e r a l f o r m a t i o n s i n which some m a r i n e o s t r a c o d e s o c c u r . The R i a c h u e l o F o r m a t i o n i s composed of l i m e s t o n e s , c l a y s t o n e s and s i l t s t o n e s w i t h b e n t h o n i c and p l a n k t o n i c f o r a m i n i fera.
Kr&unelbein and Wenger ( 1 9 6 6 ) a s s i g n e d a n Upper A p t i a n
t o A l b i a n a g e t o t h i s f o r m a t i o n b a s e d on o s t r a c o d a .
Kr6mmel-
b e i n (196 ) c i t e d from t h e R i a c h u e l o F o r m a t i o n S e r g i p e l l a t r a n s a t l a n t i c a Krb'mmelbein and A r a c a j u i a b e n d e r i Krommelbein; b o t h s p e c i e s were a l s o found i n Gabon, A f r i c a .
BASINS STAGES
IECIFE/JOAC ESSOA BASH TINOCO. 1967 I
MAASTRICHTIAN
GRAYAYE Fn.
CAMPANIAN SANTONIAN CONlAClAN
BEBERIDE Fa
SERGIPE/ALAGOAS BASIN
REC~NCAVOI TUCANO BASIN
[SCHILLER ,19691
I V I A N A e l (111.19711
/I- I
MARITUBA Yb
GROUPS
FORMATION
JATOBA BASIN
NEUQUEN BASIN
RAUN. 19661
(01GREGOR10.1972I
SAN JORGE BASIN ESlLINDFERELL0,19721
JAGUEL
MEMBERS
ALLEN CANDELEROS
SAPUCARI Mb
COUODORO RlVADAVlA
TURONIAN CENOMANIAN
RAYOSO
ALBIAN
LlMAY
UPPER APTIAN
ORTlZ
YARIZAL PlCHl PlCUN LEU1
LOWER APTIAN RIO
CHIVES Fm.
sno
v)
P E N E W Fn.
w
PACIENCIA
ILHAS
AGRlO
POJUCA
SANTIAW ILHAS
V
0 W V
BARRA DE ITIUBA Fn
SALVADOR
a SANTO
YULICHINCD
3 B
CANDEIAS
CANDEIAS
AMARO ITAPARICA
-
UORAO 00 BARF YARACANGALI
a P
T e x t F i g . 1.
CARMEN
SEBASTI~O
LT v) W
0 > U
NEOCOMIAN
UlNA DEL
JDANE'
IASSACARA
AGUA GRAND
PUINTUCOVACA MUERTA
SERGI 7 \LIANCA
P O 2 0 0129
7
S t r a t i g r a p h i c u n i t s o f S o u t h American S e d i m e n t a r y B a s i n s ,
276 The S e r g i p e / A l a g o a s s e q u e n c e s m i g h t b e c o r r e l a t e d w i t h t h e Reconcavo/Tucano b e t w e e n Zones
Basin s t r a t a ; t h e r e e x i s t s c l o s e c o r r e l a t i o n
( 0 0 1 ) t o (007) of t h e S e r g i p e / A l a g o a s B a s i n w i t h
t h e Zones R - 1 0
t o R . 2 o f t h e Reconcavo/Tucano
Basin although
some s p e c i e s a r e l a c k i n g i n b o t h a r e a s ( S c h a l l e r , 1 9 6 9 ) . B a s i n t h e basement i s
I n t h e t e c t o n i c Reconcavo/Tucano
composed o f i g n e o u s , m e t a m o r p h i c and s e d i m e n t a r y r o c k s o f P r e c a m b r i a n and P a l e o z o i c a g e . Viana
( 1 9 7 1 ) s u b d i v i d e d t h e B a h l a S u p e r g r o u p i n t o t h e Bro-
t a s , S a n t o Amaro, I l h a s and M a s s a c a r i g r o u p s ( T e x t F i g . 1 ) . The Dom J o a o , R i o d a S e r r a , A r a t u , B u r a c i c a , J i q u i A and A l a g o a s S t a g e s c o n s t i t u t e t h e Reconcavo S e r i e s w h i c h , e x c l u d i n g t h e Alagoas S t a g e , c o r r e s p o n d t o t h e rock s t r a t i g r a p h i c Bahla Supergroup.
The J u r a s s i c - C r e t a c e o u s b o u n d a r y i s n o t c l e a r l y
defined i n t h i s basin. Viana e t a l .
( 1 9 7 1 ) r e c o g n i z e d s e v e r a l o s t r a c o d e z o n e s and
s u b z o n e s which c h a r a c t e r i z e t h e Reconcavo S e r i e s i n t h i s b a s i n . Many a u t h o r s h a v e worked o u t t h e a r e a and c o n t r i b u t e d w i t h v a r i o u s d a t a a b o u t t h e o s t r a c o d e f a u n a and i t s u t i l i t y a s z o n a l m a r k e r s s u c h a s Swain ( 1 9 4 6 ) , Wicher
( 1 9 5 9 ) , P i n t o and
S a n g u i n e t t i ( 1 9 6 2 ) , Krommelbein ( 1 9 6 2 )
,
M u l l e r ( 1 9 6 6 ) , Krommel-
b e i n and Wenger ( 1 9 6 6 ) , B r a u n ( 1 9 6 6 ) , V i a n a (1969) and Viana e t a l .
(1971).
(1967) , S c h a l l e r
Other organic groups, such a s
p o l l e n , w e r e a l s o i n t e n s i v e l y s t u d i e d (Weber 1 9 6 3 , 1 9 6 4 ) . Useful o s t r a c o d e g u i d e f o s s i l s have been r e c o r d e d i n t h i s b a s i n w h i c h , a l s o a r e p r e s e n t i n n e i g h b o r i n q b a s i n s and i n Af-
rica. The Reconcavo a r e a of t h e b a s i n i s l o c a t e d n e a r t h e p r e s e n t c o a s t w h i c h p r e s e n t s f a u l t s t r u c t u r e s w h e r e a s t h e Tucano a r e a e x t e n d s i n l a n d ; c o n s e q u e n t l y more f a c i e s c h a n g e s a r e f o u n d i n t h e Reconcavo b a s i n . The Upper J u r a s s i c B r o t a s Group o v e r l i e s t h e P r e c a m b r i a n c r y s t a l l i n e complex and P a l e o z o i c r o c k s . The S e r g i F o r m a t i o n o f t h e B r o t a s Group shows a g r a d a t i o n a l c o n t a c t w i t h t h e u n d e r l y i n g A l i a n c a F o r m a t i o n a s w e l l as w i t h t h e o v e r l y i n g I t a p a r i c a
Bormation of t h e S a n t o Amaro Group of a s u p p o s e d Lower C r e t a ceous a g e .
The a g e of t h e I t a p a r i c a F o r m a t i o n w a s d a t e d by
means of o s t r a c o d e s . The S a n t o Amaro Group c o m p r i s e s t h e I t a p a r i c a and C a n d e i a s formations. The I t a p a r i c a F o r m a t i o n i s composed of v a r i e d c o l o r e d c l a y s t o n e s and f o s s i l i f e r o u s o l i v e g r e y s i l t s t o n e s . o c c u r s i n Reconcavo and s o u t h e r n Tucano.
The f o r m a t i o n
O s t r a c o d e s of t h e
g p r i d e a k e g e l i Wicher s u b z o n e a r e t h e m o s t s i g n i f i c a n t . The C a n d e i a s F o r m a t i o n i s composed of c l a y s t o n e s , s i l t s t o n e s i n t e r c a l a t e d by l i m e s t o n e s , d o l o m i t e s and t h i c k b e d s of s a n d s t o n e s ; t h e member d e n o m i n a t i o n s c o r r e s p o n d s t o t h e s e f a c i a l changes.
The most i m p o r t a n t o s t r a c o d e s a r e t h o s e o f t h e
zones o f : Theriosynoecum v a r i e t u b e r a t u m Grekoff and K r o m e l b e i n C y p r i d e a ( M o r i n i n o i d e s ) c a n d e i e n s i s Krommelbein and P a r a c y p r i d e a b r a s i l i e n s i s Krommelbein The C a n d e i a s F o r m a t i o n i s c o r r e l a t e d by Viana e t a l .
(1971)
with t h e lower p a r t of t h e Barra d e I t i u b a F o r m a t i o n of t h e Sergipe/Alagoas B a s i n . The S a l v a d o r F o r m a t i o n i s composed of f a n g l o m e r a t e s of c r y s t a l l i n e , c a l c a r e o u s , m e t a s a n d s t o n e s and m e t a s i l t s t o n e s .
It
i s l a t e r a l l y i n t e r d i g i t a t e d w i t h t h e S a n t o Amaro, I l h a s and Massacari Groups and p o s s e s s e s t h e same o s t r a c o d e c o n t e n t a s t h e f o r m a t i o n s w i t h which i t i s i n t e r f i n g e r e d .
Viana e t a l .
2.)
(2, c o r r e l a t e t h e S a l v a d o r F o r m a t i o n w i t h t h e Rio P i t a n ga Formation of t h e S e r g i p e / A l a g o a s B a s i n . The I l h a s Group c o m p r i s e s t h e Marfim and P o j u c a F o r m a t i o n s . The Marfim F o r m a t i o n i s l o c a l i z e d i n t h e s u b s u r f a c e ;
i t i s com-
posed of l i g h t g r e y t o g r e e n i s h f i n e g r a i n e d t o s i l t i c sandstones.
I n t h e Reconcavo area t h e Marfim F o r m a t i o n i s l a t e r a l -
l y i n t e r d i g i t a t e d w i t h t h e S a l v a d o r Formation.
The c o n t a c t
with t h e o v e r l y i n g P o j u c a F o r m a t i o n i s g r a d a t i o n a l . The most i m p o r t a n t o s t r a c o d e s are: P a r a c y p r i -
dea b r a s i l i e n s i s
Krommelbein, C y p r i d e a d r o m e d a r i u s K r o m e l b e i n ,
Cypridea s a l v a d o r i e n s i s K r o m e l b e i n and Candona? g r e g a r i a Krommelbein, The Marfim F o r m a t i o n i s c o r r e l a t e d by Viana e t a l .
(1971)
with t h e m i d d l e p a r t of t h e Barra d e I t i u b a F o r m a t i o n of t h e
278
S e r g i p e / Alagoas B a s i n . The main l i t h o l o g i c a l f e a t u r e s of t h e P o j u c a F o r m a t i o n a r e the varied composition greenish gray sandstones, claystones, s i l t s t o n e s and brown o o l i t i c l i m e s t o n e s which sometimes g r a d e i n t o arenaceous limestones.
The f o r m a t i o n p r e s e n t s good s t r a -
t i f i c a t i o n , with r i p p l e narks.
The c o n t a c t w i t h t h e o v e r l y i n g
Sao S e b a s t i a o F o r m a t i o n i s g r a d a t i o n a l . The most s i g n i f i c a n t o s t r a c o d e s s p e c i e s b e l o n g t o t h e b i o zones (Viana e t a l . ,
1970) :
P a r a c y p r i d e a o b o v a t a o b o v a t a Swain and C y p r i d e a ( M o r i n i n a ? ) b i b u l l a t a b i b u l l a t a Wicher The Massacara Group c o m p r i s e s t h e f o r m e r l y d e s i g n a t e d I l h a s F o r m a t i o n and t h e S a o S e b a s t i a o F o r m a t i o n .
The Sao
S e b a s t i a o F o r m a t i o n i s composed of r e d d i s h - y e l l o w f i n e t o c o a r s e s a n d s t o n e s , f r i a b l e , w i t h i n t e r c a l a t i o n s of s i l t y c l a y s t o n e s , The u p p e r p a r t i s i n some a r e a s c o n g l o m e r a t i c . The s i g n i f i c a n t o s t r a c o d e s are:
P e t r o b r a s i a m a r f i n e n s i s Krommel-
b e i n , c g r i a c i n a c o r i a c e a Krommelbein, C y p r i d e a ( S e b a s t i a n i t e s ? ) s o s t e n s i s s o s t e n s i s Krommelbein and C y p r i d e a ( S e b a s t i a n i t e s )
--
f i d a f i d a Krommelbein.
Viana e t a l .
( 1 9 7 1 ) c o r r e l a t e t h e Sao
S e b a s t i a o F o r m a t i o n w i t h p a r t of t h e Penedo F o r m a t i o n i n t h e Sergipe/Alagoas B a s i n . The M a r i z a l F o r m a t i o n i s composed of c o a r s e s a n d s t o n e s and c o n g l o m e r a t e s i n t e r c a l a t e d by s i l t s t o n e s , c l a y s t o n e s and l i m e stones,
This rock-stratigraphic u n i t i s a l s o recognized i n t h e
JatobA and M i r a n d i b a b a s i n s .
Viana e t a l . ,
(1971) correlate
t h i s f o r m a t i o n w i t h t h e Carmopolis Member of t h e Muribeca Form a t i o n of t h e S e r g i p e / A l a g o a s B a s i n . Viana e t a l . ,
( 1 9 7 1 ) b a s e d on t h e o s t r a c o d e c o n t e n t o f t h e
s e v e r a l non-marine s t a g e s of t h e Reconcavo S e r i e s p r o p o s e d t h e f o l l o w i n g C r e t a c e o u s o s t r a c o d e z o n e s , which a l s o a r e s u b d i v i d e d i n s e v e r a l subzones i n a s c e n d i n g o r d e r of a g e ( T e x t . F i g . Rio d a S e r r a S t a g e :
1):
RT-002
-
Theriosynoecum v a r i e t u b e r a t u m
RT-003
-
Cypridea(M0rininoides)candeiensis Paracypridea obovata obovata
RT-004
Aratu Stage:
RT-006
-
Buracica Stage:
RT-007
-
RT-005
Paracypridea b r a s i l i e n s i s C r i d e a (Morinina?) b i b u l l a t a 3hiiiiata Coriacina coriacea
279
RT-008
- Cypridea (Sebastianites) fida
minor Jiqui: Stage : RT-009 - Petrobrasia diversicostata Moura ( 1 9 7 2 ) described additional new species from the Reconcavo/Tucano Basin. Some of the new species occur in Africa, such as Cypridea cf. C. primaria Grekoff and Kr6mmelbein in the transitional Lower Cocobeach Series; and in other areas of Brazil such as: Cypridea (Sebastianites) fida minor, Cypridea (Pseudocypridina) fabeolata and Cypridea riojoanensis which occur in the Sergipe/ Alagoas Basin. In the interior region of Brazil a northwestern extension of the Reconcavo/Tucano Basin, the so-called Jatobi Basin, the Araripe Basin, conspicuous in its gypsum deposits, and the "Mirandiba" Basin were studied by Braun (1966). In the Jatobi Basin the Alianca (Upper Jurassic?-Lower Cretaceous), Sergi, Candeias, Ilhas (=Marfim and Pojuca formations of the Reconcavo/Tucano Basin), Sao Sebastiao, Marizal, Santana and Exu formations were recognized. The Candeias Formation overlies unconformably the Sergi Formation; for the Candeias Formation the zones of Cypridea opifera Krhmelbein Cypridea (Morininoides) candeiensis Krb'mmelbein
(z.
Cypridea ambigua KrGmmelbein were proposed by Braun cit.). The Ilhas Formation contains the zones of: Paracypridea obovata obovata Swain gpridea (Morinina) bilbullata Paracypridea brasiliensis Krdmmelbein The Aptian-Albian? Sao Sebastiao Formation in their lower section yields the zones of: Cypridea (Sebastianites) spp. Coriacina coriacea Krommelbein and Paracypridea quadrirugosa The upper Sao Sebastiao and Marizal Formation are nonfossiliferous whereas the Santana Formation yielded species of the genera Candonopsis, Paraschuleridea, Heterocypris and gsulcocypris.
-
280
In the Araripe Basin or Chapada the Araripe region (Aptian -Albian) Beurlen (1963) presented a stratigraphic sequence for this area; the section was subdivided into the Crato, Coriri, Missao Velha, Santana and Exu formations. Braun (2. G.) compares the partly evaporitic Santana Formation with the marine Riachuelo Formation of the Sergipe/ Alagoas Basin; faunally it seems similar to the Cod0 Formation in the northern located Maranhao Basin. The ostracode assemblage is composed of Candonopsis spp., Paraschuleridea spp., Heterocypris sp., and Bisulcocypris sp. In addition Bate (1971) recorded from the Aptian-Albian Santana Formation of Serra do Araripe , Ceari , northern Brasil , phosphatized ostracods; they belong to the freshwater Subfamily Cypridinae, Family Cyprididae. In northern Brazil Braun (1966) mentioned, from the MaranI hao Basin, the Cod0 Formation of Aptian-Albian age; it is correlative with the marine Riachuelo Formation of the Sergipe/ Alagoas Basin and with the Santana Formation of the Araripe region. Although Braun noted the ostracode similarities between the Santana Formation and the Cod6 Formation, the present writer has no knowledge of the ostracode content of the Cod6 Formation. In the "Rio de Peixe Basin," Paraiba, Braun (1966) found Darwinula sp. and Cypridea vulgaris Krommelbein; Braun assigned the strata to the Berriasian and correlated with the Lower Ilhas Group (-Marfim Formation) of the Reconcavo/Tucano Basin. From the western Argentina Neuqu&n Basin Musacchio (1971) recorded from La Amarga Group (Pichi Pith Leuf; Formation) non-marine ostracodes. In the Cerro China Muerta
--
area the non-marine La Amarga Group overlies concordantly and gradually the marine Agrio Formation; the Pichi P i c h LeufA Formation is mainly composed of marls. Although La Amarga Group was considered of Aptian-Albian age (Digregorio, 1972) , Musacchio (1971) based on charophytes assigned the strata of the basal La Amarga Group to the Barremiari. In the same basin Musacchio (1975) describes from the Rayoso Formation the ostracode Rayosoana quilimalensis Musacchio. Although the Rayoso Formation has been considered to be Cenoma-
281
n i a n ( D i g r e g o r l o , 1 9 7 2 ) , Musacchio ( 1 9 7 5 )
,
b a s e d on a f f i n i t i e s
of t h e new genus and i t s s p e c i e s , a s s i g n e d t h e l e v e l s i n which Rayosoana q u i l i m a l e n s i s was found t o t h e u p p e r p a r t of t h e Low-
er C r e t a c e o u s . I n t h e w e s t e r n p a r t of t h e San J o r g e B a s i n Musachio and C h e b l i (1975) r e c o r d e d from t h e c o n t i n e n t a l Chubut Group "Wealdian" t y p e o s t r a c o d e a s s e m b l a g e ; i t comes from t h e Chubut Group (Gorro F r i g i o F o r m a t i o n ) . Musacchio (1975) d e s c r i b e d t h e s e s p e c i e s . The P a s o d e I n d i o s l o c a l i t y r e p r e s e n t s t h e s o u t h e r n m o s t l o c a l i t y i n which "Wealdian t y p e " o s t r a c o d e s have been f o u n d . Musacchio (2. found a f f i n i t i e s w i t h t h e N o r t h e r n
e.)
Hemisphere a s s o c i a t i o n , p r i n c i p a l l y from t h o s e d e s c r i b e d from t h e R o c a l l o s a Mountains; Musacchio
e.) assigned
(9.
t o the
assemblage a n A p t i a n a g e . I n A r g e n t i n a m a r i n e extra-Andean Lower C r e t a c e o u s sequences a r e mentioned from t h e NeuquAn and A u s t r a l b a s i n s .
In the
NeuquAn B a s i n t h e A g r i o F o r m a t i o n was d a t e d a s H a u t e r i v i a n Barremian b a s e d on c o n t a i n e d ammonites.
The A g r i o F o r m a t i o n
c o n t a i n s m a r i n e o s t r a c o d e s which a r e b e i n g s t u d i e d .
In the
Austral Basin marine sedimentation took place s i n c e Valanginian
times. The f i r s t m a r i n e o s t r a c o d e , Novocythere s a n t a c r u c e a n a was r e c o r d e d by Malumian, Masiuk and Ross1 d e G a r c i a ( 1 9 7 2 ) from s u b s u r f a c e s t r a t a o f A p t i a n - A l b i a n a g e . Upper C r e t a c e o u s I n S o u t h America a l t h o u g h Upper C r e t a c e o u s s e d i m e n t s a r e recorded i n s e v e r a l basins, data about t h e ostracode content are scarce. Upper C r e t a c e o u s s e q u e n c e s a r e found i n t h e B r a z i l i a n Foz d e Amazonas, B a r r e i r i n h a s , R e c i f e / J o a o P e s s o a , S e r g i p e / A l a g o a s , A r a r i p e and i n t h e A r g e n t i n a NeuquAn, C o l o r a d o and A u s t r a l basins. I n t h e Amazonas B a s i n m a r i n e s e d i m e n t a t i o n i s r e p r e s e n t e d by t h e Upper C r e t a c e o u s t o P a l e o c e n e Amapi F o r m a t i o n ( S h a l l e r
et al., Petri
1 9 7 1 ) ; t h i s f o r m a t i o n i s m a i n l y made up of c a r b o n a t e s .
(1954) r e c o r d e d r a r e C r e t a c e o u s F o r a m i n i f e r a , b u t up t o
now o s t r a c o d e s have n o t b e e n s t u d i e d . The B a r r e i r i n h a s B a s i n c o n t a i n s c o n t i n u o u s s e d i m e n t a r y sequences which r a n g e i n a g e from A p t i a n t o t h e T e r t i a r y (Lima,
282
1 9 7 3 ) ; a s y e t o s t r a c o d e s have n o t been r e c o r d e d . I n t h e A r a r i p e B a s i n t h e Exu F o r m a t i o n d o e s n o t c o n t a i n f o s s i l s (Braun, 1 9 6 6 ) . I n t h e S e r g i p e / A l a g o a s B a s i n i n d i s c o r d a n c e o v e r t h e mar i n e R i a c h u e l o F o r m a t i o n s e d i m e n t s of Middle A l b i a n ? t o Lower S a n t o n i a n a r e r e p r e s e n t e d by t h e C o t i n g u i b a F o r m a t i o n . The S a p u c a r i Member, made up of c a r b o n a t e s , y i e l d e d t h e ostracode Brachycythere (Brachycythere) sapucariensis K r & m e l b e i n , of T u r o n i a n a g e .
Krijmmelbein ( 1 9 6 4 ) c o r r e l a t e s t h e S o u t h
American S a p u c a r i F o r m a t i o n w i t h t h e A f r i c a n S i b a n g F o r m a t i o n and t h e Milango F o r m a t i o n , of p r o b a b l y C o n i a c i a n a g e , b a s e d on t h e p r e s e n c e of B r a c h y c y t h e r e ( B r a c h y c y t h e r e ) s a p u c a r i e n s i s i n these formations. The C o t i n g u i b a F o r m a t i o n i s o v e r l a i n by t h e m a r i n e P i a c a bucu F o r m a t i o n , which r a n g e i n a g e from t h e Campanian t o t h e Lower Eocene.
Although p l a n k t o n i c F o r a m i n i f e r a were s t u d i e d
i n d e t a i l ( S c h a l l e r , 1 9 6 9 ) no o s t r a c o d e s h a v e been r e c o r d e d from t h i s f o r m a t i o n . F o r t h e Calumbi Member Kr&mnelbein and Wenger ( 1 9 6 6 ) a s s i g n e d a n Upper C a m p a n i a n - M a a s t r i c h t i a n a g e .
For t h e Marituba
Member ( S c h a l l e r 1 9 6 9 ) a s s i g n e d a n a g e which r a n g e from Campa-
n i a n t o Lower Eocene. I n t h e R e c i f e / J o a o P e s s o a B a s i n t h e Upper C r e t a c e o u s i s r e p r e s e n t e d by t h e f l u v i a l B e b e r i d e and m a r i n e Gramame format i o n s o f t h e P a r a i b a Group: t h e f o r m e r i s u n f o s s i l i f e r o u s . The m a r i n e Gramame F o r m a t i o n r e p r e s e n t s a t r a n s g r e s s i v e c a l c a r e o u s f a c i e s (Mabesone, Tinoco and Cou’itinho,
1968).
The
Gramame F o r m a t i o n y i e l d e d m i c r o f o s s i l s s u c h a s f o r a m i n i f e r a and ostracodes.
The C r e t a c e o u s f o r a m i n i f e r a 1 a s s e m b l a g e s w e r e s t u -
d i e d by Tinoco ( 1 9 6 7 ) : t h e o n l y known o s t r a c o d e s a r e s p e c i e s o f t h e g e n r e C y t h e r e l l a , C y t h e r o p t e r o n and C y t h e r e i s ( T i n o c o , 1967).
The o s t r a c o d e a s s e m b l a g e i s b e i n g s t u d i e d .
I n A r g e n t i n a Upper C r e t a c e o u s non-marine s e d i m e n t s e x t e n d s o v e r a l a r g e a r e a of t h e NeuquLn B a s i n , and form t h e b a s a l b e d s of t h e J a g c e l F o r m a t i o n and e q u i v a l e n t s s u c h a s t h e Lower memb e r of t h e H u a n t r a i - c o F o r m a t i o n ( B e r t e l s , 1 9 7 2 ) .
These have
y i e l d e d f r e s h - w a t e r o s t r a c o d e s s u c h a s Candona? h u a n t r a i c o e n s i s B e r t e l s , I l y o c y p r i s t r i e b e l i B e r t e l s and Wolburgia? n e o c r e t a c e a
Bertels.
283
A f t e r t h i s e p i s o d e t h e s e a e n c r o a c h e d upon t h e C o l o r a d o and N e u q u h b a s i n s .
The f r e s h w a t e r and p o l y h a l i n e d e p o s i t s
belong t o t h e Lower J a g s e l l a n S u b s t a g e w h e r e a s t h e n e a r l y m a r i n e t o t h e Upper J a g u e l i a n S u b s t a g e . From t h e J a g i i e l F o r m a t i o n B e r t e l s
( 1 9 7 4 , 1975) d e s c r i b e d
from t h e Neuqukn B a s i n Upper C r e t a c e o u s (Lower M a a s t r i c h t i a n ? ) and M i d d l e M a a s t r i c h t i a n o s t r a c o d e a s s e m b l a g e s .
The a s s i g n e d
age was b a s e d upon p l a n k t o n i c F o r a m i n i f e r a ( B e r t e l s , 1 9 7 2 ) . The d e s c r i b e d s p e c i e s a r e o n l y known from A r g e n t i n a , b u t they show c l o s e s i m i l a r i t i e s w i t h w e s t e r n A f r i c a n s p e c i e s , esp e c i a l l y w i t h t h o s e b e l o n g i n g t o Veenia
cytherels d e s c r i b e d
by Reyment
( N i g e r i a ) and A&-
( 1 9 6 0 ) and A p o s t o l e s c u
(1961,
1963). Africa Neocomian Neocomian non-marine
d e p o s i t s a r e found i n w e s t e r n A f r i c a
i n t h e Gabon, Congo and Angola s e d i m e n t a r y b a s i n s and i n S o u t h Africa. The c o n t i n e n t a l s e d i m e n t s a r e o f l a g o o n a l , l a c u s t r i n e and e s t u a r i n e o r i g i n ; t h e s e sequences reach l a r g e t h i c k n e s s ; varyi n g from 3 0 0 0 t o 6 0 0 0 meters.
They m o s t l y b e l o n g t o t h e Coco-
beach S e r i e s o f Gabon and were d e n o m i n a t e d by G r e k o f f and ( 1 9 6 7 ) as t h e West A f r i c a n W e a l d i a n S e r i e s . I n t h e Gabon B a s i n t h e Cocobeach S y s t e m ( d e K l a s z and
Krammelbein
M i c h o l e t , 1 9 7 0 ; L e C a l v e z , d e K l a s z and B r u n , 1 9 7 4 ) i s o f E a r l y C r e t a c e o u s a g e c o m p r i s i n g t h e Neocomian
(Berrisian t o Barremi-
a n ) and A p t i a n S t a g e s ; t h e i r s t r a t a e x t e n d t o t h e s o u t h i n t o t h e Congo B a s i n and A n g o l a . The Cocobeach S y s t e m o v e r l i e s t h e P r e c a m b r i a n b a s e m e n t and crops o u t i n a n o r t h w e s t - s o u t h e a s t
trend.
The Lower Cocobeach S u b s y s t e m i s s e p a r a t e d from t h e Upper Cocobeach S u b s y s t e m by a n i n p o r t a n t d i s c o r d a n c e c o n n e c t e d w i t h major t e c t o n i c e v e n t s t h a t marked a p h a s e o f t h e s e p a r a t i o n of A f r i c a and S o u t h A m e r i c a . The Lower Cocobeach S u b s y s t e m c o m p r i s e s s e v e r a l s t r a t i graphic u n i t s
(Text Fig. 2 ) .
and b r a c k i s h - w a t e r
ostracodes
These u n i t s y i e l d e d f r e s h - w a t e r
.
The o s t r a c o d e s a s s e m b l a g e s f r o m t h e Lower Cocobeach Sub-
284
s y s t e m w e r e s t u d i e d by Krgmmelbein ( 1 9 6 6 ) , G r e k o f f and Krommelbein (1967)
,
Grosdidier (1967)
,
Viana ( 1 9 6 6 ) , and d e K l a s z and
Micholet ( 1 9 7 0 ) . D e K l a s z and M i c h o l e t ( 1 9 7 0 ) p r o p o s e d t h e S e r i e s o f Kango,
Remboue, N'Toum, Gamba and S a l i f e r e D'Ezanga f o r t h e nomenclat u r e u s e d p r e v i o u s l y by G r e k o f f and Krtjmmelbein ( 1 9 6 7 ) . Kr%nmelbein ( 1 9 6 6 ) compares t h e K & l & l & ( l o w e r Kango S e r i e s ) b e d s o s t r a c o d e a s s o c i a t i o n w i t h t h a t of I t a p a r i c a and b a s a l C a n d e i a s of t h e Reconcavo/Tucano b a s i n ; i n a d d i t i o n V i a n a ( 1 9 6 6 ) f o u n d common components i n t h e a s s e m b l a g e s of t h e o v e r -
l y i n g Bokouk and Bikoumk b e d s ( u p p e r Kango S e r i e s ) . D e K l a s z and M i c h o l e t ( 1 9 7 0 ) n o t e t h a t , e x c l u d i n g t h e new
s p e c i e s , o s t r a c o d e s of t h e Bikoum;! Member a r e common t o t h o s e of t h e Candeias Formation i n B r a z i l . KrEmmelbein ( 1 9 6 6 )
, Viana
( 1 9 6 6 ) and G r e k o f f and Krommel-
b e i n ( 1 9 6 7 ) c o r r e l a t e d t h e Fourou P l a g e Member ( l o w e r Remboue S e r i e s ) w i t h t h e lower I l h a s .
The f a u n a l a s s e m b l a g e s o f t h e
Lob& Member and B i f o u n e Member ( m i d d l e Remboue S e r i e s ) a r e s i m i l a r ; Kr6mmelbein ( 1 9 6 6 ) c o r r e l a t e d b o t h members, a s w e l l a s t h e o v e r l y i n g Moundunga Member, w i t h t h e l o w e r p a r t of t h e Upp e r I l h a s Group of t h e Reconcavo/Tucano B a s i n i n B r a z i l , wherea s G r e k o f f and Krtimmelbein ( 1 9 6 7 ) compared t h e Moundounga and t h e u n d e r l y i n g B i f o u n e members w i t h t h e u p p e r p a r t o f t h e u p p e r I l h a s Group,
I n t h e Moundunga Member,
( u p p e r Remboue S e r i e s ) ,
Grekoff and Kr6mmelbein 4 1 9 6 7 ) i n a d d i t i o n found a d i f f e r e n t a s s e m b l a g e f r o m t h a t of t h e u n d e r l y i n g member and t h e a p p e a r a n c e of C y p r i d e a n a n o r o s t r a t a G r e k o f f and KrEmmelbein and t h e g e n u s P e t r o b r a s i a which announce t h e a p p e a r a n c e o f t h e o v e r l y i n g Benguie assemblage. The o s t r a c o d e a s s e m b l a g e of t h e Benguik and N'Gwanz6 Memb e r s (N'Toum S e r i e s ) c o n t a i n s e v e r a l s p e c i e s of P e t r o b r a s i a and Cypridea ( S e b a s t i a n i t G s )
.
The a s s e m b l a g e s r e c o r d e d by K r h m e l b e i n ( 1 9 6 6 ) b e l o n g t o p a r t of t h e Moundunga Member and t h e Bengui6 Member; t h e s p e -
c i e s c i t e d by Viana ( 1 9 6 6 ) and G r o s d i d i e r ( 1 9 6 7 ) came from t h e lower P a r t of t h e Bengui;! Member. The N'Gwanze Member f a u n a d o e s n o t show g r e a t d i f f e r e n c e s from t h a t of t h e u n d e r l y i n g Bengui;! Member.
TExt
I-i:.
2. S t r a t i y r a p k i c
tini
ts o f ilfrican sedinlentary basirls.
J.t
m cn
286
G r e k o f f and Kr5mmelbein ( 1 9 6 7 ) r e c o r d e d s p e c i e s common t o b o t h t h e B r a z i l and Gabon a r e a s ; t h e s e a u t h o r s made p r e c i s e c o r r e l a t i o n s w i t h t h e B r a z i l i a n Wealdian t y p e s e d i m e n t s , espec i a l l y w i t h t h o s e of t h e Reconcavo/Tucano and S e r g i p e / A l a g o a s Basins. I n t h e Congo B a s i n G r e k o f f
(1957) s t u d i e d o u t c r o p s a l o n g
t h e L u a l a b a R i v e r and b o r e h o l e s a m p l e s from Samba; b o t h a r e a s a r e l o c a t e d i n t h e n o r t h e r n p a r t of t h e Congo B a s i n .
The se-
quences y i e l d e d r i c h f a u n a l assemblages which w e r e c o n s i d e r e d t o b e u p p e r J u r a s s i c and l o w e r C r e t a c e o u s a g e ,
The same a u t h o r
(Grekoff, 1 9 6 0 ) analyzed a d d i t i o n a l l y t h e upper Jurassic-lower C r e t a c e o u s o s t r a c o d e a s s o c i a t i o n s of s a m p l e s from s e v e r a l reg i o n s of t h e Congo B a s i n s u c h a s : Yakoko and D e k e s e .
Grekoff
N'Deke, E l i s a b e t h a , U b a n g i ,
( 1 9 5 7 ) made a n e x h a u s t i v e a n a l y s i s
of t h e o s t r a c o d e a s s e m b l a g e of t h e Congo B a s i n L u a l a b a S e r i e s . The L u a l a b a S e r i e s i s s u b d i v i d e d i n t o t h e S t a n l e y v i l l e S t a g e and t h e L o i a S t a g e .
The L u a l a b a S e r i e s i s composed o f s e v e r a l
b e d s numbered from 1 t o 1 6 ( b a s e t o t o p ) .
The b e d s 1 t o 1 4
c o n s t i t u t e t h e S t a n l e y v i l l e S t a g e w h e r e a s t h e bed 1 5 i s a s s i g n e d t o t h e Loia S t a g e . c o n s i d e r e d by G r e k o f f
The H o r i z o n 1 6 o r Bokungu Beds were
(1957) a s a n i n d e p e n d e n t e n t i t y ; it i s
composed of r e d c l a y s t o n e s and s a n d s t o n e s .
The a s s i g n e d a g e
was l a t e J u r a s s i c f o r t h e S t a n l e y v i l l e " S t a g e " and lower C r e t a ceous f o r t h e Loia "Stage" of t h e Lualaba Series.
The f a c i e s
which y i e l d e d o s t r a c o d e s a r e of f r e s h - w a t e r o r b r a c k i s h - w a t e r e n v i r o n m e n t and h a v e b e e n d e s c r i b e d by M a r l i & r e ( 1 9 4 8 , 1 9 5 0 ) and Grekoff
(1957, 1960).
The L u a l a b a S e r i e s a s s e m b l a g e was f o u n d t o c o n t a i n z o n a l markers. The Samba (Congo) b o r e h o l e a s s o c i a t i o n ( G r e k o f f , 1 9 5 7 ) i s composed of 27 s p e c i e s o f f r e s h - and b r a c k i s h ? - w a t e r o s t r a c o d e s Grekoff
( 1 9 5 7 ) found s e v e r a l s p e c i e s common t o b o t h t h e
L u a l a b a S e r i e s and t h e Samba S e r i e s (1 t o 5 ) , and c o r r e l a t e d t h e h o r i z o n s 3 t o 1 4 of L u a l a b a w i t h t h e Samba S e r i e s 3 ( S t a n l e y v i l l e S t a g e ) and h o r i z o n s 1 5 and 1 5 ' o f L u a l a b a w i t h S e r i e s 3 and 4 of Samba ( L o i a S t a g e ) .
I n a d d i t i o n Grekoff
f o u n d s i x s p e c i e s common t o Gabon.
These a r e :
(1957)
Paracypridea
o b o v a t a , Cypridea d i m i n u t a , P a r a c y p r i a l o n g a e n s i s l i l o e n s i s ,
287
B r a d y c y p r i s r o t u n d a , M e t a c y p r i s s p . 390, D a r w i n u l a l e g u m i n e l l a and D a r w i n u l a o b l o n g a . The b o r e h o l e s s e q u e n c e s a t N'Deke, E l i s a b e t h a , U b a n g i , Yakoko and Dekese were c o n s i d e r e d by G r e k o f f
(1960) c o r r e l a t i v e
w i t h h o r i z o n s 1 5 and 1 6 of t h e L u a l a b a S e r i e s . Based on t h e above m e n t i o n e d o s t r a c o d e c o n t e n t G r e k o f f ( 1 9 5 7 ) c o r r e l a t e d l e v e l s 2 t o 1 5 ' of t h e L u a l a b a S e r i e s and
s e r i e s 3 t o 5 from t h e b o r e h o l e a t Samba w i t h t h e M i d d l e CocoThe Cocobeach System i s nowadays a l m o s t a l -
beach subsystem.
ways i n c l u d e d i n t h e Lower C r e t a c e o u s ; t h e r e f o r e t h e s e q u e n c e s
of t h e L u a l a b a S e r i e s would c o r r e s p o n d t o t h e Gabonese Bokou& Member of t h e Mayanga F o r m a t i o n (Kango S e r i e s ) up t o t h e Beng u i 6 Member of t h e Bingone F o r m a t i o n (N'Toum S e r i e s ) .
The
S t a n l e y v i l l e Stage could be approximately equivalent t o t h e Kango S e r i e s w h e r e a s t h e L o i a S t a g e m i g h t b e a p p r o x i m a t e l y e q u i v a l e n t t o t h e Remboue S e r i e s , b a s e d on G r e k o f f ' s d a t a (1957).
P i n t o and S a n g u i n e t t i ( 1 9 6 1 ) b a s e d on t h e p r e s e n c e of M e t a c y p r i s c o n s o b r i n a J o n e s , and v e r t e b r a t e s , b e l i e v e d t h a t h o r i z o n 1 5 ' of t h e L u a l a b a S e r i e s ( u p p e r p a r t of t h e L o l a S t a g e ) and s e r i e s 3 o f Samba a r e of A l b i a n a g e . I n t h e Congo B a s i n G r e k o f f d i a n ( C r e t a c e o u s ) assemblages : and a n u p p e r W e a l d i a n ( = B )
( 1 9 6 0 ) d i s t i n g u i s h e d two Weal-
a l o w e r W e a l d i a n (=A) "complex"
"complex."
The boundary b e t w e e n A
and B w a s n o t c l e a r l y d i s t i n g u i s h a b l e and some s p e c i e s a r e found i n b o t h a s s o c i a t i o n s . G r o s d i d i e r ( 1 9 6 7 ) s t u d i e d o s t r a c o d e s a s s e m b l a g e s from Gabon and Congo.
I n r e f e r e n c e t o t h e Congo a s s e m b l a g e G r o s d i d i e r
p o i n t e d o u t t h a t t h e y do n o t show c l o s e s i m i l a r i t i e s w i t h t h e B r a z i l i a n f a u n a ; o n t h e o t h e r hand G r o s d i d i e r (%. c &.)
be-
l i e v e d t h a t t h e y a r e s t r a t i g r a p h i c a l l y h i g h e r t h a n t h o s e of t h e m i d d l e Sao S e b a s t i a o F o r m a t i o n of t h e B r a z i l i a n Reconcavo/Tucan0 B a s i n . The new s p e c i e s d e s c r i b e d by G r o s d i d i e r ( 1 9 6 7 ) from t h e Congo B a s i n a r e C y p r i d e a l o a n g o G r o s d i d i e r , C y p r i d e a bapounou G r o s d i d i e r , Cypridea
G r o s d i d i e r , Damonella?
&-
o u s s o u e n s i s G r o s d i d i e r , "Reconcavona" b a t i k e G r o s d i d i e r , R h i n o c y p r i s k r o e m m e l b e i n i G r o s d i d i e r and O r t h o n o t a c y t h e r e m v i l i
288
Grosdidier. This association is equivalent to the N'Gwanze Member of the Bingone Formation of Gabon. In the Angola Cuanza Basin "Wealdian" type sediments were deposited (Cuvo Formation) (Brognon and Verrier, 19661, but the sequence does not contain non-marine ostracodes as far as the writer knows. In South Africa Dingle (1969, 1971) recorded for the first time Lower Cretaceous marine ostracodes from the Uitenhage region and from the continental margin Agulhas Bank. The Neocomian marine strata of the Uitenhage Group are confined to the coastal areas in the east and southeast of the country. The classic subdivision of the Cretaceous section of the Uitenhage Basin (Text.Fig. 2 ) is composed of the Enon Conglomerate, followed by Variegated Marls and Wood Beds (Wealdian facies but without ostracodes) and then by the Sundays River Formation. This Cretaceous section overlies the Cape Sequence and is covered by Tertiary strata (Dingle, 1969). Dingle (1969) found the age of the Sundays River Formation to be at least partly Upper Valanginian. From the continental margin Agulhas Bank (Dingle 1971) recorded marine ostracodes. The samples studied by Dingle were dated by means of Foraminifera and ammonite fragments as Barremian?. Aptian In western Africa, Aptian sequences are recognized in Gambia, Nigeria, Gabon, Congo, Angola and South Africa. Apostolescu (1963) studied subsurface samples from Gambia and Senegal. In both regions the samples yielded marine ostracodes. In Gambia from the borehole at Sara-Kunda 1, Apostolescu (1963) recorded ostracodes from several levels which range in age from Aptian to Maastrichtian. Based on the ostracodal content of the Sara-Kunda 1 sequence Apostolescu proposed five faunizones. Faunizone 1 comprises the Aptian-Albian interval; the Aptian Stage was dated by means on Choffatella decipiens Schl. It only yielded the marine ostracode genus Schuleridea.
289
I n t h e Gabon B a s i n t h e f l u v i o - l a c u s t r i n e Gamba S e r i e s i s s e p a r a t e d from t h e Lower Cocobeach S u b s y s t e m by a d i s c o r d a n c e which marks a n i m p o r t a n t p h a s e i n t h e s e p a r a t i o n o f A f r i c a and S o u t h America ( d e K l a s z and M i c h o l e t , 1 9 7 0 ) ; t h e f i r s t m a r i n e i n f l u e n c e s a r e m a n i f e s t e d i n t h e Gamba/Ezanga b o u n d a r y e v i denced by l a m i n a t e d c l a y s t o n e s , w i t h s a l t c r y s t a l s , of t h e S c h i s t e s d e Cocobeach Member w h i c h , o t h e r w i s e , c o n s t i t u t e s t h e b a s e o f t h e Ezanga S e r i e s . I n t h e Gamba S e r i e s o s t r a c o d e s a r e r a r e ; t h e y were r e c o r d ed by Krdmmelbein
( 1 9 6 6 ) a n d by G r o s d i d i e r
(1966).
The Ezanga S e r i e s is t r a n s i t i o n a l b e t w e e n t h e f l u v i o l a c u s t r i n e b e d s and t h o s e d o m i n a n t l y m a r i n e .
This series has
not y i e l d e d o s t r a c o d e s . I n t h e Congo B a s i n t h e Exanga? S t a g e , w h i c h i s l a t e r a l l y e q u i v a l e n t t o t h e s a l t - b e a r i n g s t r a t a i n Gabon d i d n o t y i e l d o s t r a c o d e s a s f a r a s t h e w r i t e r knows. I n t h e Angola Cuanza B a s i n d u r i n g A p t i a n t i m e s , s u b s i d e n c e of t h e c e n t r a l p a r t of t h e b a s i n c o n t r o l l e d t h e c y c l i c a l d e p o s i t i o n of c a r b o n a t e - e v a p o r i t i c
b e d s (Brognon a n d Verrier
,
1966)
.
No o s t r a c o d e s h a v e b e e n r e c o r d e d from t h e s e f a c i e s . From t h e c o n t i n e n t a l m a r g i n o f S o u t h A f r i c a D i n g l e ( 1 9 7 1 ) recorded marine Aptian o s t r a c o d e s .
The s a m p l e s came from t h e
Agulhas Bank and y i e l d e d o s t r a c o d e s . The a g e o f t h e s a m p l e w a s d a t e d by t h e f o r a m i n i f e r E p i s t o -
& m
(Brotzenia) alveata.
Although t h e a g e r a n g e s from Barre-
mian t o A l b i a n , a n A p t i a n a g e i s p r e f e r r e d by D i n g l e . Albian A l b i a n d e p o s i t s a r e w i d e l y d i s t r i b u t e d i n West A f r i c a : t h e y a r e e n c o u n t e r e d i n Gambia, N i g e r i a , Cameroon, Gabon, Congo, Angola a n d S o u t h A f r i c a . I n West A f r i c a , A l b i a n a n d y o u n g e r C r e t a c e o u s s e q u e n c e s a r e m o s t l y o f m a r i n e o r i g i n ; t h e s e s t r a t a a r e d a t e d by means o f a m m o n i t i e s and p l a n k t o n i c F o r a m i n i f e r a . From Gambia A p o s t o l e s c u
( 1 9 6 3 ) r e c o r d e d , from s u b s u r f a c e
s a m p l e s o f t h e Sara-Kunda w e l l , t h e f o l l o w i n g o s t r a c o d e s : thonotacythere sp. A . , Asciocythere sp. A . ,
sp. B.,
Cytheropteron sp. A . ,
or-
sp. B.,
and C y t h e r e i s s p . A .
T h i s a s s e m b l a g e b e l o n g t o t h e F a u n i z o n e 1 of A p t i a n - A l b i a n age.
290
I n t h e N i g e r i a B a s i n and i n Cameroon, t h e f i r s t d a t a b l e m a r i n e sequence b e g i n s i n A l b i a n t i m e s b u t o s t r a c o d e s have n o t b e e n s t u d i e d from t h e s e a r e a s . I n t h e Gabon B a s i n m a r i n e A l b i a n d e p o s i t s a r e r e p r e s e n t e d by t h e MadiAla S e r i e s o f t h e N'Komi System.
According t o d e
K l a s z and M i c h o l e t ( 1 9 7 0 ) t h e MadiAla S e r i e s r e p r e s e n t a p e r i o d of g r a d u a l r e g r e s s i o n .
This S e r i e s , p a r t i c u l a r l y i t s upper
p a r t , i s s e c u r e l y d a t e d by ammonites
(Reyment, 1 9 6 6 ) ; t h e o s -
t r a c o d e s a r e b e i n g s t u d i e d ( d e K l a s z and M i c h o l e t , 1 9 7 0 ) . I n t h e Congo B a s i n Upper C r e t a c e o u s s t r a t a a r e m e n t i o n e d by G r e k o f f
( 1 9 6 0 ) from t h e s o u t h w e s t e r n p a r t o f t h e B a s i n ; t h e y
a r e r e p r e s e n t e d by t h e Kwango S e r i e s . ed i n t o two s t a g e s : Stage.
This Series i s subdivid-
t h e l o w e r I n z i a S t a g e and t h e u p p e r Nsele
A c c o r d i n g t o G r e k o f f i n some areas t h e r e i s a d i s c o n -
t i n u i t y b e t w e e n t h e L u a l a b a S e r i e s and t h e o v e r l y i n g Kwango S e r i e s ; i n t h e t y p e a r e a a t Kwango t h e L u a l a b a S e r i e s i s un-
known.
The I n z i a S t a g e was s u b d i v i d e d i n t o f i v e h o r i z o n s a c -
cording t o t h e i r l i t h o l o t i c a l f e a t u r e s .
The f o s s i l i f e r o u s ho-
r i z o n i s Kwango 5 from h o r i z o n 5 of t h e I n z i a S t a g e ; t h i s l e v e l y i e l d e d f r e s h - w a t e r o s t r a c o d e s s u c h a s : Ilyoc y p r i s compressa G r e k o f f , I l y o c y p r i s l u z u b i e n s i s G r e k o f f , Cypridea ( C y p r i d e a ) h i l a r i e n s i s G r e k o f f , Darwinula kwangoensis G r e k o f f , A f r o c y t h e r e ? K 536 G r e k o f f and " M e t a c y p r i s " K 3099 Grekoff.
Taking i n t o a c c o u n t t h a t t h e o v e r l y i n g N s e l e S t a g e
c o n t a i n s t h e h o r i z o n i n w h i c h P . d e S a i n t S e i n e found a b i r d b e l o n g i n g t o t h e F a m i l y D e r c e t i d a e , known o n l y from t h e Cenomanian, it i s t o s u s p e c t t h a t t h e u n d e r l y i n g I n z i a S t a g e c o u l d b e of A l b i a n a g e . I n t h e Angola Cuanza B a s i n t h e A l b i a n S t a g e s e q u e n c e s a r e r e p r e s e n t e d by t h e T u e n z a , Catumbela and Q u i s s o n d e F o r m a t i o n s (Brognon and V e r r i e r , 1 9 6 6 ) , b u t o s t r a c o d e s h a v e n o t b e e n rec o r d e d from them. Cenomanian Cenomanian s t r a t a a r e f o u n d i n Gambia, N i g e r i a , Cameroon, Gaban, Congo B a s i n and Angola. The West A f r i c a n Cenomanian was c h a r a c t e r i z e d by a p e r i o d of g e n e r a l r e g r e s s i o n i n a l l c o a s t a l b a s i n s
(Hourq, 1 9 6 4 ) .
I n Gambia A p o s t o l e s c u ( 1 9 6 3 ) s t u d i e d t h e s u b s u r f a c e seq u e n c e s ; a c o m p l e t e s e c t i o n of m a r i n e s t r a t a r a n g i n g i n a g e
291
from A p t i a n t o M a a s t r i c h t i a n a r e f o u n d .
The Cenomanian t o
T u r o n i a n s e q u e n c e s y i e l d e d a n a s s e m b l a g e of o s t r a c o d e s which b e l o n g t o t h e F a u n i z o n e 2 of A p o s t o l e s c u ( 1 9 6 3 ) I n t h e N i g e r i a B a s i n w e l l d a t e d m a r i n e Cenomanian i s o n l y known from C a l a b a r ; it y i e l d e d ammonites and m i c r o f o s s i l s ; t h e l a s t o n e s h a v e n o t y e t b e e n s t u d i e d (Reyment, 1 9 6 6 ) .
From
Cameroon Reyment ( 1 9 6 5 ) m e n t i o n e d t h e Mundeck S a n d s t o n e Format i o n of Albian-Cenomanian a g e . c o r d e d from t h i s f o r m a t i o n .
No m i c r o f o s s i l s h a v e b e e n re-
I n t h e Gabon B a s i n t h e Cenomanian
i s a p e r i o d of r e g r e s s i o n i n which r e d f a c i e s h a v e a l a r g e geo-
graphic extend.
The s t r a t a of t h e Cap Lopez S e r i e s c o n t a i n a
c h a r a c t e r i s t i c microfauna.
Up t o d a t e no o s t r a c o d e s h a v e b e e n
studied. I n t h e Congo B a s i n t h e l i t h o l o g i c s e q u e n c e o f t h e Nsele S t a g e i s composed of v e r y f i n e s a n d s t o n e w i t h c l a y s t o n e s a t t h e i r b a s e and l o c a l l y w i t h s i l i c i f i e d b e d s .
The o s t r a c o d e
f o s s i l i f e r o u s h o r i z o n i s a t t h e b a s e of t h e N s e l e S t a g e . horizon provided t h e following:
This
D o l e r o c y p r i s k i n k o e n s i s Grek-
o f f and P a r a c y p r i a makawensis G r e k o f f .
The o s t r a c o d e s came
from h o r i z o n s i n w h i c h P . d e S a i n t S e i n e found b i r d s b e l o n g i n g t o t h e F a m i l y D e r c e t i d a e known from t h e Cenomanian i n o t h e r areas (Grekoff, 1 9 6 0 ) .
I t i s important t o note t h a t almost a l l
C r e t a c e o u s o s t r a c o d e s p e c i e s o f t h e Congo B a s i n a r e d i f f e r e n t t o t h o s e found i n a d j a c e n t b a s i n s .
I n t h e Angola Cuanza B a s i n
d u r i n g Cenomanian t i m e s t h e Cab0 Led0 F o r m a t i o n was d e p o s i t e d . The l i t h o l o g i c c o m p o s i t i o n of t h i s f o r m a t i o n a r e s i l t y s h a l e s w i t h i n t e r c a l a t e d l i m e s t o n e s (Brognon and V e r r i e r , 1 9 6 6 ) . s t r a t a c o n t a i n o s t r a c o d e s and f o r a m i n i f e r s
The
(Reyment, 1 9 6 6 1 , b u t
t h e o s t r a c o d e s have n o t been s t u d i e d . Turonian The T u r o n i a n was a p e r i o d of e x t e n s i v e t r a n s g r e s s i o n i n West A f r i c a ; s t r a t a of t h i s a g e a r e r e c o r d e d from Gambia, N i -
g e r i a , Cameroon, Gabon and Angola.
Reyment ( 1 9 6 6 ) m e n t i o n e d
s e q u e n c e s o f t h i s a g e from N i g e r R e p u b l i c , Chad and I v o r y C o a s t I n Gambia s u b s u r f a c e b o r e h o l e s a m p l e s a t S a r a Kunda 1 y i e l d e d T u r o n i a n o s t r a c o d e s ; t h e y were r e c o r d e d by A p o s t o l e s c u (1963)
.
292
I n N i g e r i a t h e Turonian s e a s p r e a d o v e r a wide area; duri n g t h i s t i m e s e q u e n c e s o f t h e Eze-Aku
S h a l e and Makurdi Forma-
t i o n i n t h e Benue V a l l e y i n s o u t h e a s t e r n N i g e r i a , and from t h e P i n d i g a , G o n g i l a , Dukul and J e s s u f o r m a t i o n s i n N o r t h e a s t e r n N i g e r i a were d e p o s i t e d (Reyment, 1 9 6 6 ) ; d a t a b l e ammonite f a u n a s a r e common i n t h e s e s t r a t a , The Lower T u r o n i a n Eze-Aku S h a l e of N i g e r i a y i e l d e d o s t r a c o d e s and b e n t h o n i c F o r a m i n i f e r a which h a v e n o t y e t b e e n g i v e n d e t a i l e d s t u d y (Reyment, 1 9 6 5 ) . I n Cameroon t h e T u r o n i a n , C o n i a c i a n , S a n t o n i a n , and Camp a n i a n s t r a t a o f t h e Mungo R i v e r F o r m a t i o n a r e w e l l d a t e d w i t h ammonites and p l a n k t o n i c F o r a m i n i f e r a
(Belmonte, 1 9 6 6 ) .
Ostra-
c o d e s have n o t b e e n s t u d i e d up t o t h e p r e s e n t , t o t h e w r i t e r ' s knowledge. I n t h e Gabon B a s i n t h e m a r i n e T u r o n i a n A z i l ;
Series yield(Reyment, 1966) ,
ed g u i d e p l a n k t o n i c F o r a m i n i f e r a and o s t r a c o d e s b u t t h e o s t r a c o d e s have n o t y e t been s t u d i e d .
I n Angola t h e s e d i m e n t s o f t h e Lower Itombe F o r m a t i o n were d e p o s i t e d d u r i n g T u r o n i a n t i m e s , c o n d i t i o n s b e i n g much t h e same as d u r i n g Cenomanian t i m e . N o o s t r a c o d s have b e e n r e c o r d e d from t h e Lower Itombe F o r m a t i o n . Senonian, sensu l a t o I n t h e S e n o n i a n , s . l., a r e i n c l u d e d t h e C o n i a c i a n , Sant o n i a n and Campanian S t a g e s . I n Gambia s u b s u r f a c e s e q u e n c e s y i e l d e d C o n i a c i a n t o C a m panian ostracode assemblages.
The C o n i a c i a n a s -
semblage c o n s t i t u t e s A p o s t o l e s c u ' s F a u n i z o n e 3 , w h e r e a s h i s F a u n i z o n e 4 c o m p r i s e s S a n t o n i a n t o Campanian s t r a t a An i m p o r t a n t c o n t r i b u t i o n was made by A p o s t o l e s c u ( 1 9 6 1 ) ; t h i s a u t h o r s t u d i e d Upper C r e t a c e o u s o s t r a c o d e f a u n a s from S C n e g a l , I v o r y C o a s t , Togo, Dahomey and M a l i .
The s t u d i e d
C r e t a c e o u s s e q u e n c e s r a n g e i n a g e from t h e S e n o n i a n t o t h e M a a s t r i c h t i a n , t h e M a a s t r i c h t i a n b e i n g o n l y found i n t h e TogoDahomey b a s i n . I n t h e S e n e g a l B a s i n t h e S e n o n i a n s e q u e n c e s a r e found i n s u b s u r f a c e ; t h e s t r a t a a r e m a i n l y composed of c l a y s and i n t e r c a l a t e d sandstones.
293
I n t h e I v o r y C o a s t B a s i n t h e S e n o n i a n o s t r a c o d e s come from The known s p e c i e s a r e c h a r a c t e r i s t i c bore h o l e s a t Yokobov;. from p r e - M a a s t r i c h t i a n
s t r a t a o f S e n e g a l , Cameroon and N i g e r i a
.
( i . e . S a n t o n i a n t o Campanian) The Dzodze a r e a o f Ghana, a t t h e b o r e h o l e a t Q u i b a y e i l d e d Campanian M a a s t r i c h t i a n o s t r a c o d e s . I n t h e N i g e r i a B a s i n S e n o n i a n , s.l., s t r a t a a r e w i d e s p r e a d Reyment ( 1 9 6 0 ) s t u d i e d t h e N i g e r i a n Upper C r e t a c e o u s Senonian and M a a s t r i c h t i a n o s t r a c o d a ; t h e s t u d i e d m a t e r i a l comes from o u t c r o p s and b o r e h o l e s from n o r t h e r n , w e s t e r n , and e a s t e r n Nigeria and from Cameroon. , The S a n t o n i a n S t a g e i s p r e s e n t i n t h e N i g e r i a B a s i n .
As
r e g a r d s t o t h e m i c r o p a l e o n t o l o g y of t h e N i g e r i a B a s i n , which was a t i m e of r e g r e s s i o n , s e v e r a l o s t r a c o d e s p e c i e s have been d e s c r i b e d from t h e Lamja S h a l e of n o r t h e a s t e r n N i g e r i a (Reyment, 1 9 6 0 ) and p o s s i b l e S a n t o n i a n l i m e s t o n e s i n Benue P r o v i n c e of c e n t r a l N i g e r i a ; from t h e Benua P r o v i n c e , Reyment (1966) cited Cytherella a u s t i n e n s i s Alexander, Brachycythere Reyment, B u n t o n i a vanmorkhoveni Reyment
.
From t h e Lam] a For-
mation Reyment ( 1 9 6 0 ) r e c o r d e d s p e c i e s o f O v o c y t h e r i d e a , Cythe r e l l a aus t i n e n s i s Alexander
,
M e t a c y t h e r o p t e r o n paganum (Rey-
ment) and B u n t o n i a s p p . Reyment s t a t e d t h a t t h e s e a s s o c i a t i o n s c o n t a i n many s p e c i e s i n common w i t h t h e E g y p t i a n Upper C r e t a ceous.
Undoubtedly, Campanian i s known i n N i g e r i a from t h e
Duala Embayment (Reyment, 1 9 6 5 ) . I n Cameroon t h e o n l y known S e n o n i a n s t r a t a came from t h e Mungo R i v e r F o r m a t i o n : t o t h e s e s t r a t a a Campanian a g e was assigned (Reyment, 1965) ,' b u t o s t r a c o d e s were n o t r e c o r d e d . I n t h e Gabon B a s i n t h e S e n o n i a n s t r a t a a r e r e p r e s e n t e d by t h e A n g u i l l e S e r i e s and P o i n t e C l a i r e t t e S e r i e s ; o s t r a c o d e s have n o t been c i t e d from t h e d e p o s i t s ( d e X l a s z and M i c h o l e t , 1970).
From t h e Congo B a s i n Reyment ( 1 9 6 5 ) r e c o r d e d S a n t o n i a n s t r a t a from t h e Vonso a r e a ( L e o p o l d v i l l e ) . N o o s t r a c o d e s have
294
been mentioned. In Angola the sedimentation continued during the Senonian. During this time the Itombe, Teba and Dio Dande formations were deposited. The sequences are dated by means of planktonic foraminifera; no ostracodes have been recorded. In South Africa from the Agulhas Bank, located at the continental margin Dingle (1971) recorded two ostracode species: ?Amphicytherura sp. and Brachycythere agulhasensis Dingle. The assigned age was early-middle Senonian based on the planktonic foraminifera Globotruncana spp. ex. gr. 5. marginata (Reuss) Maastrichtian In western Africa the Maastrichtian was marked by a wide transgression which, during its climax seems to have stretched across West Africa to North Africa (Reyment, 1966). Maastrichtian sequences were recorded from Gambia, Senegal, Ghana, Dahomey, Togo, Nigeria, Cameroon, Gabon, Angola and South Africa. From Gambia Apostolescu (1963) recorded from boreholes at Sara Kunda 1 an assemblage of Faunizone 5 of Apostolescu
.
(2. G.) .
In Senegal the borehole at Sangalkam yielded the Maastrichtian ostracode Bradleya vesiculosa?. In Ghana the section at Kuli Jabafi contains the following Maastrichtian ostracodes: Ovocytheridea nuda Grekoff, Veenia? ughelli Reyment, Veenia? varriensis Reyment and Brachycythere oguni Reyment (Apostolescu, 1961, 1963). Several borehole sequences from Togo and Dahomey such as those at Attitogon, Bopa, Issoba, Lokossa and Sehouk were studied by Apostolescu (1961). The Maastrichtian sections were found to be equivalent to the planktonic foraminifera1 Zone of Abathomphalus mayaroensis. They yielded the ostracode Brachycythere armata Reyment. In western Nigeria the oldest sediments, dated by a m o r nites, are of upper Maastrichtian age; they lie directly on the Precambrian basement. Reyment (1961) studied surface samples from the Aule River locality near Auchi and from subsurface at the localities of
29 5
Sbekevo and A r a r o m i .
Reyment ( 1 9 6 0 ) o b s e r v e d t h a t a l a r g e
number o f new s p e c i e s would r e f l e c t l o c a l d e v e l o p m e n t s . I n e a s t e r n N i g e r i a , during t h e M a a s t r i c h t i a n , coal-forming c o n d i t i o n s d e v e l o p e d i n t h e Anambra Embayment. I n t h i s region t h e c o a l m e a s u r e s e q u e n c e b e g i n s w i t h t h e m a r i n e Nkporo S h a l e , p a s s i n g upward i n t o t h e l a r g e l y non-marine Mamu F o r m a t i o n (Reyment, 1 9 6 6 ) . The c o a l l e v e l s may e x t e n d i n t o t h e Danian ( S t o l k , 1963). I n Gabon t h e M a a s t r i c h t i a n Ewongu& S e r i e s c o n t a i n s o n l y arenaceous F oram inifera I n Angola t h e M a a s t r i c h t i a n S t a g e i s r e p r e s e n t e d by t h e lower p a r t of t h e Rio Dande F o r m a t i o n .
No o s t r a c o d e s h a v e been
mentioned from t h e s e s t r a t a . From S o u t h A f r i c a D i n g l e ( 1 9 7 1 ) r e c o r d e d m i d d l e t o l a t e M a a s t r i c h t i a n a s s e m b l a g e s from t h e A g u l h a s Bank. Summary During t h e u p p e r J u r a s s i c and C r e t a c e o u s P e r i o d s t h e cont i n e n t a l s e d i m e n t a r y s e q u e n c e s which a c c u m u l a t e d on b o t h s i d e s of t h e b o r d e r l a - i d s o f t h e p r e s e n t A t l a n t i c Ocean a r e i n t i m a t e l y related t o tectonic events.
These t e c t o n i c e v e n t s began i n
l a t e J u r a s s i c t i m e s and c o n t i n u e d d u r i n g t h e C r e t a c e o u s and Cenozoic, p r o d u c i n g s e v e r a l t e c t o n i c s t r u c t u r e s s u c h a s f a u l t s , g r a b e n s and r i f t v a l l e y s which c o n t r o l l e d t h e s e d i m e n t a r y depos i t i o n a l areas. During t h e Neocomian f r e s h , b r a c k i s h , and m a r i n e d e p o s i t s a r e r e g i s t e r e d i n b o t h t h e S o u t h American and A f r i c a n c o n t i nents. I n t h e S o u t h American c o n t i n e n t t h e f r e s h and b r a c k i s h water sedimentary sequences are confi ned t o t h e n o r t h e a s t e r n a r e a of B r a z i l and A r g e n t i n a .
The s e d i m e n t a r y b a s i n s i n which
non-marine Neocomian s t r a t a a r e r e c o r d e d i n S o u t h America a r e the Sergipe/Alagoas
,
Reconcavo
(Tucano , J a t o b k and Almada ba-
s i n s i n B r a z i l ; i n A r g e n t i n a non-marine Neocomian i s found i n t h e Neuqu6n and San J o r g e s e d i m e n t a r y b a s i n s .
During t h e Neocomian m a r i n e i n f l u e n c e s a r e c o n f i n e d t o t h e s o u t h e r n m o s t S o u t h American a r e a s ; m a r i n e s e q u e n c e s o c c u r i n t h e N e u q u h and A u s t r a l b a s i n s .
296
In the African Continent non-marine strata are found in Cameroon, Gabon, Congo, Angola and South Africa. Marine Neocomian strata are also restricted to the southernmost area of Africa, being found in South Africa. In summary, Neocomian marine transgressions took place in regions where the present South American and African continents were not united to form the Gondwana Supercontinent in Lower Cretaceous time. The ostracodal assemblages of both continents show close relationships and a large part of the associations are formed by species common to both continents. Non-marine ostracodes are at this time useful zonal markers. Text Fig. 3 summarizes the marine and non-marine sedimentary basins during Neocomian times. During Aptian and Albian epochs, the first marine transgressions were registered; the transgression left its record in thick evaporitic beds which are common in Gabon, Congo, Angola in Africa and in the Sergipe/Alagoas and Reconcavo/Tucano basins in South America. After this event the sea encroached over some areas although not contemporaneously in all basins; the first transgression recorded by mainly marine ostracodes is of Aptian and Albian times in South America (Sergipe/Alagoas Basin - Riachuelo Formation), and in western Africa in Ivory Coast, Nigeria, Cameroon, Gabon and Angola. Common African and South American marine ostracodes are recorded during Aptian-Albian times. Although Upper Cretaceous sedimentation is recorded in several sedimentary basins, there are few data about the ostracoda1 content. During the Maastrichtian a wide transgression was registered in several South American and African basins. The ostracode assemblages from this Stage show endemic features. The known assemblages from western Africa and those from South America (Argentina) evidences a common origin as shown by the great similarities between the assemblages, especially by the presence of the genera Veenia (E?igeria),Anticythereis and Togoina in both continents.
291
-1I
rext Fic. 3.
Neocomiaii, m a r i n e a n d n o n - m a r i n e
60.
a.
I
1
sedimentary basins.
,
10.
I
AmTIAN-ALIIAM-YIIINE \*ON
~
-
*.G
\ ,
U W l l CRETACEOUS-
T e x t F i g . 4. t j p t i a n 9 Albian and Upper Cretaceous iiarine and non-marine sedimentary basins.
298
Text Fig. 4 synthesizes the marine and non-marine sedimentary basins during the Aptian-Albian and Upper Cretaceous times. References Apostolescu, V., 1961. Contribution a 1'6tude pal6ontologique (Ostracodes) et Stratigraphique des Bassins Cr&tac&s et Tertfaires de 1'Afrique occidentale. Rev. Inst. FranFais du Petr., v. XVI, no. 7-8, p. 779-867. 1963. Essai de Zonation par les Ostracodes dans le Cr&tak du Bassin du Shegal. Rev. Inst. Franyais du P&tr., v. XVIII, no. 12, p. 1675-1694. Bate, R. H., 1971. Phosphatized ostracods from the Cretaceous of Brazil, Nature, v. 230, no. 5293, p. 397-398. Belmonte, Y. C., 1966. Stratigraphie du Bassin S6dimentaire du Cameroon. Proc. 2nd W . African Micropal. Coll. Ibadan, 1966, P. Bertels, A,, 1972. Ostricodos de agua dulce del Miembro Inferior de la Formaci6n Huantrai-co (Maastrichtiano inferior) Provincia del Neuqu&n, Repdblica Argentina. Ameghiniana, T. IX, no. 2, p. 173-182. , 1974. Upper Cretaceous (lower Maastrichtian?) ostracodes from Argentina. Micropaleontology, v. 20, no. 4, p. 385-397. , 1975. Upper Cretaceous (middle Maastrichtian) ostracodes of Argentina. Micropaleontology, v. 21, no. 1, pp. 97130. Beurlen, K., 1963. Geologia e Estratigrafia da Chapada do Araripe. XVII Congreso Brasileiro de Geologia, Recife, P. Braun, P. G. O., 1966. Estragrafia dos sedimentos da parte interior da regiao nordeste do Brasil. Minist. Minas e Energia. Divi. Geol. e Mineral. Bol. no. 236, p. 5-79. Brognon, G. and Verrier, G., 1966. Tectonique et skdimentation dans le bassin du Cuanza (Angola). In: Bassins sedimentaires du littoral africain. I'ere partie: Littoral Atlantique. AsSOC. des Services G6ol. Africains., p. 207-252. de Klasz, I., and Micholet, J., 1970. Elements nouveaux concernant la Biostratigraphie du Bassin Gabonais. IV Colloque African Micropal., p. 109-141. Digregorio, J. H., 1972. Neuquen. En: Geologia Regional Argentina, Acad. Nac. Ciencias. Cordoba., p. 439-505. Dingle, R. V., 1969. Marine Neocomian Ostracoda from South Africa. Trans. roy. S O C . S. Afr., v. 38, p. 2, p. 139-163. , 1971. Some Cretaceous Ostracodal assemblages from the Agulhas Bank (South African continental margin). Trans. roy. SOC. S. Afr., v. 39, p. IV, p. 393-418. Grekoff, N., 1957. Ostracodes du Bassin du Congo. Annales du Muske Royal du Congo Belge, Ser. in-8, v. 19, p. 1-97. 1960. Ostracodes du Bassin du Congo. 11. CrktacC. Ann. I&s&e Royal Congo Belge. Ser. in-8, v. 3 5 , p. 1-70. Grekoff, N. and KrEmmelbein, K., 1967. Etude comparee des O s tracodes M&ozoYques continentaux des bassins Atlantiques: Serie de Cocobeach, Gabon et S&rie de Bahia, Brksil. Rev. Inst. Francais Petr., v. X X I , no. 9 , p . 1307-1353. Grosdidier , E. , 1967. Quelques ostracodes nouveaux de la S&ie Ant&-SalifLre ("Wealdienne") des bassins cottiers du Gabon et du Congo. Rev. Micropal., v. 10, no. 2, p. 107-118.
299
Hourq, V . , 1 9 6 4 . L e s g r a n d s t r a i t s d e l a g & o l o g i e d e s b a s s i n s c o t i e r s du g r o u p e & q u a t o r i a l . Symp. W . A f r . S e d . B a s . , 2 2 I n t e r n . C o n g r e s s G e o l . N e w D e l h i , p . 171-178. K r h n e l b e i n , K . , 1 9 6 4 a . O s t r a c o d e n a u s d e r m a r i n e n "KiistenK r e i d e " B r a s i l i e n s . 1: B r a c h y c y t h e r e ( B r a c h y c y t h e r e ) s a p u c a r i e n s i s n . s p . a u s dem Turonium. Senck. L e t h . , V . 4 5 , p . 489495. 1964b. Neue A r t e n d e r O s t r a c o d e n - G a t t u n g P a r a c y p r i d e a Swain atis d e r B a h i a - S e r i e d e s Reconcavo B a h i a n o ( O b e r j u r a ? ) U n t e r k r y i d e , Wealden-Fazies, NE-Brasilien. Bol. Paranense d e G e o g r a f i a . , n o s , 10-15, p . 139-160. , 1 9 6 4 c . Uber e i n i g e n e u e A r t e n d e r O s t r a c o d e n - G a t t u n g Reconcavona Kr&mmelbein 1962 a u s d e r N E - b r a s i l i a n i s c h e n B a h i a - S e r i e . Senck. L e t h . , v . 45, p . 29-41. , 1 9 6 6 a . On "Gondwana Wealden" O s t r a c o d a from NE B r a z i l and West A f r i c a . P r o c . 2nd. West A f r i c a n M i c r o p . C o l l . I b a d a n , p . 113-118. , 1 9 6 6 b . P r e l i m i n a r y r e m a r k s o n some m a r i n e C r e t a c e o u s Proceed. O s t r a c o d e s f r o m n o r t h e a s t e r n B r a z i l and West A f r i c a . 2nd. West A f r i c a n M i c r o p . C o l l . I b a d a n , 119-121. , 1967. O s t r a c o d e n a u s den marinen"Kikten-Kreide" B r a s i l i e n . 2 : S e r g i p e l l a t r a n s a t l a n t i c a n . g . n . s p . und c a j u i a b e n d e r i n . g . n . s p . a u s dem Ober-Aptium/Albium. S e n c k . L e t h . , v . 48, p . 525-533. Suy q u e l q u e s a n a l o g i e s K r & u n e l b e i n , K . , - a n d Wenger, R . , 1 9 6 6 . r e m a r q u a b l e s d a n s l e s m i c r o f a u n e s c r b t a c e e s du Gabon e t du B r b s i l o r i e n t a l B a h i a e t S e r g i p e , p . 193-196. L e C a l v e z ~ , Y . , d e K l a s z I . , and B r u n , L . , 1 9 7 4 . N o u v e l l e c o n t r i b u t i o n a l a c o n n a i s s a n c e d e s m i c r o f a u n e s du Gabon. Rev. E s pafiola d e M i c r o p a l . , v . V I , n o . - 3 , p . 381-400. R e g i b n e x t r a n d i n a d e ChuL e S t a , P . J . , and F e r e l l o , R . , 1 9 7 2 . b u t y n o r t e d e S a n t a Cruz. I n : Geologia Regional A r g e n t i n a , p c a d . Nac. C i e n c i a s C o r d o b a . , p . 601-653. Lima, E . C . , 1 9 7 3 . B i o e s t r a t i g r a f i a d a Bacia d e B a r r e i r i n h a s . A n a i s d o X X V I C o n g r e s o S O C . B r a s . G e o l . , v . 3 , p . 81-91. Mabesone, J . M . , T i n o c o , I . M . , and C o u t i n h o , P . N . , 1 9 6 8 . The M e s o z o i c - T e r t i a r y Boundary i n N o r t h e a s t e r n B r a z i l . P a l e o g e o graphy, Palaeoclimatology, Palaeoecology, v. 4 , p . 161-185. Marliere, R . , 1 9 4 8 . O s t r a c o d e s e t P h y l l o p o d e s du Syst&me du K a r r o ? a u Congo B e l g e . An. Mus. Roy. Congo B e l g e . T e r v u r e n . S c . G e o l . , v . 2 , p . 1-15. , 1 9 5 0 . O s t r a c o d e s , e L P h y l l o p o d e s du Systkme du Karr o o a u Congo B e l g e e t l e s r e g i o n s a v o i s i n a n t e s . An. M u S . Roy. Congo B e l g e . T e r v u r e n . S e r . i n - 8 . S c . G 6 O l . , , V . 6 , p . 1-43. Moura, J . A . , 1 9 7 2 . Algumas e s p & i e s e s u b e s p e c i e s n o v a s d e o s t r a c o d e s d a b a c i a Reconcavo/Tucano. B e o l . Tecn. P e t r o b r A s . R i o d e J a n e i r o , v . 1 5 , n o . 3 , p . 245-263. Musacchio, E . A.,1971 Hallazgo d e l gbnero C r i d e a (Ostracoda) en Argentina y consjderaciones e s t r a t i g r f i c a s sobre l a F o r m a c i 6 n L a Amarga ( C r e t a c i c o I n f . ) e n l a P r o v . d e Neuquen. A m e g h i n i a n a , T . V I I I , no. 2 , p . 105-125. M u s a c c h i o , E . A . , a n d F h e b l i , G . , 1 9 7 5 . O s t r i c o d o s no m a r i n o s y c a r o f i t a s d e l C r e t a c i c o I n f e r i o T e n l a s p r o v i n c i a s d e Chub u t y Neuquen, A r g e n t i n a . 1. O s t r a c o d o s y c a r o f i t a s d e l Grupo C h u b u t . 2 . Rayosoana q u i l i m a l e n s i s nov. g e n . nov. s p . d e l a F o r m a c i 6 n Rayoso, N e u q u h . A m e g h i n i a n a , T . X I I , no. 1, p . 7096.
,
e-
.
+
300
1954. F o r a m i n i f e r o s f o s s e i s da B a c i a do Maraj6. Univ. Sao P a u l o . F a c u l t . F i l o s o f i a , C i e n c i a s e L e t r a s . B o l . 1 7 6 , G e o l . , no. 11, p . 1-173. P i n t o , I . D . , and S a n g u i n e t t i , Y . T . , 1 9 6 1 . O b s e r v a c i o n e s on M e t a c y p r i s ( O s t r a c o d a ) from t h e Mesozoic of North America and A f r i c a . Esc. Geol. P . A l e g r e . Bol. 9 , p . 3-14. , 1 9 6 2 . A c o m p l e t e r e v i s i o n of t h e Genera B i s u l c o c y p r i s and Theriosynoecum ( O s t r a c o d a ) w i t h t h e world g e o g r a p h i c a l and s t r a t i g r a p h i c a l d i s t r i b u t i o n . Esc. Geol. P . A l e g r e , Pub. Esp: no. 4 , p. 1-165. Reyment, R . A . , 1 9 6 0 . S t u d i e s on N i g e r i a n Upper C r e t a c e o u s and Lower T e r t i a r y O s t r a c o d a . P a r t 1: Senonian and M a e s t r i c h t i a n O s t r a c o d a . Acta Univ. Stockholm. v . V I I , p . 1-238. , 1965. A s p e c t s of t h e Geology of N i g e r i a . The s t r a t i graphy of t h e C r e t a c e o u s and Cenozoic d e p o s i t s . I b a d a n Univ. P r e s s . , p . 1-145. , 1 9 6 6 . B r i e f Review of t h e s t r a t i g r a p h i c s e q u e n c e s of West A f r i c a (Angola t o S e n e g a l ) . P r o c e e d . 2nd West A f r . Microp. Coll. I b a d a n , p . 162-175. S c h a l l e r , H . , 1 9 6 9 . Revisao e s t r a t i g r i f i c a da b a c i a d e S e r g i p e Alagoas. Bol. Tec. P e t r o b r a s , Rio d e J a n e i r o , v. 1 2 , no. 1, p . 21-86. S c h a l l e r , H . , V a s c o n c e l o s , D . N I , and C a s t r o , J . C . , 1 9 7 1 . E s t r a t i g r a f i a p r e l i m i n a r d a Bacia s e d i m e n t a r da Poz do Rio Amaz o n a s . S O C . B r a s . Geol. Anais do XXV Congreso, v . 3 , p . 1 8 9 Petri, S.,
202.
Swain, F. M . , 1946. Middle Mesozoic non-marine o s t r a c o d e s . J o u r . P a l e o n t o l o g y , v . 2 0 , no. 6 , p . 543-555. Tinoco, I . de M . , 1 9 6 7 . Micropaleontologia da f a i x a sedimentar B o l . SOC. B r a s . Geol. 1 6 , no. c o s t e i r a Recife-Joao Pessoa 1, p . 81-85. V i a n a , C . G . , 1966. 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 of o s t r a c o d a i n t h e Bahia Supergroup ( B r a s i l ) , P r o c . 2nd. W . A f r . M i c r o p a l . C o l l . I b a d a n . p . 2 4 0 , 257. V i a n a , G . F . , d a Gama, E . G . , J r . , A r a u j o Simoes, J . , J o u r a , J . A . , dos R e i s F o n s e c a , J . , A l v e s , R . J., 1 9 7 1 . Revisao est r a t i g r a f i c a d a b a c i a Reconcavo/Tucano. Bol. Tec. P e t r o b r a s , v . 1 4 , no. 3 / 4 , p . 157-192. Plate I. (Stereographic paired.*photographs) F i g . 1 Cypridea, ( M o r i n i n o i d e s ) c a n d e i e n s i s Krommelbein, x 5 3 , a ) l a t e r a l view; b) d o r s a l view S a n t o Amaro Zone ( 0 0 3 ) . Occurs i n S e r g i p e / Alagoas J a t o b a and A f r i c a . F i g . 2 . Paracypridea b r a s i l i e n s i s Kr'&mmelbein, x45, a ) l a t e r a l view; b ) d o r s a l v i e w Zones ( 0 0 3 ) - ( 0 0 4 ) Occurs i n S e r g i p e / A l a g o a s Reconcavo/ Almada and A f r i c a . Fig. 3 Ilhasina torosa Tucano Kr'cimmelbein Zones ( 0 0 4 ) ( 0 0 5 ) - Upper S a n t o Ama-ower I l h a s , x68. Occurs i n Reconcavo/Tucano and A f r i c a , a ) l a t e r P a r a c y p r i d e a o b o v a t a oboa l view; b) d o r s a l view. Fig. 4 v s (Swain) , x38, 1 9 4 6 Zones ( 0 0 5 ) Lower J l h a s . O c c u r s x S e r g i p e / A l a g o a s - Reconcavo/Tucano J a t o b a and A f r i c a . a ) l a t e r a l view; b ) d o r s a l view. P l a t e 11. ( S t e r e o g r a p h i c p a i g e d p h o t o g r a p h s ) F i g . 1 - S a l v a d o r i e l l a r e d u n c a redunca Krommelbein, x52, 1963 Zone ( 0 0 6 ) Upper I l h a s . Occurs i n Reconcavo/Tucano and A f r i c a . a ) l a t e r a 1 view; b ) d o r s a l view. F i g 2 Parac r i d e a quadriru osa w e b e r i Kr&rmelbein, x31 , a ) i a t e r a r s a l view? Zones ( 0 0 6 ) - ( 0 0 7 ) . Upper I l h a s Lower Sao S e b a s t i a o . O c c u r s i n S e r g i p e / A l a g o a s and i n A f r i c a . F i g . 3 Petrobrasia
-
-
-
-
-
-
.
-
-
-
-
-
-
-
-
301
marfinensis Kr&nmelbein, 1962, x63, a) lateral view; b) dorsal view, Zone (007) - Sao Sebastiao. Occurs in Sergipe/ Alagoas and Reconcavo/Tucano. Fig. 4. Cypridea (Sebastianites) sostensis sostensis Krommelbein, x53, a) lateral viyw; b) dorsal view, Zones (007)-(008) - Sao Sebastiao - Jiquia. Occurs in Sergipe/Alagoas and Reconcavo/Tucano. Plate 111. Fig. 1 - NOVOC there santacruceana Malumian, Musiuk and Rossi de Garcia-tral Basin - Upper Aptian Albian. Fig. 2 - Alatac there? rocana Bertels, x84, Neuquen Basin - (Lower? &.aM J m a n o Stage. Fig. 3 Platycythereis? velata Bertels , x84 , Neuqukn Basin - (Lower? Maastrichtian) Jagueliano Staye. Fig. 4 - Veenia (Nigeria) inornata Bertels , xl00 , Neuquen Basin - (Middleaastrichtian) Jagceliano Stage. Fig. 5 - Anticythereis venusta Bertels, x 8 4 , Neuquen Basin - (Middle Maastrichtian) Jaglieliano Stage. Fig, 6 - Tumidoleberis australis Bertels , xl00 , NeuquAn Basin - (Middle Maastrichtian) Jagueliano Stage. Fig. 7 Wichmannella magna Bertels, x100, N e u q u h Basin - (Middle Maastrichtian) Jagieliano Stage. Fig. 8 - Veenia (Nfgeria) tumida Bertels , x100, N e u q u h Basin - (Middle Maastrichtian) Jagueliano Stage. Fig. 9 - Veenia (Ni eria) a uelensis Bertels , xl00 , Neuquen B a s i n T d d l e g M a a s t r j h jaguelensis Bertels, x100, Neuquen Basin - (Middle Maastrichtian) Jagceliano Staye. Ffg. i0 - Veenia (Nigeria) unctata Bertels, x100, Neuquen Basin - (Midd-astrichtian’Jagiieliano Stage.
302
PLATE
I
303
PLATE I 1
304
PLATE 111
305
MESOZOIC R A D I O L A R I A FROM THE A T L A N T I C BASIN A N D ITS BORDERLANDS Helen P. Foreman Department of Geology, Oberlin College, Oberlin, Ohio 44074
Abstract Mesozoic Radiolaria, primarily from Deep Sea Drilling cores, are surveyed. Maps indicate localities, ages, and usefulness of the occurrence. The new Arnphipyndax tylotus Zone is defined and the next lower Arnphipyndax enesseffi Zone is emended.
Introduction It has long been recognized that Radiolaria i n the Atlantic Basin are not as abundant or as well
preserved as those f r o m the Pacific (Murray, J., 1876). This has again been pointed out by Johnson
(1976) who suggested that the poor preservation, at least, may be due t o the greater amount of silica depleted detritus being deposited in the Atlantic Basin. Examination of Deep Sea Drilling Project Atlantic Basin Mesozoic cores again confirms this diagnosis. With only a few exceptions, paucity and poor preservation seem to be the rule.
Localities The following four maps of the Atlantic Basin and i t s borderlands indicate with circles the locations of Deep Sea Drilling Project Sites, together w i t h supplementary localities from which Mesozoic radiolarian bearing sediments have been recovered. The dark circles represent holes or land-based supplementary localities which contain sediments w i t h the occasionally well-preserved Radiolaria, or Radiolaria i n useful sequences. DSDP Legs 1 through 4, 10 through 1 5 , 4 0 and 41, are included in this survey. Although Legs
36 and 39 in the southwest Atlantic and Leg 44 along t h e east coast of the United States also recovered Mesozoic radiolarian bearing sediments, their results are not included because the Initial Reports have not yet been published. The localities on the maps (Figures 1-41 and the chart of occurrences (Figure 5) from which Mesozoic radiolarian bearing sediments have been recovered are listed below:
306 Deep Sea D r i l l i n g Project A t l a n t i c Basin localities: Leg 1 Leg 2
Site 4
24"28.68'NI 73'47.52'W
Site 5
24"43.59'N, 73"38.46'W
at depth of 5319.5 meters at depth of 5361 meters
Site9
32O46.4' N, 59O11.7' W
at d e p t h of 4965 meters
Leg 3
Site 13
06'02.4'
N, 18O13.71 'W
at depth of 4588 meters
Leg 4
Site 24
06" 16.5 8'S,3 0'5 3.46'W
at depth of 5148 meters
Site 28
2Oo35.19'N, 65'37.33'W
at depth of 5521 meters
Site 95
24'09.00'N. 86'23.85'W
a t depth o f 1633 meters
Site 97
23'53.05'NI 84'26.74'W
at d e p t h o f 2930 meters
Leg 10 Leg 11
Site 100
24'41.28".
73'47.95'W
a t depth of 3525 meters
Site 101
25'1 1.93".
74O26.31 'W
at depth of 4868 meters
Site 105
34"53.72'N, 69"10.40'W
at depth o f 5251 meters
Leg 12
Site 118
45'02.65'N, 09"00.63'W
at d e p t h o f 4901 meters
Leg 13
Site 120
36'41.39'N, 11'25.94'W
at d e p t h o f 1711 meters
Leg 14
Site 136
34'10.1 3".
16"18.19'W
at depth of 4169 meters
Site 138
25'55.37".
25"33.79'W
at depth o f 5288 meters
Site 140
21O44.97".
21'47.52'W
at depth o f 4483 meters
Site 144
09"27.23'N, 54'20.52'W
at depth of 2957 meters
Site 146
15'06.99'N. 69'22.67'W
at depth of 3949 meters
Site 150
14'30.69'N, 69"21.35'W
at depth of 4545 meters
Site 152
15'52.72'N, 74"36.47'W
at depth of 3899 meters
Site 153
l3"58.33'NI 72O26.08.W
at depth of 3932 meters
Site 361
35"03.97'S, 15'26.91 'E
at depth o f 4549 meters
Site 363
19"38.75'S, 09'02.80'E
at d e p t h of 2248 meters
Site 364
11'34.32's. 1 l"58.30'E
at depth of 2448 meters
Site 367
12O29.2' N,2O002.8' W
a t d e p t h of 4748 meters
Site 368
17O30.4' N,21"21.2' W
at depth o f 3366 meters
Site 369
26O35.5' N. 14'59.9' W
at depth o f 1752 meters
Leg 15
Leg 40
Leg 41
Non DSDP Atlantic Basin localities: L a m o n t V-18-129
41'42's. 56"35'W
at depth o f 2039 meters
L a m o n t V-17-144
40"34'S, 55OlO'W
at d e p t h of 2503 meters
Marginal A t l a n t i c Basin localities:
B191, Pre-Habana f o r m a t i o n Cuba (Foreman, 1968) RM312, Val Dorbia, Umbria, Italy (Foreman, 1968) Marac Well # 1, Trinidad (Foreman, 1968)
BR1025, Via Blanca f o r m a t i o n Cuba (Bt6nnimann & Rigassi, 1963) BR1028, Via Blanca (?) f o r m a t i o n Cuba (Bronnimann & Rigassi, 1963)
307
-
Figure 1. Berriasian Late Jurassic localities. These are, as would be expected, few, dependent as they are on location, and drilling expertise. All the material so far examined was poorly preserved with abundances generally rare, occasionally common. However, Holes 367 and 100 are shaded to indicate useful sequences. Legs 1, 11, and 41 sampled this sequence.
308
P
1
4-- I
I
II
0
c
if
-----I-----
.- .
I
NEOCOMIAN LOCAL ITlES (exclusive of Berriasian)
,
1
Figure 2. Neocomian localities, exclusive of Berriasian. Only at Site 101 of Leg 1 1 have Radiolaria i n significant quantities in a useful sequence been recovered. A t Site 105 there is a long sequence, but the Radiolaria are generally rare and poorly preserved.
The Serhocapsa rrachyosrraca Zone could be identified in Core 18 of Hole 105. This Zone could also be recognized in Core 29 of Hole 367. Holes 363 and 361 had, i n general, only very rare, very poor
Radiolaria, and no Neocornian Zones could be recognized. For the Neocornian, as for the Berriasian
- Late
Jurassic, only Legs 1, 1 1 , and 41 are represented, together with the very minor occurrences of Leg 40.
309
ns
9
P
A1
41
0137
I I
Figure 3. Cenomenian and late Early cretaceous localities. With a few additions the map for the Cenomanian and late Early Cretaceous is very similar t o that for the Neocomian with faunas generally poor and sparse. A n exception might be Site 5 where Radiolaria from an Albian core show moderate preservation. Localities from Legs 1, 10, 11, 13, 14,40, and 41 are identified.
31 0
FIGURE 4 LATE CRETACEOUS LOCALITIES (exclusive of Cenornanian)
Figure 4. Late Cretaceous localities. These are by far the most abundant. The material is in general much better preserved and the shaded localities indicate in almost all instances moderate to good preservation in either a single core or in a sequence of a few cores. No long sequences are known except those from the Carribean Leg 15 which are poorly preserved and were considered by the shipboard scientists to be redeposited. For this time, some excellent land based localities with very good preservation are known. A single CampaniadSantonian sample (8191) collected by Bermudez from Habana, Cuba (Foreman, 1966) and some samples (BR), Campanian/Maestrichtianfrom Br'dnnimann and Rigassi (1963) also from Cuba, and early Santonian Globorruncana concavara Zone, sample (M-1) from Trinidad (Foreman, 1968).
31 1 What emerges is a pattern which finds Radiolaria present rather consistently in sediments of late Late Cretaceous age, minor occurrences in the Cenomanian-Albian and the l a t e Neocomian, and again Radiolaria consistently present in the few holes that sampled the Early Neocomian/Late Jurassic (Tithonian-Kimmeridgian and possibly Oxfordian). This is in rather great contrast to the Pacific where Radiolaria in the late Late Cretaceous are generally lacking or when present poorly preserved with Maestrichtian Radiolaria completely missing. I n the Pacific fairly common but relatively poor early Late Cretaceous and late Early Cretaceous Radiolaria are recognized and rich sequences of Aptian and Late Neocomian Radiolaria are known. No Radiolaria older than Tithonian have been recovered from the Pacific.
Radiolarian zonation For some time now, three major groups of Late Cretaceous radiolarian fossils have been recognized: A MaestrichtiadLate Campanian fauna typically represented by the forms described by Foreman (1968) and recognized by Kozlova (1972) from Leg 14 of the Deep Sea Drilling Project; an Early Campanian fauna partly described by Pessagno (1963) from Puerto Rico and recognized in samples from Cuba collected by Bermudez, Bronnimann and Rigassi as well as samples from numerous Deep Sea Drilling localities; and a Cenomanian/Albian
fauna, elements of which have been variously described by
Dumitrica (1970), Aliev (19651, and Foreman (1975). In the same way three major groups of Early Cretaceous Radiolaria have in the last few years been recognized: the Albian/Cenomanian fauna mentioned above, a Barremian to Hauterivian or Valanginian fauna, and an Early Neocomian to Late Jurassic one. With the impetus provided by the long sequences recovered by the Deep Sea Drilling Project, a number of zonal schemes have recently been introduced in an attempt to break down this very coarse, very inadequate division. However, the radiolarian zonation of the Cretaceous (Figure 5) i s s t i l l very much in a fledgling state. The first coarse zonation by Moore (1973) based on the Radiolaria recovered from Leg 17 divided the whole of the Cretaceous and late Late Jurassic into seven zones. Poor preservation, the lack of comparative material and inadequate calcareous control made the resulting zones difficult to apply. Riedel and Sanfilippo (1974) using a broader base introduced a coarse zonation, again of seven zones for the same period of time. Some of these zones were widely applicable but they again had very imprecise calcareous control. This zonation was modified by Foreman (1975) and i s here modified further to make it applicable to the sequences studied from the Atlantic basin. While these studies based largely on Deep Sea Drilling Project material were being made, Pessagno (in press) and Dumitrica (1975) produced, respectively, detailed zonations, based on well-preserved material with good calcareous control for the Late Cretaceous of California and the Cenomanian of Roumania. Dumitrica's zones were open-ended and his Late Cenomanian Holocryptocanium nanum-Excentropyloma cenomana Zone may be partly comparable to the lower part of Pessagno's Turonian Alievium superbum
Zone, and his Holocryptocanium barbui-H. tuberculatum Zone probably extends into the Albian. Figure 5 presents the various zonations in chronological order from left to right with the oldest zonation scheme recorded here at the left. In the figure the zones of Pessagno which could, for the most part, not be recognized in the Atlantic are correlated with the later zonations on the basis of age. The resulting uncertainty is indicated by dashed boundary lines. Also, the base of the Theocapsomma comys [group] is certainly lower than indicated by Riedel and Sanfilippo, the dashed line introduced here between it and the next lower zone again indicates uncertainty.
ZlE Figure 5. Radiolarian zonations
The best DSDP Atlantic sequence so far of MaestrichtiadLate Campanian Radiolaria was recovered from Hole 369A, Cores 35-39. While preservation is quite good, the Radiolaria are not common. This together with the incursion of a boreal fauna in Cores 35 and 37 leaves the accuracy of many first and last appearances open to question unless there is some outside back-up control. Nevertheless, this sequence, because it did have calcareous control, was used as a base together with other samples, with and without calcareous control, throughout the Late Cretaceous to contruct a chart of occurrences (Figure 6 ) in an attempt to pin down the ranges or a t least the order of appearance and extinction for some of the more common members of Late Cretaceous assemblages. This chart gives some detail for the CampaniadMaestrichtian section but i s deficient in that the early Late Cretaceous samples are too few in number. Five Zones are recognized: The
Cenomanian
Cenomanian
Holocryptocanium
Holocryptocanium
barbui-H.
tuberculatum
nanum-fxcentropyloma
Zone and the
cenomana
Zone,
both
Late of
Dumitrica; the Artostrobium, urna and Amphipyndax enesseffi Zones of Riedel and Sanfilippo emended here and the new Amphipyndax tylotus Zone. This new Amphipyndax tylotus Zone and the consequently emended Amphipyndax enesseffi Zone are defined and amplified as follows. All first appearances and last occurrences are morphological unless
otherwise stated. Arnphipyndax tylotus Zone Foreman, new zone The base is defined as the first evolutionary appearance of Amphipyndax tylotus. It includes near i t s
31 3
I
4
I
; : 1
N
p
; c m 1
m
1'1
:Imp I 8.111
,, Early Camp
1
I
C8mpmlan
,
La18 Campanm I Ewly U r s l r k h l i m
I
b
M...tr~'an
R
314
base the first appearance of Lihomelissa (?) hoplifes and Lophophaena polycyrfis. Also included are the last occurrences
of
Clafhropyrgus
firthium
and Afens
liriodes. The
top
is defined
as the
MaestrichtiadDanian boundary recognized by the last occurrence of the many species described from the California Maestrichtian (Foreman, 1968) which are recognized here, among them Lophophaena (?) polycyrfis, Theocampe bassilis and Theocapsomma feren. Arnphipyndax enesseffi Zone Riedel and Sanfilippo, emend. Foreman The base i s defined by the first morphological appearance of Amphipyndax enesseffi which may be approximately coincident with the first appearance of Lifhosfrobus puncfulafus. It includes near i t s base the first appearance of Clafhropyrgus riffhiurn and Theocampe apicafa. Also included are the first appearances of Theocampe daseia, Afens liriodes and Theocampe bassilis and the last occurrence of Arfosfrobium urna. The top is defined by the base of the next higher,A. fylotus Zone. The Amphipyndax rylorus Zone is recognized by the presence of i t s nominal species alone or in greater abundance than its ancestor A. enesseffi in all the Atlantic Basin and Atlantic margin samples of comparable age, except the two high latitude samples V-17-144 and V-18-129. It also cannot be recognized by i t s nominal species in California. It can, however, be identified in all the California localities of appropriate age, i.e. Maestrichtian, or Lower Maestrichtian-?Campanian of Foreman (1968, Table 1 ), and the two high latitude Atlantic samples, by the presence of LophophaenaOJpoIycyrfis and/or Lifhomelissa(?)hoplifes (Figure 7 ) .
Figure 7. Ranges for some diagnostic species of the Arnphipyndax fylotus and A . enesseffi Zones.
315
Taxonomic List Species included in the chart of occurrences (Figure 6) are listed here in alphabetical order with one or two references to the literature where they are more completely described or a more complete synonomy is given. Acidnomelos proapteron Foreman, in press. Illustrated here on Plate 1, Figure 12. Afensliriodes Riedel andsanfilippo, 1974,~.775,pl. 11,fig. 1 l ; p l . 13,figs. 14-16. Alievium gallowayi (White) in Pessagno, 1972, p. 299, pl. 25, figs. 4-6; pl. 26, fig. 5; pl. 31, figs. 2, 3. Alievium superbum (Squinabol) in Pessagno, 1972, p. 302, pl. 24, figs. 5-6; pl. 25, fig. 1; text-fig. 1. Amphipyndax enesseffi Foreman, 1966, p. 356, text-figs. 7-11. Illustrated here on Plate 1, Figure 2. Amphipyndax stocki (Campbell and Clark) in Foreman, 1968, p. 78, pl. 8, figs. 12a-c. Amphipyndax tylotus Foreman, in press. Illustrated here on Plate 1, Figure 1. Artostrobium urna Foreman, 1971, p. 1677, pl. 4, figs. 1-2. Cinclopyramis sanjoaquinensis (Campbell and Clark) 1944, p. 22, pl. 7, fig. 2. Foreman, in press. Clafhropyrgus bumasrus Riedel and Sanfilippo, 1974, p. 775, pl. 12, figs. 6-8. Clathropyrgus ritrhium Riedel and Sanfilippo, 1974, p. 775, pl. 3, fig. 12; pl. 12, figs. 9-12. Dicryomirraduodecimcosrafa (Squinabol) in Foreman, 1975, p. 614, pl. 1G. fig. 5; pl. 7, fig. 10.
Dictyomitra koslovae Foreman, 1975, p. 614, pl. 7, fig. 4. Dictyomitra lamellicosfata Foreman, 1968, p. 65, pl. 7, figs. 8a-b. Dicfyomitrapseudomacrocephala (Squinabol) in Foreman, 1975, p. 614, PI. 7, fig. 10. Druppafractona sp. A Foreman, in press. Illustrated here on Plate 1, Figure 3. Ellipsoxiphus sp. A Foreman, in press. Illustrated here on Plate 1, Figure 4. Ellipsoxiphus sp. B Foreman, in press. Illustrated here on Plate 1, Figure 5. Hagiastrin cf. Staurolonchidium tuberosum Rust in Riedel and Sanfilippo, 1974, p. 779, pl. 14, figs. 5-8. Holocryptocanium ruberculatum Dumitrica, 1970, pl. 16, figs. 102. 103ac. Lithomelissa ( ? ) heros Campbell and Clark in Foreman, 1968, p. 25, pl. 3, figs. 5a,b; text-fig. 1, fig. 7. Lithomelissa ( ? ) hoplires Foreman, 1968, p. 26, pl. 3, figs. 2a-c. Illustrated here on Plate 1, Figure 7. Lithomelissa ( ? ) pefilla Foreman, 1975, p. 616, pl. lG, figs. 2,3; pl. 6, fig. 3. Lithostrobus sp. A Foreman, in press. Illustrated here on Plate 1, Figure 9. Lithostrobus punctularus Pessagno, 1963, p. 210 (partim) pl. 1, fig. 1. Foreman, in press. Illustrated here on Plate 1, Figure 6. Lophophaena ( ? ) polycyrris Foreman, 1968, p. 23, pl. 3, figs. 3a-c. Illustrated here on Plate 1, Figure 8. Pseudoaulophacus lenriculatus (White) in Pessagno, 1963, p. 202, pl. 2, figs. 8-9. Pseudoaulophacuspargueraensis Pessagno, 1963, p. 204, pl. 2, figs. 4-7; pl. 6, figs. 4,5. Rhopalosyringium colpodes Foreman, 1968, p. 57, pl. 6, fig. 6. Rhopalosyringium sparnon Foreman, 1968, p. 56, pl. 6, fig. 5. Spongosaturnalis hueyi (Pessagno), in press. Foreman, 1975, p. 641, pl. 1A, fig. 6; pl. 4, fig. 10. Spongosarurnalis ( ? ) preclarus Foreman, 1975, p. 61 1, pl. 1A, figs. 4,5; pl. 4, fig. 8. Stichomitra asymmetra Foreman, in press. Illustrated here on Plate 1, Figure 10.
31 6 Stichomitra sp. A Foreman, in press. Illustrated here on Plate 1, Figure 11. Stichopilidium teslaense Campbell and Clark, in Foreman, 1968, p. 70, pl. 8 , fig. 13. Theocampe altamontensis (Campbell and Clark) in Foreman, 1968, p. 53, pl. 6, figs. 14,a.b. Theocampe apicata Foreman, 1971, p. 1679, pl. 4, fig. 6. Theocampe ascalia Foreman, 1971, p. 1678, pl. 4, fig. 4. Theocampe bassilis Foreman, 1968, p. 50, pl. 6, fig. 10. Theocampe daseia Foreman, 1968, p. 48, pl. 6, figs. 9a,b. Theocampe lispa Foreman, 1968, p. 49, pl. 6, fig. 11. Theocampe salillum Foreman, 1971, p. 1678, pl. 4, fig. 5. Theocapsomma comys Foreman, 1968, p. 29, pl. 4, figs. 2a-c; Theocapsomma comys group, in Foreman, in press. Theocapsomma teren Foreman, 1968, p. 32, pl. 4, fig. 4. Triactinosphaera sp. Although not tabulated because of i t s rare occurrence, this form i s illustrated here on Plate 1, Figure 13. It represents one of the few boreal forms present in the Atlantic Basin material as well as in the midcontinent Late Campanian.
Footnotes to Figure 6 These footnotes give the age or ages assigned to various samples together with a reference to the source. Letters "N" and "F"
indicate that the information is based respectively on nannofossil or
foraminifera1 data. 1. F: Maestrichtian, Abathomphalus mayaroensis 2.. Krasheninnikov (in press). hi.- Maestrichtian, ( ? ) Micula mura 2.. Cepek (in press). 2. F.- Maestrichtian, Globotruncana gansseri 2.. Krasheninnikov (in press). N.- Maestrichtian, Lithraphidites quadratus 2..Cepek (in press). 3. N.- Late Campanian/Early Maestrichtian, Arkhangelskiella cymbiformis 2 . . Cepek (in press). 4. N: Campanian, Milow (1970). Maestrichtiadcampanian, Tetralithus nitidus trifidus Z . , Bukry and Bramlette (1970). 5. F.- Campanian/Early Maestrichtian, Globofruncana fornicata - stuartiformis 2.. Beckman (1972). N.Late Campanian/Early Maestrichtian, Tetralifhus gothicus trifidus Z . , Roth and Thierstein (1972). Bukry (1972). Although samples 144-3-2,8486 and 144A4,CC are considered to be similar in age on the basis of nannofossil data, the presence in 144A4,CC of common, well-developed Amphipyndax enesseffi unaccompanied by A . fylotus suggests an earlier age and the latter sample is here considered to be Campanian. 6. N: Late Campanian/Early Maestrichtian, Tetralithus trifidus Z . , Cepek (in press). 7. F: Campanian, Globotruncana elevata 2 . . Krasheninnikov (in press). N.- Campanian, Eiffelithus eximus 2..Cepek (in press). 8 . F.- Early Campanian, Globotruncana elevata Z . , McNeely (1973). N.- Early CampaniadLate Santonian, Bukry (1973). 9. Campanian/E. Maestrichtian (age of Via Blanca form.) Bronnimann & Rigassi (1963). 10. F.- Campanian/Santonian, Pre Habana form., Bermudez (Foreman, 1968). 11. N: Senonian, Milo (1970). 12. F.- Senonian, Saito (1970). N.-Senonian, Milo (1970). 13. F.- Early Santonian, Globotruncana concavata 2 . . J.B. Saunders (Foreman, 1968). 14. F.- Cenomanian, M.A. Chierici (Foreman, 1968). 15. R: Cenomanian, Kozlova (1972).
31 7
Acknowledgements Partial financial support for this study was provided b y the National Science Foundation, Grant no.
DES75-19288.
References Aliev, KhSh., 1965. Radioliarii nizhnemelovykh otlozhenii severovostochnogo Azerbaidzhana i ikh Stratigraficheskoe znachenie. (Radiolarians of the Lower Cretaceous deposits of northeastern Azerbaidzhan and their stratigraphic significance.): Izd. Akad. Nauk Azerbaidz. SSR, Baku, p. 3-124. Beckman, J.P., 1972. The foraminifera and some associated microfossils of Sites 135 t o 144. In Hayes, D.E., Pimm, A.C., e t al., Initial Reports of the Deep Sea Drilling Project, v. 14, Washington (US. Government Printing Office). D. 389420. Bronnimann, P. and Rigassi, D., 1963. Contribution t o the geology and paleontology of the area of the city of La Habana, Cuba, and i t s surroundings: Eclog. Geol. Helv. v. 56, no. 1, p. 193-480, pl. 1-25. Bukry, D., 1972. Coccolith stratigraphy-Leg 14, Deep Sea Drilling Project. In Hayes, D.E., Pimm, A.C., e t al., Initial Reports of the Deep Sea Drilling Project, v. 14, Washington ( U S . Government Printing Office), D. 487494. Coccolith statigraphy, Leg 10, Deep Sea Drilling Project. In Worzel, J.L., Bryant, W., et at., -1973. Initial Reports of the Deep Sea Drilling Project, v. 10, Washington ( U S . Government Printing Office), p. 385406. Bukry, D. and Bramlette, M.N., 1970. Coccolith age determinations Leg 3, Deep Sea Drilling Project. In Maxwell, A. E e t al., Initial Reports of the Deep Sea Drilling Project, v. 3, Washington (U.S. Government Printing Office), p. 589-61 1. Cepek, Pave!, in press (See Lancelot, Y . and Seibold. E. e t al., in press.) Dumitrica, P., 1970. Cryptocephalic and cryptothoracic Nassellaria in some Mesozoic deposits of Romania: Rev. Roum. Geol., Geophys., Geogr., Ser. Geol., v. 14, p. 45-124. -1975. Cenomanian Radiolaria a t Podul Dimbovitei (excursion 6). In Micropaleontological guide to the Mesozoic and Tertiary of the Romanian Carpathians, 14th Europ. Micropal. Colloq., Bucharest (Inst. Geol. and Geophys.) Foreman, H. P., 1966. Two Cretaceous radiolarian genera: Micropaleontology, v. 12, p. 355-359. 1968. Upper Maestrichtian Radiolaria of California: Palaeontol. Assoc., London, Spec. Paper 3, p. iv + 1-82. -1971. Cretaceous Radiolaria. In Winterer, E. L., Riedel, W. R., e t al., Initial Reports of the Deep Sea Drilling Project, v. 7, Washington ( U S . Government Printing Office), p. 1673-1693. -1975. Radiolaria from the North Pacific. Deep Sea Drilling Project, leg 32. In Larson, R.L., Moberly, R., e t al., 1975. Initial Reports of the Deep Sea Drilling Project, v. 32, Washington (U.S. Government Printing Office), p. 579876. in press. Mesozoic Radiolaria in the Atlantic Ocean, o f f the west coast of Africa, Deep Sea Drilling Project, Leg 41. Johnson, T.C.. 1976. Controls on the preservation of biogenic opal in sediments of the eastern tropical Pacific: Science, v. 192, p. 887890. Kozlova, G. E., 1972 (See Petrushevskaya, M. G. and Kozlova, G. E., 1972.) Krasheninnikov, V., in press. (See Lancelot, Y . , and Seibold, E. et al., in press). Lancelot, Y . , and Seibold, E. e t al., in press. Initial Reports of the Deep Sea Drilling Project, V. 41, Washington (US. Government Printing Office). Maxwell, A. E. et al., 1970. Initial Reports of the Deep Sea Drilling Project, v. 3, Washington (US. Government Printing Office) x x + 806 p.
-
-
318
McNeely, B. W., 1973. Biostratigraphy of the Mesozoic and Paleogene pelagic sediments of the Campeche embankment area. In Worzel, J. L., Bryant, W., e t al., Initial Reports of the Deep Sea Drilling Project, v. 10, Washington (US. Government Printing Office) p. 679895. Milow, D., 1970. (See Maxwell, A. E. e t al. 1970). Moore, T. C., 1973. Radiolaria from Leg 17 of the Deep Sea Drilling Project. In Winterer, E. L., Ewing. J. L., e t al., Initial Reports of the Deep Sea Drilling Project, v. 17, Washington (US. Government Printing Office), p. 797869. Murray, John, 1876. Preliminary reports to Professor Wyville Thomson, F.R.S., Director of the civilian staff, on work done on board the 'Challenger': Roy. SOC. London, Proc., v. 24, p. 471-544. Pessagno, E. A,, Jr. 1963. Upper Cretaceous Radiolaria from Puerto Rico: Micropaleontology, v. 9, no. 2, p. 197-214. 1972. Pseudoaulophacidae Riedel from the Cretaceous of California and the Blake-Bahama Basin (JOIDES Leg 1) . In Cretaceous Radiolaria: Bull. Am. Paleontol., v. 61, p. 281-328. in press. Radiolarian zonation and stratigraphy of the Upper Cretaceous portion of the Great Valley sequence, California Coast Ranges. Petrushevskaya, M. G. and Kozlova, G. E., 1972. Radiolaria: Leg 14, Deep Sea Drilling Project. In Hayes, D. E., Pimm, A.C. e t al., Initial Reports of the Deep Sea Drilling Project, v. 14, Washington (US. Government Printing Office), p. 495878. Riedel, W. R. and Sanfilippo, A., 1974. Radiolaria from the southern Indian Ocean, DSDP Leg 26. In Davies, T. A., Luyendyk, B . P., e t al., Initial Reports of the Deep Sea Drilling Project, v. 26, Washington (US. Government Printing Office), p. 771814. Roth, P. H. and Thierstein, H., 1972. Calcareous nannoplankton: Leg 14 of the Deep Sea Drilling Project. In Hayes, D. E., Pimm, A. C., e t al., Initial Reports of the Deep Sea Drilling Project, v. 14, Washington (US. Government Printing Office), p. 421485. Saito, T., 1970. (See Maxwell, A. E., e t al., 1970.)
Plate 1. All figues are magnified 214x. Fig. 1. Amphipyndax tylotus: 13A-2-1.84-88. SI. 2,01910. Fig. 2 . Amphipyndax enesseffi: 369A-39-3,69-70. SI. 1, €2413. Fig. 3. Druppatractona sp. A : 369A-37,CC. SI. 2, L3013. Fig. 4. Ellipsoxiphus sp. A: 369A-35,CC. SI. 4, D1912. Fig. 5. Nlipsoxiphus sp. B: 369A-37,CC. SI. 2, T2810. Fig. 6. Lithostrobus punctulatus: 369A-38,CC. SI. 1, 01810. Fig. 7. Lithomelissa (?) hoplites: 369A-37,CC. SI. 2 , V21/2. Fig. 8. Lophophaena polycyrtis V-18-129,78 cm. St. 1241, D39/0. Fig. 9. Lithostrobus sp. A : 369A-36-5.6587. SI. 1, J2811. Fig. 10. Stichomitra asymmetra: 13A-2-1,8486. SI. 1, L2414. Fig. 11. Stichomitra sp. A : 369A-36,CC. SI. 5, N3711. Fig. 12. Acidnomelosproapteron: 369A-38,CC. SI. 1, R2411. Fig. 13. Triactinosphaera sp.: 369A-37,CC. SI. 1, M2114.
319 Plate 1
320
Discussion W . Hay: Can t h e A . mid-continent?
D r . W.
t y l o t u s Zone b e r e c o g n i z e d i n t h e
Foreman: No, t h e N o r t h and S o u t h Dakota R a d i o l a r i a I have examined of comparable a g e a r e a l m o s t c o m p l e t e l y d i f f e r e n t . The few s p e c i e s t h a t a r e common t o b o t h s e q u e n c e s a r e o n l y s p o r a d i c a l l y p r e s e n t i n t h e A t l a n t i c B a s i n s e q u e n c e and have n o t been t a b u l a t e d . One form i s i l l u s t r a t e d on P1. 1, f i g u r e 1 3 .
321 JURASSIC PALYNOSTRATIGRAPHY OF OFFSHORE EASTERN CANADA
JONATHAN P. BUJAK and GRAHAM L . WILLIAMS A t l a n t i c G e o s c i e n c e C e n t r e , D a r t m o u t h , Nova S c o t i a ABSTRACT The S c o t i a n S h e l f and Grand Banks, o f f s h o r e s o u t h e a s t e r n Canada, c o v e r a p p r o x i m a t e l y 175,000 s q u a r e m i l e s and e x t e n d o v e r 850 m i l e s f r o m n o r t h e a s t t o southwest.
More t h a n 100 e x p l o r a t o r y w e l l s d r i l l e d t o d a t e
have e n c o u n t e r e d Lower P a l e o z o i c t o R e c e n t s e d i m e n t s , w h i c h f r e q u e n t l y c o n t a i n r i c h and age d i a g n o s t i c p a l y n o m o r p h a s s e m b l a g e s .
Based upon
t h e a n a l y s i s o f t w e n t y w e l l s , t e n p a l y n o m o r p h zones have been d e f i n e d within the Jurassic.
These zones have been c o r r e l a t e d w i t h t h e s t a n d a r d
European J u r a s s i c s t a g e s , a l l o f w h i c h a r e r e c o g n i s e d , a l t h o u g h t h e H e t t a n g i a n - S i n e m u r i a n and T o a r c i a n - A a l e n i a n c a n n o t , as y e t , be s e p a r a t e d . B o t h m a r i n e ( d i n o f l a g e l l a t e s ) and n o n - m a r i n e ( s p o r e s ) p a l y n o m o r p h s a r e u t i l i z e d f o r z o n a t i o n and f o r t h e i n t e r p r e t a t i o n o f e n v i r o n m e n t s .
On
t h e S c o t i a n S h e l f t h e o l d e s t e n c o u n t e r e d s e d i m e n t s , t h e E u r y d i c e Forma t i o n , a r e R h a e t i a n - H e t t a n g i a n , and a r e o v e r l a i n b y t h e A r g o S a l t , p r o b a b l y o f H e t t a n g i a n - S i n e m u r i a n age.
The s u c c e e d i n g n o n - m a r i n e o r
m a r g i n a l l y m a r i n e I r o q u o i s F o r m a t i o n c o n t a i n s d i s t i n c t i v e s p o r e assemblages w h i c h have no known c o r r e l a t i v e s , b u t a r e t e n t a t i v e l y d a t e d S i n e murian-Pliensbachian.
On t h e Grand Banks, m a r g i n a l l y m a r i n e a n d e v a -
p o r i t i c H e t t a n g i a n - S i n e m u r i a n s t r a t a a r e o v e r l a i n b y t h e m a r i n e Whale U n i t , whose d i n o f l a g e l l a t e s c l o s e l y compare w i t h P l i e n s b a c h i a n - T o a r c i a n assemblages f r o m E u r o p e .
T h r o u g h o u t t h e M i d d l e and L a t e J u r a s s i c , p r e -
d o m i n a n t l y s h a l l o w m a r i n e d e p o s i t i o n o c c u r r e d i n b o t h a r e a s , w i t h some non-marine episodes.
D i n o f l a g e l l a t e species d i v e r s i t y , low i n the Early
J u r a s s i c and A a l e n i a n - B a j o c i a n ,
i n c r e a s e d t o a peak i n t h e K i m m e r i d g i a n
and s u b s e q u e n t l y d e c l i n e d i n t h e P o r t l a n d i a n . SOMMAI R E
La p l a t e f o r m e c o n t i n e n t a l e de l a N o u v e l l e i c o s s e e t l e s Grands Bancs, s i t u 6 s a u l a r g e de l a p a r t i e s u d - e s t d u Canada, r e c o u v r e n t e n v i r o n
322
175,000 milles carr6s et s'gtendent, du nord-est au sud-ouest, sur plus de 850 milles. Jusqu'ici, plus de 100 sondages d'exploration ont permis d'6chantilloner des sediments dont l'origine s'6chelonne depuis le Palgozoyque infgrieur jusqu'au Quaternaire, et qui renferment frgquemment des riches assemblages de palynomorphes permettant d'en 6tablir l'bge. L'analyse de vingt sondages a permis de distinguer dix zones palynomorphiques au sein du Jurassique. On a 6tabli la correlation entre ces zones et les 6tages jurassiques classiques de l'Europe, qui ont tous 6t6 identifies, bien qu'on ne puisse pas encore s6parer 1'Hettangien-Singmurien et le Toarcien-Aal6nien. On se sert h la fois des palynomorphes marins (dinoflagellgs) et non marins (spores) pour gtablir les zones et pour interprgter les environnements. Les sgdiments les plus anciens de la plateforme continentale de la Nouvelle icosse, ceux de la formation Eurydice, datent du Rhetien-Hettangien, e t sont recouverts par la formation Argo ("Argo Salt"), qui remonte probablement h 1 'Hettangien-Singmurien. La formation Iroquois, non marine ou marginalement marine, qui y succsde renferme des assemblages distinctifs de spores qui n'ont pas de correlatif connu, mais qu'on date expgrimentalement du Singmurien-Pliensbachien. Sur les Grands Bancs, des strates hettangiennes-sinemuriennes, gvaporitiques et marginalement marines sont recouvertes par le d6p6t marin Whale ("Whale Unit"), dont les dinoflagell6s correspondent de trPs prPs aux assemblages pliensbachienstoarciens de 1 'Europe. Durant tout le Jurassique moyen et supgrieur, i l s'est fait dans les deux secteurs des dgp6t.s marins surtout en eau peu profonde, avec quelques 6pisodes non marins. Le nombre d'espsces diff6rentes de palynomorphes, faible au Jurassique infgrieur et 'a l'Aal6nien-Bajocien, s'est accru pour atteindre un maximum pendant le Kimmeridgien, et a diminu6 par la suite au Portlandien. INTRODUCTION The eastern Canadian continental shelf extends from the northern part o f Georges Bank in the southwest to the Arctic Ocean in the north. Its southern part primarily consists o f the Scotian Shelf and Grand Banks. These two areas cover more than 175,000 square miles and extend over 850 miles from northeast to southwest (text-figure 1). Since the first two exploratory wells were drilled on the Grand Banks in 1966, over 100 wells have provided a unique opportunity for detailed geological study o f the western North Atlantic margin. Intensive investigations o f many of
323
L
I Zoom
Text-figure 1 .
Scotian Shelf-Grand Banks wells in which J u r a s s i c sections have been studied.
these wells have provided l i t h o s t r a t i g r a p h i c , b i o s t r a t i g r a p h i c and geochemical data which have been i n t e g r a t e d with various geophysical parameters. B i o s t r a t i g r a p h i c s t u d i e s have been undertaken on foraminifera, o s t r a c o d s , nannoplankton, and various organic-walled m i c r o f o s s i l s , the l a t t e r including chitinozoa and a c r i t a r c h s in the Early Paleozoic, a c r i t a r c h s and spores in the Late Paleozoic, a n d d i n o f l a g e l l a t e s , spores, The presence of dinoand a c r i t a r c h s in t h e Mesozoic a n d Cenozoic. f l a g e l l a t e s , chitinozoa, and a c r i t a r c h s i s usually i n d i c a t i v e of marine deposition. Spores, although of t e r r e s t r i a l o r i g i n , may a l s o be present in marine sediments and thus permit c o r r e l a t i o n of marine a n d non-marine deposits. This paper presents t h e palynological zonation, based on dinof l a g e l l a t e s and spores, o f the J u r a s s i c studied in eleven Scotian Shelf The Scotian Shelf wells a r e and nine Grand Banks wells ( t e x t - f i g u r e 1 ) . Mobil-Tetco Cohasset 0-42, Mobil-Tetco Oauntless D-35, Shell Argo F-38, Shell Eurydice P-36, Shell Mic Mac 5-77, Shell Missisauga H-54, Shell Mohawk B-93, Shell Mohican 1-100, Shell Naskapi N-30, Shell Oneida 0-25,
I l l
l
!
'
L
J U RASSlC
~
l
I
I
I
~~
~~
vermiculata
~
~~
~
~~
_
~- .
~
~
~~~
-
_
~~
~~~
~
ionvaulacvsta cladoohora
Aeiourogonyaulax deflandrei ___ llipsoidictyum tinctum
~
~~
~~
~
_
~
-
___
~~
-- -
lolvsteohaneDhorus Daracalathus ;onyaulacysta lurassica
I nealei
iystrichogonyaulax cornigera _
:leistosphaeridium tribuliferum. ;en et so 2 Gocht 1970
~
idnatosphaeridium caulleryi ~i aemulum ~
_
-~
-
_ _ ~ -
~
:ordosphaeridium costatum.
.ndoscrinium eisenacki iystematophora areolata
~
:tenidodinium ornatum .enua sp B
1.
1 ovulum
lalensiella ampulla
.ithodinia jurassisa
.eptodinium subtile . subtile ssp pectinigerum
~
____
.~thodiniavalensii ~. :tenidodinium continuum Gnyaulacysta aldorfensis
~~
~~~
a f f tenellum .eptodinium regale
:tentdodimurn 'enua rioulti
:tenidodinium pachydermum
dendicodinium reticulaturn
~~~~
.uehndea spinosa daturodinium inornatum ~~dannoceratoose oracilis
PZE
~
~
-
_
Herendeenia pisciformis Prolixosphaeridium granulosum Muderongia simplex Gplosphaera reticulospinosa Gen et SD 2 Gonyaulacysta aculeata Gonyaulacysta ehrenbergi lmbatodinium antennatum Wanaea spectabills Dingodinium jurassicum Prolixosphaeridium deirense Psaligonyaulax apatela lmbatodinium kondratjevi Polystephanephorus sarleantii AmDhorula metaelllDtlCa Ctenidodinium culmulum Ctenidodiniurn panneum Lanterna sportula Hystrichodinium pulchrum Oligosphaeridium dividuum Systematophora schindewolfi Ctenidodinium schizoblatum Pyxidiella sp
Scriniodinium dictyotum Leptodinium arcuatum Gonyaulacysta ambigua
~-
Endoscrinium luridum Gen et sp 1 Gonyaulacysta granulata G _ aranuliaera ,epioc n -m egeren ... Parvocavar-s trDeros.s -
.
-
Gonyaulacysta lurassica SSP longicornls Systematophora fasciculigera
Zcisucysta sp A Tenua hystrix Stephanelytron caytonense S scarburghense Hexagonifera )urass\ca Stephanelytron redcliffense Acanthaulax paliuros Taeniophora iunctispina Tenua villersense
I
21
I
EARLY
I MIDDLE
J URASSlC
I
-i
LATE
I
.
-.
~~~
~-
~~
_ _
'ilosisporites trichopapillosus k t v .o p h v.l l i d i t e s eauiexinus
'ilosisporites s p A Iensoisporites velatus
~
_-
-
9ZE
2icatiicosisporites australiensis 4&$~triradites spinulosus -~~
-eptolepidites malor h o n a t i s p o r a valdensis .eptolepidites psarosus
Indosporites jurassicus
rrilobosporites lurassicus jtaplinisporites caminus
:ircularaesporites cerebroides 3allialasporites dampieri Zamerosporites secatus h t i o n i s o o r i t e s SD
ixesipollenites tumulus
~~
~.
Ichinitosporites ~cf iliacoides 2erebropollenites mesozoicus Zadaroasoorites verrucosus :amarozonosporites rudis
'Cycaaop _.ies sp B
Vitreisporites pallidus 7 Cycadopites s p A 3vcadoDites subaranulosus
:,camp ies aeter .s 2 r c a a o p ies ct anscn
2orollina meyeriana Zonvolutisoora klukiforma
Verrucosisporites cf cheneyi Porcellispora longdonensis
327
AGE
ZONE Ctenidodinium panneum PORTLANDIAN W~ K~MMER~DG~A ' GN Gonyaulacysta cladophora 0 '1 Gonyaulacysta jurassica OXFORDIAN -. Valensiella vermiculata CALLOVI A N cn. W 1 Gonyaulacysta filapicata BATHONIAN cn n a BAJoCIAN f Mancodinium semitabulatum ~
LT
3
AALENIAN
---
TOARCl A N PLIENSBACHIAN
Nannocera topsis gracilis
3
7
SINEMURIAN HE T TA N G I A N I-
RHAETIAN
T e x t - f i g u r e 4.
---
Echinitosporites cf. iliacoides Cycadopites subgranulosus Corollina meyeriana ( P E A K ) GSC
J u r a s s i c p a l y n o m o r p h z o n a t i o n , S c o t i a n S h e l f - G r a n d Banks.
and S h e l l Wyandot E-53.
The Grand Banks w e l l s a r e Amoco-Imp B i t t e r n
M-62, Amoco-Imp C o r m o r a n t N-83, Amoco-Imp Heron H-73, Amoco-Imp P e t r e l A-62,
A m o c o - I m p - S k e l l y E g r e t K-36, A m o c o - I m p - S k e l l y O s p r e y H-84, Amoco
I O E E i d e r M-75, Amoco-IOE M u r r e G-67, and M o b i l - G u l f F l y i n g Foam 1-13. The s t r a t i g r a p h i c r a n g e s o f J u r a s s i c d i n o f l a g e l l a t e s and s p o r e s , as d e t e r m i n e d f r o m t h e s e w e l l s , a r e shown i n t e x t - f i g u r e s 2A, 2B and 3 . Ten f o r m a l zones a r e r e c o g n i z e d i n t h e J u r a s s i c o f t h e S c o t i a n S h e l f - G r a n d Banks, b a s e d o n t h e 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 o f d i n o f l a g e l l a t e s and s p o r e s ( t e x t - f i g u r e 4).
N i n e o f t h e s e a r e assemblage
zones a n d one i s a peak zone as d e f i n e d i n t h e A m e r i c a n Commission o n S t r a t i g r a p h i c N o m e n c l a t u r e , 1961, A r t i c l e s 209, 21, J u r a s s i c zones w e r e e r e c t e d b y W i l l i a m s ( 1 9 7 5 ) .
The f o u r h i g h e s t
The r e m a i n i n g s i x zones
are proposed f o r m a l l y h e r e i n f o r t h e f i r s t time, although
H. S a b r y
v e r b a l l y presented a paper, d i s c u s s i n g t h e E a r l y J u r a s s i c Scotian S h e l f p a l y n o s t r a t i g r a p h y , a t t h e E i g h t h Annual M e e t i n g o f t h e A m e r i c a n Associ a t i o n o f S t r a t i g r a p h i c P a l y n o l o g i s t s a t H o u s t o n i n 1975.
The p r e s e n t
328 s t u d y i s based p r i m a r i l y o n c u t t i n g s samples, so t h a t i t i s n e c e s s a r y t o use f o s s i l " t o p s " ( i . e . ,
t h e l a t e s t o r highest occurrences o f a species)
t o d e f i n e zones a n d c o r r e l a t e between w e l l s .
Each assemblage zone t a k e s
i t s name f r o m a f o s s i l n o t f o u n d i n s e d i m e n t s above t h a t zone.
Where
i t can be e s t a b l i s h e d w i t h some c e r t a i n t y , as when u s i n g s i d e w a l l o r
conventional cores, t h e "base", o r o l d e s t occurrence, o f a species i s a l s o u s e d t o d e l i n e a t e zones a n d f o r c o r r e l a t i o n .
Peak zones a r e
named a f t e r s p e c i e s w h i c h a t t a i n t h e i r maximum abundance w i t h i n them. The m a j o r i t y o f S c o t i a n S h e l f - G r a n d Banks J u r a s s i c s p e c i e s have been p r e v i o u s l y d e s c r i b e d f r o m European s u r f a c e s e c t i o n s .
The s t r a t i -
g r a p h i c r a n g e s o f t h e s e s p e c i e s p e r m i t d a t i n g o f t h e zones r e l a t i v e t o European s t a g e t e r m i n o l o g y .
I t m u s t be n o t e d , however, t h a t a l l age
a s s i g n m e n t s a r e p r o v i s i o n a l and may b e s u b j e c t t o m o d i f i c a t i o n as new d a t a become a v a i l a b l e . C o r r e l a t i o n o f t h e p a l y n o l o g i c a l and f o r a m i n i f e r a 1 z o n a t i o n s i s presented i n Gradstein ( t h i s volume). BIOSTRATIGMPHY
CoroZZina meyeriana Peak Zone Type s e c t i o n : S h e l l E u r y d i c e P-36, 8920-9700 f t . Other wells:
A r g o F-38, Heron H-73, O s p r e y H-84.
C h a r a c t e r i s t i c s p e c i e s : S p o r e s ; Convolutispora k l u k i f o r m a ( N i l s s o n ) S c h u l z , Corollina meyeriana ( K l a u s ) V e n k a t a c h a l a a n d G6czdn abundance ( p l a t e 1, f i g u r e l ) , C. torosus ( R e i s s i n g e r ) K l a u s , Verrucosisporites cheneai C o r n e t and T r a v e r s e , I/errucosisporites sp. A ( p l a t e 2, f i g u r e 3 ) . S p e c i e s b a s e s : None e s t a b l i s h e d . S e l e c t e d s p e c i e s t o p s : Spores; C. rneyeriana abundance. D i s c u s s i o n : Some h o r i z o n s f r o m t h i s zone may be d o m i n a t e d by C. torosus w i t h f e w e r C. rneyeriana.
The assemblages c o n t a i n i n g a b u n d a n t C, meyer-
iana a r e c l o s e l y c o m p a r a b l e t o t h o s e documented b y C o r n e t and T r a v e r s e (1975) f r o m t h e S h u t t l e Meadow F o r m a t i o n o f t h e H a r t f o r d B a s i n , e a s t e r n United States.
These a u t h o r s g i v e a d e t a i l e d d i s c u s s i o n o f t h e age o f
t h e S h u t t l e Meadow F o r m a t i o n a n d a s s i g n a p r o b a b l e R h a e t o - L i a s s i c (Rhaet i a n - H e t t a n g i a n ) age f r o m m a c r o - and m i c r o f o s s i l e v i d e n c e a n d i s o t o p e dating.
The C. meyeriana Peak Zone i s r e c o g n i s e d i n o n l y f o u r w e l l s
( A r g o F-38, E u r y d i c e P-36,
Heron H-73, O s p r e y H - 8 4 ) .
I n B i t t e r n M-62,
C o r m o r a n t N-83, a n d P e t r e l A-62 t h e p r e - l a t e P l i e n s b a c h i a n L i a s s i c c a n n o t be s u b d i v i d e d .
329
Cyeadopites subgranulosus Assemblage Zone Type s e c t i o n : Shell Eurydice P-36, 2640-8814 f t .
Other wells: Argo F-38, Mohican 1-100. Selected species bases: Spores; CerebropolZenites mesozoicus (Couper) Ni 1 sson, Cycadopites subgranulosus (Couper) comb. n o v . , ?Cycadopites spp. A and B ( p l a t e 2 , f i g u r e s 4 , 7 , 8 ) , E c h i n i t o s p o r i t e s c f . iZiacoides
Schulz a n d Krutzsch, i(raeuse2isporite.s r e i s s i n g e r i ( H a r r i s ) Ilorbey ( p l a t e 1 , f i g u r e 2 ; p l a t e 2, f i g u r e s 5 , 6 ) . Selected species tops : Spores ; Cycadopites subgranuZosus, ?Cycadopites spp. A a n d B , Verrucosisporites cheneyui, V e r r u e o s i s p o r i t e s sp. A , Discussion: Laevigate, g r a n u l a t e , a n d verrucate species of Cycado?ites a n d ?Cycadopites a r e frequently common a n d c h a r a c t e r i s e t h i s zone. Assemblages a r e mostly dominated by CoroZZina torosus while C. meyerianu i s uncommon. The predominance of C. torosus over C. me3eria.w t h r o u g h -
o u t t h e Picadopites subgranulosus Zone indicates t h a t t h e zone i s no o l d e r t h a n Hettangian, following the reasoning of Cornet a n d Traverse (1975), The presence of Cerebrosollenites mesozoicus supports a n age no o l d e r t h a n L i a s s i c a n d t h e occurrence of KraeuselLsporites reissingeri a n d Cycadopites subgranuZosus i n the C. subgranulosus Zone indicates t h a t the zone i s Hettangian-Sinemurian in age. Dinoflagellates have n o t been recorded from t h i s zone. E c h i n i t o s p o r i t e s c f . ilicrccia'es Assemblage Zone
Type s e c t i o n : Shell Eurydice P-36, 1765-2575 f t . Other wells: Argo F-38. Selected species bases: None e s t a b l i s h e d . Selected species tops: Spores; Convolutispora klukiforma, Corollina medericrna, E e h i n i t o s p o r i t e s c f . iiiaco<des ( p l a t e 1 , f i g u r e s 3 , 4 ; p l a t e 2 , f i g u r e s 9 , 1 2 ) , E c h i n i t o s p o r i t e s sp. A ( p l a t e 2 , f i g u r e s l O , l l ) , KraeuseZisporites r e i s s i n g e r i . Discussion: Monosulcate pollen assigned t o E c h i n i t c s p o r i t e s c f . i l i a c o i des a r e common in t h i s zone. E. iZiaco<des has only been described from Late T r i a s s i c sediments ( s e e Scheuring, 1970, f o r d i s c u s s i o n ) . 2oonvcZutCspora kZukiforma i s c h a r a c t e r i s t i c of the European L i a s s i c
(Cornet and Traverse, 1975), and Geiger a n d Hopping (1968) noted t h a t Xraeuselispc:+ites r e i s s i n g e r i has a known range of Rhaetian-Pliensbachian.
The zone i s t e n t a t i v e l y dated Sinemurian-early Pliensbachian.
330
Nannoceratopsis g r a c i l i s Assemblage Zone Type s e c t i o n : Amoco-IOE M u r r e G-67, 6450-7870 ft. Other w e l l s :
A r g o F-38, B i t t e r n M-62, C o r m o r a n t N-83, E i d e r M-75,
Heron H-73, M o h i c a n 1-100. S e l e c t e d s p e c i e s b a s e s : D i n o f l a g e l l a t e s ; Luehndia spinosa M o r g e n r o t h ,
Mancodiniwn semitabulatwn M o r g e n r o t h , Maturodinim inornatw? M o r g e n r o t h , Mendicodinim reticuZatw? M o r g e n r o t h , Nannoceratopsis g r a c i l i s A1 b e r t i , Pareodinia eeratophora D e f l a n d r e .
S p o r e s ; C a l l i a l a s p o r i t e s dampieri
(Balme) Dev, Contignisporites sp. ( p l a t e 2, f i g u r e 2 ) , K l u k i s p o r i t e s
pseudoreticulatus Couper
.
S e l e c t e d s p e c i e s t o p s : D i n o f l a g e l l a t e s ; Luehndia spinosa, Maturodinim
inomatwn, Nannoceratopsis g r a c i l i s . D i s c u s s i o n : D i n o f l a g e l l a t e s f i r s t a p p e a r i n t h e Nannoceratopsis g r a c i l i s Zone on t h e S c o t i a n S h e l f and i n d i c a t e a l a t e P l i e n s b a c h i a n - T o a r c i a n / A a l e n i a n age, as t h e y compare c l o s e l y w i t h assemblages o f t h i s age f r o m n o r t h w e s t Europe (see W i l l i a m s , i n p r e s s , f o r d i s c u s s i o n ) .
On t h e
Scotian Shelf, t h e coeval deposits are devoid o f d i n o f l a g e l l a t e s , b u t can be dated u s i n g spores.
The f i r s t a p p e a r a n c e o f C a l l i a l a s p o r i t e s
dampieri, C o n t i g n i s p o r i t e s sp. a n d K l u k i s p o r i t e s p s e u d o r e t i c u l a t u s i s t a k e n as t h e base o f t h e N , g r a c i l i s Zone.
H e r n g r e e n and de B o e r
( 1 9 7 4 ) n o t e d t h a t C o n t i g n i s p o r i t e s problematicus f i r s t a p p e a r s i n t h e Upper P l i e n s b a c h i a n .
Pocock ( 1 9 7 0 ) p l a c e d t h e b a s e o f C. dumpieri i n
w e s t e r n Canada w i t h i n t h e P l i e n s b a c h i a n - T o a r c i a n .
Vancodiniwn semitabulatwn Assemblage Zone Type s e c t i o n : Amoco-IOE E i d e r M-75, 8545-10,240 Other w e l l s :
ft.
A r g o F-38, B i t t e r n M-62, C o r m o r a n t N-83, M o h i c a n 1-100,
M u r r e G-67. S e l e c t e d s p e c i e s b a s e s : D i n o f l age1 l a t e s ; Ctenidodiniwn pachydermwn ( D e f l a n d r e ) Gocht, C. a f f . t e n e l l w n D e f l a n d r e , Gonyaulacysta f i l a p i e a t a Gocht, Leptodiniwn regale Gocht. S e l e c t e d s p e c i e s t o p s : D i n o f l a g e l l a t e s ; Mancodiniwn semitabulatwn,
Mendicodiniwn r e t i c u l a t w n . D i s c u s s i o n : Ctenidodiniwn pachydermwn and Gonyaulacysta f i l a p i c a t a f i r s t appear i n t h e Bajocian o f southern England (personal o b s e r v a t i o n , G.L.
W i l l iams).
Mancodi?iwn semitabulatwn a n d Mendicodiniwn r e t i c u l a t w n
were o r i g i n a l l y d e s c r i b e d f r o m t h e L a t e P1 i e n s b a c h i a n o f Germany b y Morgenroth (1970).
The a s s o c i a t i o n o f a l l f o u r s p e c i e s i n t h i s zone
i n d i c a t e s a B a j o c i a n age.
The M . semitabulatwn Zone i s r e c o g n i s e d i n
331
T e x t - f i g u r e 5.
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 o f J u r a s s i c sediments i n t h e examined Grand Banks we1 1s
T e x t - f i g u r e 6.
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 o f J u r a s s i c sediments i n t h e examined S c o t i a n S h e l f w e l l s .
.
332
s i x o f t h e w e l l s , f o u r on t h e Grand Banks a n d t w o o n t h e S c o t i a n S h e l f ( t e x t - f i g u r e s 5,6).
GorqauZaqsta f i l a p i c a t a Assemblage Zone Type s e c t i o n : Amoco-Imp B i t t e r n M-62, 6600-8230 f t . Other w e l l s :
Argo F-38, C o r m o r a n t N-83, E i d e r M-75, Heron H-73, Mohawk
8-93, M o h i c a n 1-100, M u r r e G-67, Wyandot E-53. S e l e c t e d s p e c i e s b a s e s : D i n o f l a g e l l a t e s ; Cordosphaeridi-un costaturn (Davey and W i 11 i a m s ) Gorka, C t e n i d o d i n i m omaturn ( E i s e n a c k ) D e f l a n d r e ,
Gonyaulacksta a l d o r f e n s i s G o c h t , Lithodinia jurassica E i s e n a c k ,
Y a l e n s i e l l a ovulvr! ( O e f l a n d r e ) E i s e n a c k , V . vermiculata G o c h t . S e l e c t e d s p e c i e s t o p s : D i n o f l a g e l l a t e s ; GongauZacysta f i l a p i c a t a . D i s c u s s i o n : B a t h o n i a n d i n o f l a g e l l a t e assemblages have been d e s c r i b e d f r o m B u l g a r i a b y Dodekova ( 1 9 7 5 ) a n d f r o m Germany b y G o c h t ( 1 9 7 0 ) .
Most
o f t h e s p e c i e s d e s c r i b e d by Gocht, i n c l u d i n g Gonyaulacysta f i l a p i c a t a , o c c u r i n t h e 9. f i l a F i c a t a Zone.
The zone i s t h e r e f o r e d a t e d B a t h o n i a n .
A d e t a i l e d d i s c u s s i o n o f t h e B a t h o n i a n i n d e x s p e c i e s documented b y G o c h t i s given i n Williams ( i n press).
T h e r e i s a marked i n c r e a s e i n d i n o -
f l a g e l l a t e s p e c i e s d i v e r s i t y f r o m n i n e i n t h e Mancodiniwn semitabuZatm Zone ( B a j o c i a n ) t o 22 i n t h e G. f i l a p i c a t a Zone ( B a t h o n i a n ) , as shown i n t e x t - f i g u r e 7.
VaZensiella vermiculata Assemblage Zone Type s e c t i o n : S h e l l N a s k a p i 1.1-30, 6590-6992 f t . Other w e l l s :
A r g o F-38, B i t t e r n M-62, C o h a s s e t D-42, C o r m o r a n t N-83,
E i d e r M-75, H e r o n H-73, M i c Mac 5-77, Mohawk B-93, Mohican 1-100, M u r r e G-67, Oneida 3-25, Wyandot E-53. S e l e c t e d s p e c i e s b a s e s : D i n o f l a g e l l a t e s ; Gen. e t sp. 2 G o c h t , 1970,
GonyauZacysta cladophora ( D e f l a n d r e ) Dodekova, G. j u r a s s i c a ( D e f l a n d r e ) N o r r i s and S a r j e a n t . Scriniodiniwn c r y s t a l l i n m ( D e f l a n d r e ) Klement,
Sgstematcphora o r b i f e r a K l e m e n t .
The d i n o f l a g e l l a t e s Hezagonifera
jurassica G i tmez and S a r j e a n t , Stephanelgtron caytonense S a r j e a n t a n d Spores; S . scarburghense S a r j e a n t a p p e a r i n t h e u p p e r p a r t o f t h i s zone. T r i l o b o s p o y i t e s j u r a s s i c u s Pocock. S e l e c t e d s p e c i e s t o p s : D i n o f l a g e l l a t e s ; GonyauZacysta aldorfensis,
L e p t o d i n i m regale, L. s u b t i l e , Lithodinia jurassica, StephaneZytron caytonense, S. scarburghense, Y a l e n s i e l l a ovulwn, V . vermiculata. Spores ; Circularaesporites cerebroides. D i s c u s s i o n : W i l l i a m s ( i n p r e s s ) d i s c u s s e d t h e known r e c o r d s o f d i n o f l a g e l l a t e s f r o m t h e t y p e C a l l o v i a n s e c t i o n and e l s e w h e r e .
Comparison
333
-
SPECIES
D N O . OF BASES Text-figure 7 .
W N O . OF "TOPS"
TOTAL N0.
Species d i v e r s i t y of J u r a s s i c dinocysts, Scotian ShelfGrand Banks.
of the V . vermiculata Zone assemblages with t h i s data s u b s t a n t i a t e s i t s This zone i s widespread in the J u r a s s i c of Callovian age assignment. offshore eastern Canada and i s apparently always marine with diverse The zone d i n o f l a g e l l a t e assemblages, 41 species having been recorded. i s recognised in f i v e Grand Banks and seven Scotian Shelf wells. Son3nulacysta j u r u s s i c a Assemblage Zone Type s e c t i o n : Shell Naskapi N-30, 6390-6590 f t . Other wells: Argo F-38, B i t t e r n M-62, Cohasset D-42, Dauntless D-35, Egret K-36, Eider M-75, Flying Foam 1-13, Heron H-73, Mic Mac 5-77, Missisauga H-54, Mohawk 8-93, Mohican 1-100, Murre G-67, Oneida 0-25, Wyando t E -53. Selected species bases: D i n o f l a g e l l a t e s ; E n d o s c r i n i m Zuridur; (Deflandre) Gocht, Gonyaulacysta granulata ( K 1 ement) S a r j e a n t , C;. granuZigera ( K 1 ement) Sarjeant , Leptodiniwn egemenii G i tmez, S c r i n i o d i n i ' m dietyotun? Cookson and The spores Cicatricosisporites a u s t r a l i e n s i s (Cookson) Eisenack.
Potoni6, De%Soispo?iteS v e l a t u s Weyland and Krieger, and P i l o s i s p o r i t e s sp. A ( p l a t e 2 , f i g u r e 1 ) appear in the upper p a r t of t h i s zone. Selected species t o p s : D i n o f l a g e l l a t e s ; A d n a t o s p h a e r i d i m cau2ler;ii
334
(Def 1 andre) Wi 11 iams and Downie, C o r d o s p h a e r i d i m costatwn, Ctenidodinium o m a t w r , E n d o s c r i n i w eisenacki, Gen. e t S p . 2 Gocht, 1970, Gonyaulacdsta j w a s sica
.
Discussion: This zone i s dominated by Gonyaulacysta iurassica on the northeastern Grand Banks. Elsewhere species of C t e n i d o d i n i m replace -i. r . ;urassica . as the dominant form. Marine conditions apparently preThe age of vailed t h r o u g h o u t offshore eastern Canada during t h i s time. t h i s zone i s discussed by Williams (1975), who assigned an Oxfordian, possibly in p a r t Early Kimmeridgian, age. Gonyaulacysta cladophora Assemblage Zone
Type s e c t i o n : Shell Oneida 0-25, 10,005-11,000 f t . Other w e l l s : Argo F-38, B i t t e r n M-62, Cohasset D-42, Cormorant N-83, Dauntless D-35, Egret K-36, Eider M-75, Flying Foam 1-13, Mic Mac 5-77, Missisauga H-54, Mohawk 8-93, Mohican 1-100, Murre G-67, Naskapi N-30, Wy an do t E - 53. Selected species bases: D i n o f l a g e l l a t e s ; Dingodiniim jurassieuv Cookson and Ei senack , E p i p l o s p h a e r a r e t i c u l o s p i n o s a K1 ement , I m b a t o d i n i m kondratjeo’i Vozzhenni kova, PoZystephanophorus sarjeantii Gi tmez, P s a l i gon~ou7,axa p a t e l a (Cookson and Eisenack) S a r j e a n t .
The d i n o f l a g e l l a t e s
4 q 1 h o r u h metaelliptica Dodekova, C t e n i d o d i n i m culrnulm ( H o r r i s ) Lentin
and Williams, C. p a n n e m ( H o r r i s ) Lentin and Williams, and Shsternatophora schindewolfi ( A l b e r t i ) Downie a n d Sarjeant appear in the upper p a r t o f
t h i s zone. Spores; Pilosisporites trichopapillosus ( T h i e r g a r t ) Delcourt and Sprumont. Selected species tops: Dinoflagellates; Endoscriniwn luridwn, GonjauZaq s t a cladophora, G. granuligera, Hexagonifera jurassica, L e p t o d i n i w n
egemenii, Systernatophora turonica ( A 1 b e r t i ) Downie and S a r j e a n t . Discussion: Dinoflagellate species d i v e r s i t y reaches a maximum (49) f o r
The zone i s c l e a r l y delineated the J u r a s s i c in the I;. cladophora Zone. by the f i r s t appearance of 18 species and t h e l a s t occurrence of 27 species. Dinoflagellates a r e c o n s i s t e n t l y present, permitting recognit i o n of the zone in 16 o f the 1 7 wells havins a Late J u r a s s i c s e c t i o n . Kimmeridgian d i n o f l a g e l l a t e s nave been well documented from Europe (Gitmez, 1970; Gitmez and S a r j e a n t , 1972) and show close a f f i n i t i e s with assemblages from the G. cladopkora Zone. J u s t i f i c a t i o n of the Kimmeridgian age assignment f o r t h i s zone i s given in Williams (1975).
335
Ctenidodiniwn panneuv Assembl age Zone Type s e c t i o n : S h e l l Mohawk 8-93, Other w e l l s :
5315-5395 f t .
Argo F-38, Cohasset D-42, Cormorant N-83, D a u n t l e s s D-35,
E g r e t K-36, F l y i n g Foam 1-13, M i c Mac 5-77, M i s s i s a u g a ti-54,
?Murre
G-67, Naskapi N-30, Oneida 0-25, Wyandot E-53. S e l e c t e d s p e c i e s bases: D i n o f l a g e l l a t e s ; P y z i d i e l l a sp. W i l l i a m s , 1975. S e l e c t e d s p e c i e s t o p s : D i n o f l a g e l l a t e s ; Amphoruh
metaelliptica,
Ctenidodiniwn culrnulwn, C. pannewn, Systematophora areolata Klement. Spores ; Couperisporites j u r a s s i c u s Pocock, Dictyophy ZZidites equiexinus (Couper) Dettmann, Endosporites jurassicus Pocock, P i l o s i s p o r i t e s s p . A,
Tril o bos por i t es j u r a s s i c u s . D i s c u s s i o n : D i n o f l a g e l l a t e assemblages f r o m t h i s zone d i f f e r f r o m t h o s e o f t h e u n d e r l y i n g L a t e J u r a s s i c zones i n c o n t a i n i n g f e w e r s p e c i e s .
This,
t o g e t h e r w i t h t h e p a u c i t y o f specimens, p o s s i b l y r e f l e c t s l o c a l changing environmental conditions, apparently r e s u l t i n g from a marine regression. However, s i m i l a r o b s e r v a t i o n s i n o t h e r areas suggest a w o r l d w i d e d e c l i n e phase i n d i n o f l a g e l l a t e p o p u l a t i o n s a t t h i s t i m e .
O n l y 12 J u r a s s i c
d i n o f l a g e l l a t e s p e c i e s p e r s i s t i n t o t h e Cretaceous i n o f f s h o r e e a s t e r n Canada.
The Cte nidodini-m pannewn Zone has been d a t e d P o r t l a n d i a n by
W i l l i a m s (1975) on t h e b a s i s o f t h e d i n o f l a g e l l a t e s p r e s e n t .
T h i s zone,
widespread on t h e S c o t i a n S h e l f and p r e s e n t on t h e e a s t e r n Grand Banks i s g e n e r a l l y absent on t h e w e s t e r n and c e n t r a l Grand Banks. Several S c o t i a n S h e l f w e l l s c o n t a i n a d i s t i n c t i v e spore assemblage between 200 and 300 ft. above t h e C. panneum Zone.
This i s t y p i c a l l y
c h a r a c t e r i s e d by numerous specimens o f CalZiaZasporites ciampieri,
Cont i gni s por i t es cooksonii (Balme) Dettmann, Densoisporites v e l a t u s , Kl uki s por i t es pse udore tic ulatus and sometimes an abundance o f Coro Zlina t or os us , as i n Argo F-38.
The i n t e r v a l i s p r o v i s i o n a l l y i n c l u d e d i n t h e
B e r r i a s i a n because o f l a c k o f d a t a r e g a r d i n g i t s t r u e age, whereas t h e age o f t h e C. pmnewn Zone can be j u s t i f i e d by comparison w i t h s o u t h e r n England. PALEOECOLOGY Lower J u r a s s i c s t r a t a o n t h e S c o t i a n S h e l f commonly c o n t a i n abundant spores o f low d i v e r s i t y and a r e d e v o i d o f d i n o f l a g e l l a t e s .
During t h e
E a r l y J u r a s s i c , t h e E u r y d i c e , Argo, and I r o q u o i s Formations were d e p o s i ted.
The E u r y d i c e F o r m a t i o n i s d a t e d R h a e t i a n - H e t t a n g i a n and, a c c o r d -
i n g t o Jansa and Wade (1975) accumulated under p r e d o m i n a n t l y a r i d cond i t i o n s i n a d e s e r t environment.
The o v e r l y i n g Argo Formation, c o n s i s -
336 t i n g m a i n l y o f s a l t d e p o s i t s , i s a s s i g n e d a H e t t a n g i a n - S i n e m u r i a n a g e and i s p o s s i b l y c o e v a l w i t h t h e Louann S a l t o f t h e G u l f o f M e x i c o ( H . S a b r y , personal communication).
M a r g i n a l l y m a r i n e c o n d i t i o n s have been sugges-
t e d f o r t h e s u c c e e d i n g I r o q u o i s F o r m a t i o n ( J a n s a and Wade, 0. cit.), which i s h e r e i n dated Sinemurian-early Pliensbachian. M a r i n e 1a t e P1 i e n s b a c h i an-Aal e n i a n s e d i m e n t s (,"Janvoceratopsis
gracC2is Zone), r e c o g n i s e d i n f i v e Grand Banks w e l l s , c o n t a i n s e v e r a l d i a g n o s t i c d i n o f l a g e l l a t e species.
These s p e c i e s a r e a b s e n t f r o m t h e
examined S c o t i a n S h e l f w e l l s , s o t h a t i t i s n o t c e r t a i n i f l a t e P l i e n s b a c h i a n and A a l e n i a n s e d i m e n t s w e r e d e p o s i t e d i n t h i s a r e a .
Bajocian
d i n o f l a g e l l a t e s ( ; k m c o d i w i w sepitczbulatwn Zone) h a v e been o b s e r v e d i n f o u r Grand Banks w e l l s , b u t i n o n l y two S c o t i a n S h e l f w e l l s , Argo F-38 and Mohican 1-100.
Species d i v e r s i t y i s low, making c o r r e l a t i o n w i t h
t h e European t y p e s e c t i o n s d i f f i c u l t .
From B a t h o n i a n t o K i m m e r i d g i a n
t i m e , m a r i n e c o n d i t i o n s i n c r e a s i n g l y p r e v a i l e d on b o t h t h e S c o t i a n S h e l f and Grand Banks.
This i s r e f l e c t e d i n t h e steady increase i n dino-
f l a g e l l a t e s p e c i e s d i v e r s i t y , w h i c h a t t a i n e d a peak i n t h e K i m m e r i d g i a n (text-figure 7).
The d e p o s i t i o n a l e n v i r o n m e n t i s i n t e r p r e t e d a s
predominantly i n n e r n e r i t i c .
More r e s t r i c t e d m a r i n e t o n o n - m a r i n e
environments c h a r a c t e r i s e d t h e P o r t l a n d i a n and p r e v a i l e d i n t o t h e E a r l y C r e t a c e o u s on t h e S c o t i a n S h e l f .
The Grand Banks s u c c e s s i o n a t t h i s
time i s complicated by a major unconformity.
ACKNOWLEDGEMENTS The a u t h o r s a r e g r a t e f u l t o Hassan S a b r y f o r f r u i t f u l d i s c u s s i o n s o n E a r l y J u r a s s i c p a l y n o s t r a t i g r a p h y , t o M. S. B a r s s , D. Umpleby and
J. Wade f o r t h e i r c o n s t r u c t i v e c r i t i c i s m o f t h i s p a p e r , a n d t o G. L. Cook a n d G. M. G r a n t f o r d r a f t i n g t h e t e x t - f i g u r e s .
PLATE 1
M a g n i f i c a t i o n shown on f i g u r e s .
1.
Corollina qeleriana ( K l a u s ) V e n k a t a c h a l a and G6czhn. S h e l l E u r y d i c e P-36, c u t t i n g s s a m p l e 2840-2880 f t : H e t t a n g i a n - S i n e GSC No. 47929. murian.
2.
KraeuseSispcrites r e i s s i n g e r i ( H a r r i s ) Morbey. Shell Eurydice P-36, c u t t i n g s sample 2840-2880 f t : H e t t a n g i a n - S i n e m u r i a n . GSC No. 47930.
3,4.
k c h i n < t o s p r i : e s c f . i l i a c o i d e s S c h u l z and K r u t z s c h . Shell E u r y d i c e P-36, c u t t i n g s sample 2840-2880 f t : H e t t a n g i a n - S i n e GSC No. 47931 and 47932. murian.
337
REFERENCES American Commission on S t r a t i g r a p h i c Nomenclature, 1961. Code o f s t r a t i g r a p h i c nomenclature: A m . Assoc. P e t r o l . Geol. B u l l . , V . 4 5 , p . 645-660. C o r n e t , B . and T r a v e r s e , A., 1975. P a l y n o l o g i c a l c o n t r i b u t i o n s t o t h e chronology and s t r a t i g r a p h y of t h e H a r t f o r d Basin i n C o n n e c t i c u t and M a s s a c h u s e t t s : Geoscience and Man, v . 1 1 , p . 1-33. Dodekova, L . , 1975. New Upper Bathonian d i n o f l a g e l l a t e c y s t s from n o r t h e a s t e r n B u l g a r i a : P a l e o n t . S t r a t i g r . and L i t h o l . , v . 2 , p . 17-32. G e i g e r , M . E . and Hopping, C . A . , 1968. T r i a s s i c s t r a t i g r a p h y of t h e s o u t h e r n North Sea b a s i n : P h i l o s . T r a n s . R . S O C . London, s e r B , v. 254, p . 1-39. Gitmez, G . U . , 1970. D i n o f l a g e l l a t e c y s t s and a c r i t a r c h s from t h e b a s a l Kimmeridgian (Upper J u r a s s i c ) o f England, S c o t l a n d and France: B u l l . Br. Mus. n a t . H i s t . ( G e o l . ) , v . 1 8 , p . 231-331. Gjtmez, G.U. and S a r j e a n t , W.A.S., 1972. D i n o f l a g e l l a t e c y s t s and a c r i t a r c h s from t h e Kimmeridgian (Upper J u r a s s i c ) o f England, S c o t l a n d and F r a n c e : B u l l . Br. Mus. n a t . H i s t . ( G e o l . ) , v . 2 1 , p . 171-257. Gocht, H . , 1970. D i n o f l a g e l l a t e n - Z y s t e n aus dem Bathonium des E r d o l f e l d e s Aldorf (NW-Deutschland): P a l a e o n t o g r a p h i c a , Abt. B , v . 129, p . 125-165. PLATE 1
338
Herngreen, G.F.W. and de Boer, K.F., 1974. Palynology of Rhaetian, Liassic and Dogger strata in the Netherlands with emphasis on the Achterhoek area:Geologie en Mijnbouw, v. 53, no. 6, p. 343-368. Jansa, L.F. and Wade, J.A., 1975. Geology o f the continental margin off Nova Scotia and Newfoundland; i n Offshore Geology o f Eastern Canada, Vol. 11: Geol. Surv. Can., Paper 74-30, v. 2 , p. 51-105. Morgenroth, P., 1970. Dinoflagellate cysts from the Lias Delta o f Luhndel Germany: Neues Jahrb. Geol. Palxontol., Abh., v. 136, no. 3, p.345359. Pocock, S.A.J., 1970. Palynology o f the Jurassic sediments o f western Canada. Part 1 : Terrestrial species: Palaeontographica, Abh. B, V. 130, p. 12-72. Scheuring, B.W., 1970. Palynologische und palynostratigraphische Untersuchungen des Keupers im Bolchentunnel (Solothurner Jura): Schweiz. PalB’ont. Abh., v. 88, p. 1-119. Williams, G.L., 1975. Dinoflagellate and spore stratigraphy of the Mesozoic-Cenozoic, offshore eastern Canada; i n Offshore Geology o f Eastern Canada, Vol. 1 1 : Geol. Surv. Can., Paper 74-30, v . 2 , p. 107161,
, In Press. Dinocysts: Their palaeontology, biostratigraphy and palaeoecology; in Oceanic Micropalaeontology, Academic Press, London, England.
PLATE 2 1.
Figures 1-3 x 440; Figures 4-12 x 715.
PlZosisporites sp. A.
Mobil-Tetco Cohasset D-42, cuttings sample 10,570-10,600 ft: Kimmeridgian. GSC No. 47933. ContignisporTites sp. Mobil-Tetco Cohasset D-42, cuttings sample 2. 11,070-11,100 ft: Kimmeridgian. GSC No. 47934. 3. Verrucosisporites sp. A. Shell Eurydice P-36, cuttings sample 6600-6640 ft: Hettangian-Sinemurian. GSC No. 47935. 4. ? C y m d o p i t e s sp. A. Shell Eurydice P-36, sidewall core 5048 ft: Hettangian-Sinemurian. GSC No. 47936. 5,6 Krcleuselisporites reissingeri (Harris) Morbey, Shell Eurydice P-36, sidewall core 5048 ft: Hettangian-Sinemurian. GSC No. 47937. 7,8 ?Cgcadopites sp. B.7, Shell Eurydice P-36, sidewall core 2911 ft: Hettangian-Sinemurian. 8, Shell Eurydice P-36, sidewall core 3229 ft: Hettangian-Sinemurian. GSC No. 47938 and 47939. 9,12 Z c h i n i t o s p o r i t e s cf. i l i a c o i d e s Schulz and Krutzsch. Shell Eurydice P-36, sidewall core 2575 ft: Sinemurian-early Pliensbachian. GSC No. 47940 and 47941. 10,ll E c h i n i t o s p o r i t e s sp. A . Shell Eurydice P-36, cuttings sample 2840-2880 ft: Hettangian-Sinemurian. GSC No. 47942.
339
PLATE 2
This Page Intentionally Left Blank
341 COMPARISON OF LOWER AND MIDDLE CRETACEOUS PALYNOSTRATIGRAPHIC ZONATIONS I N THE WESTERN NORTH ATLANTIC
D a n i e l Habib Department o f E a r t h and Environmental S c i e n c e s Queens C o l l e g e , N e w York C i t y
ABSTRACT Cretaceous palynomorph z o n a t i o n s , based r e s p e c t i v e l y on t h e s t r a t i g r a p h y of d i n o f l a g e l l a t e c y s t s and sporomorphs, are p r e s e n t e d f o r s i x Deep Sea D r i l l i n g P r o j e c t s i t e s i n t h e weste r n North A t l a n t i c . E i g h t d i n o f l a g e l l a t e zones and f o u r sporomorph zones are d e s c r i b e d from t h e r e f e r e n c e s e c t i o n a t s i t e 105, which r a n g e s i n age from B e r r i a s i a n ( c o r e 30) t o Cenomanian ( c o r e 9 ) . P u b l i s h e d n a n n o f o s s i l and p l a n k t o n i c f o r a m i n i f e r a 1 s t u d i e s of s i t e 105 p e r m i t t h e d i r c c t comparison of p a r t of t h e palynomorphic z o n a t i o n s w i t h a p u b l i s h e d g e o c h r o n o l o g i c a l s c a l e . The sporomorph z o n a t i o n i s compared w i t h t h a t publ i s h e d f o r t h e l a r g e l y nonmarine Lower Cretaceous f a c i e s of t h e a d j a c e n t U. S . A t l a n t i c c o a s t a l p l a i n . INTRODUCTION
Zonations a r e proposed f o r two groups o f palynomorphs, d i n o f l a g e l l a t e c y s t s and sporomorphs, o c c u r r i n g i n Lower and Middle C r e t a c e o u s s e c t i o n s r e c o v e r e d a t s i x
s i t e s i n t h e w e s t e r n North A t l a n t i c by t h e Deep Sea D r i l l i n g P r o j e c t ( F i g u r e 1 ) . The d i n o f l a g e l l a t e z o n a t i o n p u b l i s h e d by Habib (1976) f o r Neocomian assemblages i s rev i s e d i n p a r t and extended t o t h e Cenomanian, and a sporomorph z o n a t i o n r a n g i n g from B e r r i a s i a n t o Cenomanian
i s proposed.
The z o n a t i o n s a r e based p r i m a r i l y on t h e
s t r a t i g r a p h y a t s i t e 1 0 5 , a l t h o u g h two d i n o f l a g e l l a t e subzones are i n c l u d e d from species which are s t r a t i g r a p h i c a l l y c o n t i n u o u s a t s i t e 391 o n l y .
The d i n o f l a g e l l a t e
and sporomorph z o n a t i o n s e s t a b l i s h e d a t s i t e 105 are corr e l a t e d f o r a t l e a s t p a r t o f t h e s e c t i o n , through s i t e s
342
9o"w 60"N
10 " N
90"w
E 1" W 6O"N
*IO0N 6 0"W
F i q u r c 1. G ~ o o r a p h i cd i s t r i b u t i o n o f i n v e s t i g a . L c d s i t e s , Deep Sea D r i l l i n g P r o j e c t .
343
99, 100, 101, 387, and 391 (cf. Habib, 1976, Text-Figure 4).
A total of 161 samples were studied, collected from 2 1 cores representing the reference section at site 105.
This interval is approximately 250 meters thick, ranging from the stratigraphic level of core 30 (Berriasian) through that of core 9 (Cenomanian). The cores immediately below and above the investigated interval are essentially devoid of palynofossils.
How-
ever, the investigated interval is highly palyniferous, with respect to dinoflagellates, sporomorphs, and small acritarchs.
The latter group is currently under i.nvesti-
gation. BIOSTRATIGRAPHY The ranges of 65 species of dinoflagellates and sporomorphs in the Lower and Middle Cretaceous section at DSDD site 105 are presented in Figure 2.
These species are
considered to be of chronostratigraphic value, based on their stratigraphic ranges in the section or their ages as derived from the literature.
Separate dinoflagellate and
sporomorph zonations are proposed from the ranges of selected stratigraphically continuous species at site 1 0 5 (Figure 3 ) .
Separate zonations are preferable to a single
combined palynomorph zonation for the following reasons. First, a separate sporomorph zonation permits the potential correlation of North Atlantic sections with the largely nonmarine Lower Cretaceous of the adjacent U.S. Atlantic Coastal Plain, where dinoflagellates are rare or absent.
Also, in the study of a number of sections
within the North Atlantic the relative abundance of
TRITHYRODNIUM SUY'ECTUM DEFLANDREA ECHINCIDEA
VESTITA
ODONTOCHITINA WERCULATA DRWjGlDUM RHABDCRETICULATUM
Y
n
I
CLAVATIPOLLENITES
I
I
4
DRUGG1 U M DEFLANDREI
EPHEDRMTES
DRUGGIDUM
WICOPAUCICUM
BICRBFERP
---
JOHNEWINGII
Figure 2. Stratigraphic range chart and zonations at site 105. Sporomorph taxa indicated by a .
-
MULTICOSTATUS
345
SPOROM ORPHS
DINOFLAG E LLAT ES
STRITHYRODINIUMI
SUSPECTUM DEFLANDREA ECHINOlDEA
C OMPLEXlOPOLL l S
cn W
PSILATRICOLPORITES
DEFLANDREA VESTITA TRICOLPITES MINUTUS
ODONTOCHITINA
C L AVATl P OL L ENITES
0 PE RCUL ATA DRUGGIDIUM R HA BDORET IC ULAT UM
0RUGGIDlUM DEFLANDREI E PHEDRlPlT ES DRUGGIDIUM A PIC0PAUCICUM
M ULT IC OSTAT US
BlORBlF ERA JOHN EW lNGll Figure 3. Dinoflagellate and sporomorph zonations at site 105.
346
dinoflagellates and sporomorphs varies, in some cases with one palynomorph group represented to the almost total exclusion of the other. The Cretaceous (Berriasian-Cenomanian) section at site 105 is valuable as a reference section for palynomorph stratigraphy because of the number of cores recovered, the small stratigraphic intervals between collected samples, the paleontological ages provided by other groups of fossils, and that by ranging from Berriasian to Cenomanian, it is the most complete section.
The section
is dated in large part 6n the basis of the nannofossil studies of Thierstein ( 1 9 7 6 ) , Wilcoxon ( 1 9 7 2 ) , and
Bukry
(1972);
the planktonic foraminifers studied by Luterbacher
(1972);
and the ammonite aptychi studied by Renz ( 1 9 7 2 ) .
It is dated otherwise by dinoflagellate cysts, from the study of Lower Cretaceous stratotype sections by Millioud (1967,
1 9 6 9 ) and Davey and Verdier (1971, 1 9 7 4 ) or inde-
pendently dated sections (e.g. Verdier, 1 9 7 5 ) . The interval represented by cores 33-22 has been dated as Berriasian-Valanginian (Thierstein, 1976, figure 5).
Thierstein (1976) dated core 20 within the ages of
early Hauterivian to early Barremian.
Core 18 has been
dated by ammonite aptychi within the ages of late Valanginian-Hauterivian (Renz, 1972) and by nannofossils as Hauterivian (Wilcoxon, 1972; Bukry, 1 9 7 2 ) .
It is con-
sidered in this paper to be not older than late Hauterivian based on the first appearance of the dinoflagellate Odontochitina operculata (Wetzel), which has its lowest
occurrence in the uppermost Hauterivian in the Angles section of France, according to Millioud ( 1 9 6 9 ) .
Core 1 7
is considered to be of Barremian or Aptian age.
Wilcoxon
347
(1972) dated the core as Barremian on the basis of nannofossils, and Luterbacher (1972) recovered an assemblage of small and primitive foraminifers which he stated are similar to faunules occurring in sections ranging from late Hauterivian to early Aptian. The interval represented by cores 16-11 is considered to be Albian, based on the stratigraphic range of the dinoflagellate D e f l a n d r e a v e s t i t a (Brideaux). According to Sverdlove and Habib (1974) this species became phyletically extinct in the time represented by core 11, which has been dated on the basis of foraminifers as latest (Vraconian) Albian (Luterbacher, 1972).
This age assign-
ment is supported by the first occurrence of tricolpate pollen grains, e.g. R e t i t r i c o l p i t e s g e o r g e n s i s Brenner, in core 16.
Core 9 is considered to be of Cenomanian age,
based on the first appearance of the dinoflagellate T r i t h y r o d i n i u m s u s p e c t u m (Manum and Cookson) and on the
occurrence of the first Nomapolles pollen genera, C o m p l e x i o p o l l i s and A t l a n t o p o l l i s .
DINOFLAGELLATE ZONATION Eight dinoflagellate zones are distinguished in the reference section at site 105, based on the stratigraphic first appearance or phylogenentic appearance of a selected dinoflagellate species defining the lower boundary of the corresponding zone (Figure 2).
The Neocomian zonation
proposed by Habib (1976) in the western North Atlantic is extended to the Cenomanian Stage, with the description of the following new zones:
DEFLANDREA VESTITA, DEFLAN-
DREA ECHINOIDEA, TRITHYRODINIUM SUSPECTUM Zones.
348
The OLIGOSPHAERIDIUM COMPLEX Zone proposed by Habib (1976) is modified, based on the dinoflagellate stratigraphy occurring elsewhere in the North Atlantic.
Its
two original subzones, the lower DRUGGIDIUM RHABDORETICULATUM and higher ODONTOCHITINA OPERCULATA, are proposed as zones in its place because they are correlatable at sites for which there is adequate core recovery, e.g. site Also at site 391 (Habib, i n p r e s s ) , O l i g o s p h a e r i d i u m
391.
c o m p l e x (White) appears stratigraphically below the first
occurrence of D r u g g i d i u m r h a b d o r e t i c u l a t u m . 1. BIORBIFERA JOHNEWINGII Zone.
Defined as the interval from the lowest occurrence of B i o r b i f e r a j o h n e w i n g i i up to (but not including) the first appearance of D r u g g i d i u m a p i c o p a u c i o u m (Habib, 1976).
This interval ranges from the stratigraphic levels
represented by samples DSDP-105-30-2, 90-92 cm. through 105-27-2, 64-66 cm.
The zone is dated as Berriasian to
early Valanginian, based on the nannofossil zonation published by Thierstein (1976) for this site. A number of species which are restricted to the Neocomian in the western North Atlantic have their first stratigraphic appearance in the BIORBIFERA JOHNEWINGII zone.
In addition to the nominative species, these in-
clude D i a c a n t h u m h o l l i s t e r i i Habib, and P y x i d i n o p s i s c h a l l e n g e r e n s i s Habib.
The Upper Jurassic species
Amphorula m e t a e l l i p t i c a Dodekova has its highest strati-
graphic occurrence in this zone.
Other species which
appear in the zone include P o l y s p h a e r i d i u m w a r r e n i i Habib, G o n y a u l a c y s t a h e l i c o i d e a (Eisenack and Cookson) and
349
W a l l o d i n i u m k r u t z s c h i i (Alberti).
Stratigraphically per-
sistent species which appear in the BIORBIFERA JOHNEWINGII Zone at site 391, but which were not found at sites 99, 100, 101, and 105, include P h o b e r o c y s t a n e o c o m i c a (Gocht)
and P s e u d o c e r a t i u m p e l l i f e r u m Gocht. M.
Muderongia sp. cf.
s i m p l e x Alberti was found restricted to this zone at
site 391 only, as well. 2.
DRUGGIDIUM APICOPAUCICUM Zone.
Defined by Habib (1976) as the interval from the lowest occurrence of the nominative species up to the phylogenetic appearance of D r u g g i d i u m d e f l a n d r e i (Millioud). At site 105, this zone is represented as the interval from samples DSDP-105-27-2, 13-15 cm. through 105-24-1, 40-42 an.
It is dated as early Valanginian to
late Valanginian based on the nannofossil zonation of site 105 (Thierstein, 1976).
S c r i n i o d i n i u m a t t a d a l e n s e Cookson and Eisenack, S. campanula Gocht, D i n g o d i n i u m c e r v i c u l u m Cookson and Eisenack, C y c l o n e p h e l i u m d i s t i n c t u i n Deflandre and Cookson, and H y s t r i c h o d i n i u m v o i g t i i (Alberti) appear in the DRUGGIDIUM APICOPAUCICUM Zone at site 105. 3.
DRUGGIDIUM DEFLANDREI Zone. Phylozone characterized as the interval from the
phylogenetic appearance of D r u g g i d i u m d e f l a n d r e i up to the phylogenetic appearance of D . r h a b d o r e t i c u l a t u m Habib. At site 105, O l i g o s p h a e r i d i u m c o m p l e x appears at the same stratigraphic horizon as D . r h a b d o r e t i c u l a t u m ; elsewhere
350
it appears at a lower level (Habib, i n p r e s s ) .
In the
western North Atlantic, the stratigraphic ranges of B i o r b i f e r a j o h n e w i n g i i , P y x i d i n o p s i s c h a l l e n g e r e n s i s , and D i a c a n t h u m h o l l i s t e r i i terminate within this zone. M e i o u r o g o n y a s u l a x stoveri Millioud first occurs in this zone at site 105. The DRUGGIDIUM DEFLANDREI Zone is represented as the interval from samples DSDP-105-23 core catcher through 105-20-1, 101-103 cm.
It is considered to range in age
from late Valanginian at its lower boundary (cf. Thierstein, 1976) to Hauterivian (or early Barremian). 4. DRUGGIDIUM RHABDORETICULATUM Zone.
New zone defined as the interval from the phylogenetic appearance of the nominative species up to the stratigraphic appearance of O d o n t o c h i t i n a o p e r c u l a t a .
At
site 105, this zone is represented as the interval from samples DSDP-105-20-1, 63-65 cm. to core 18.
The zone is
dated as Hauterivian or Barremian. This zone was originally described as a subzone by Habib (1976).
It is elevated to the status of a zone
based on its geographic range in the western North Atlantic (Habib, i n p r e s s ) . 5.
ODONTOCHITINA OPERCULATA Zone. New zone defined by the interval from the strati-
graphic appearance of the nominative species up to the
.
appearance of D e f l a n d r e a v e s t i t a (Brideaux)
This inter-
val is represented at site 105 from the stratigraphic
351
level of sample DSDP-105-18 core catcher through that of sample 105-16 core catcher. It is dated as late Hauterivian or Barremian to early Albian.
The zone was origin-
ally described as a subzone (Habib, 1976).
Subtilisphaera
p e r l u c i d a (Alberti) is restricted to the ODONTOCHITINA
OPERCULATA Z m e in the western North Atlantic appearing at its base and disappearing at or very near the upper boundary.
At site 391, P h o b e r o c y s t a n e o c o m i c a and
P s e u d o c e r a t i u m p e l l i f e r u m disappear within the zone.
The
stratigraphic disappearances of P . n e o c o m i c a and S . p e r l u c i d a within the ODONTOCHITINA OPERCULATA Zone at site 391 serve to divide it into two subzones there (Figure 4 ) ;
a lower PHOBEROCYSTA NEOCOMICA Subzone which on the basis of dinoflagellate evidence is dated as late Hauterivian or Barremian to early Aptian, and a higher SUBTILISPHAERA PERLUCIDA Subzone which is dated as Aptian to early Albian (Habib, i n p r e s s ) . In the western North Atlantic, C a l l a i o s p h a e r i d i u m a s y m m e t r i c u m (Deflandre and Courteville) appears within
the ODONTOCHITINA OPERCULATA Zone (and within the PHOBEROCYSTA NEOCOMICA Subzone at site 391).
A number of species
either appear or disappear stratigraphically in the upper half of the zone in the western North Atlantic.
Those
which disappear include W a l l o d i n i u m k r u t z s c h i i , P o l y s p h a e r i d i u m w a r r e n i i , Druggidium a p i c o p a u c i c u m , D. def l a n d r e i , D i n g o d i n i u m c e r v i c u l um, M e i o u r o g o n y a u l a x stoveri
and G o n y a u l a c y s t a c a s s i d a t a (Eisenack and Cookson)
.
Species which appear stratigraphically in the upper half are C r i b r o p e r i d i n i u m m u d e r o n q e n s i s (Cookson and Eisenack), Hystrichokolpoma ferox
(Deflandre), C h l a m y d o p h o r e l l a nyei
(Cookson and Eisenack), C l e i s t o s p h a e r i d i u m a n c o r i f e r u m (Cookson and Eisenack) , S p i n i f e r i t e s c i n g u l a t u s (Wetzel),
352
S. ramsus
(Ehrenberg), H y s t r i c h o s p h a e r i d i u m arundum
Eisenack and Cookson, and H . cooksonii Singh.
This pheno-
menon apparently represents a large evolutionary change of dinoflagellate species near the boundary between the ODONTOCHITINA OPERCULATA and overlying DEFLANDREA VESTITA Zmes.
Alternatively, it may be related to the change of
lithofacies in the western North Atlantic, from nannoplankton ooze below to carbonaceous clay above (Lancelot, Hathaway, and Hollister, 1972).
This change in species
and lithofacies occurs within the ODONTOCHITINA OPERCULATA one elsewhere in the North Atlantic as well, e.g. sites 99 and 101 (OLIGOSPHAERIDIUM COMPLEX Zone of Habib, 1976).
6.
DEFLANDREA VESTITA Zone.
New zone defined as the interval from the appearance of the nominative species below up to the phylogenetic appearance of D e f l a n d r e a e c h i n o i d e a Cookson and Eisenack above.
It is represented at site 105 from the strati-
graphic level of sample DSDP-105-16-2, 110-112 cm. through that of 105-11-3, 20-22cm.
Sample DSDP-105-11-2, 75-77
cm. contains the first morphotype assigned to D . e c h i n o i d e a ; sample 1 0 5 - 1 1 - 1 ,
morphotype of D . v e s t i t a .
130-132 cm. contains the last
This zone is dated as Albian to
latest (Vraconian) Albian. The following species appear in the DEFLANDREA VESTITA Zone: G o n y a u l a c y s t a e x i l i c r i s t a t a Davey, H e x a g o n i f e r a c h l a m y d a t a Cookson and Eisenack, P a l a e o h y s t r i c h o p h o r a i n f u s o r i o i d e s Deflandre, L i t o s p h a e r i d i u m s i p h o n i p h o r u m
(Cookson and Eisenack), Achomosphaera r a m u l i f e r a (Deflandre), E p e l i d o s p h a e r i d i a s p i n o s a Davey, Nystricho-
353
s p h a e r o p s i s ovum Deflandre, and P r o t o e l l i p s o d i n i u m sp. P h o b e r o c y s t a c e r a t i o i d e s (Deflandre) was observed in this
zone at sites 101 and 391. 7.
DEFLANDREA ECHINOIDEA Zone. New zone defined as the interval from the phylo-
genetic appearance of the nominative species below up to the stratigraphic appearance of T r i t h y r o d i n i u m s u s p e c t u m . At site 105, this interval is represented from the level of sample DSDP-105-11-2, 75-77 cm. through that of 105-96, 80-82 cm.
It is dated as latest Albian-Cenomanian.
Species which appear in this zone include T r i g o n o p y x i d i a g i n e l l a (Cookson and Eisenack) , D i p l o f u s a g e a r -
l e n s i s Cookson and Eisenack, Dinogymnium a c u m i n a t u m Evitt,
Clarke, and Verdier, and S u b t i l i s p h a e r a p o n t i s - m a r i a e (Deflandre) 8.
.
TRITHYRODINIUM SUSPECTUM Zone.
New z m e defined from the appearance of the nominative species in core 9 (DSDP-105-9-6, 38-40 cm.).
The
interval above core 9 is not known, as the cores immediately above are essentially devoid of dinoflagellate fossils.
It is dated as Cenomnnian. M i c r o d i n i u m v e l i g e r u m
Davey occurs in this zone. The dinoflagellate zonation presented above is correlatable, at least in part, through sites 99, 100, 101, 387 and 391, as well as site 105.
Figure 4 shows the
zonation in the western North Atlantic within the framework of a chronostratigraphic scale.
354
I
TRI T H YRODI NI UM SUSPECTUM
CENOMANIAN VRACONIAN
ECHINOIDEA
!I !
DEFLANDREA
ALBIAN
VESTITA
SUBTILISPHAERA PE RLUCIDA
APTIAN
P H OB E ROCYS TA
N EOCOMICA
DRUG GI Dl UM RHABDORETICULATUM
BARREMIA N
HAUTERIVIAN
DR U GGlDlUM D E F L ANDRE1 DRU GGlDl UM API C OPAUCl C U M
BI ORBIFER A JOHNEWlNGll ~~
VA LA NGI NI A N
BERRlAS1A N
Figure 4. D i n o f l a g e l l a t e z o n a t i o n comvared a g a i n s t a c h r o n o s t r a t i graphic scale.
355
SPOROMORPH ZONATION Four sporomorph zones are distinguished based on the overlapping first occurrences of pollen species defining the lower boundaries of their respective zones.
In as-
cending stratigraphic order, these are the EPHEDRIPITES MULTICOSTATUS, CLAVATIPOLLENITES, RETITRICOLPITES GEORGENSIS, and COMPLEXIOPOLLIS Zones.
The RETITRICOLPITES GEOR-
GENSIS Zone is divided into a lower TRICOLPITES MINUTUS Subzone and a higher PSILATRICOLPORITES Subzone. Pollen in the sporomorph genus E p h e d r i p i t e s are reported from early Neocomian (Berriasian-Valanginian) sediments in the North Atlantic.
E p h e d r i p i t e s mu1 t i c o s t a t u s
Brenner is stratigraphically persistent in these sections and, hence, is considered to be a valuable zonal species. E p h e d r i p i t e s has long been considered a useful strati-
graphic indicator for dating sediments as Barremian or younger.
Couper (1964) indicated that "specimens of
E p h e d r a - l i k e pollen" enter the chronostratigraphic column
in the upper Hauterivian or lower Barremian.
Wolfe and
Pakiser (1971) indicated a maximal age around the Hauterivian-Barremian boundary for assemblages in the Atlantic Coastal Plain containing taxa such as E p h e d r i p i t e s (and C l a v a t i p o l l e n i t e s h u g h e s i i Couper)
.
In the Salisbury
Embayment of the Atlantic Coastal Plain, E p h e d r i p i t e s occurs stratigraphically well below the first specimens of Clavatipollenites.
In their study of the Bethards well in
the Salisbury Embayment, Robbins, Perry, and Doyle (1975) report E p h e d r i p i t e s in their lowest sample studied, up to approximately 730 meters below the lowest occurrence of C l a v a t i p o l l e n i t e s and just 40 meters approximately above
the base of the sedimentary section.
356
The stratigraphically lowest specimens assigned to C l a v a t i p o l l e n i t e s (as C l a v a t i p o l l e n i t e s sp. 1 in the North
Atlantic are characterized by monosulcate grains with a subcircular to elliptical outline in polar view.
The
exine is clearly tectate-columellate with a continuous tectum, as revealed in optical section.
These grains dif-
fer from C l a v a t i p o l l e n i t e s h u g h e s i i Couper in the continuity of the tectum layer, and from C l a v a t i p o l l e n i t e s cf. C. h u g h e s i i sensu Doyle, van Campo, and Lugardon (1975) which possesses an exine structure intermediate between semitectate and tectate-perforate. C l a v a t i p o l l e n i t e s sp. first appears in the western North Atlantic in assemblages of Barremian or late Hauterivian age (very near the base of the ODONTOCHITINA OPERCULATA zone at site 105) C. h u g h e s i i appears higher, in the SUBTILISPHAERA PERLUCIDA Subzone at site 391. Reticulate tricolpate and tricolporoidate pollen grains first appear in Albian sediments (DEFLANDREA VESTITA zone) in the North Atlantic, in species of the genera R e t i t r i c o l p i t e s and T r i c o l p i t e s .
the Cenomanian.
These species range into
They occur in successive horizons with
tricolporate species ( P s i l a t r i c o l p o r i t e s ) and triporate species of the Normapolles group ( C o m p l e x i o p o l l i s ) .
1. EPHEDRIPITES MULTICOSTATUS Zone New zone defined by the interval from the first appearance of the nominative species up to (but not including) the first occurrence of C l a v a t i p o l l e n i t e s .
It is
represented at site 105 as the interval from samples DSDP105-29 core catcher through 105-20-1, 63-65 cm.
357
A number of sporomorph s p e c i e s o c c u r i n t h e EPHEDRIPITES MULTICOSTATUS ZDne, which d a t e it as n o t o l d e r t h a n
Early Cretaceous.
These i n c l u d e L e p t o l e p i d i t e s e p a c r o r -
n a t u s N o r r i s , C i c a t r i c o s i s p o r i tes p o t o m a c e n s i s Brenner , C . b r e v i l a e s u r a t u s Couper, C . h u q h e s i i Dettmann, Monosul-
c i t e s q l o t t u s Brenner , A p p e n d i c i s p o r i t e s p r o b l e m a t i c u s ( B u r g e r ) , A . p o t o m a c e n s i s Brenner, and P a r v i s a c c i t e s r a -
d i a t u s Couper.
However, e x c e p t f o r M . g l o t t u s and p . r a -
d i a t u s , t h e s e s p e c i e s a r e r a r e o r have a d i s c o n t i n u o u s s t r a t i g r a p h i c range i n t h e w e s t e r n North A t l a n t i c , and hence, a r e of l i m i t e d v a l u e f o r z o n a t i o n .
N o r r i s (1969)
r e p o r t e d s e v e r a l of t h e s e s p e c i e s from t h e approximate p o s i t i o n of t h e J u r a s s i c / C r e t a c e o u s boundary i n t h e nonmarine Purbeck f a c i e s of s o u t h e r n England. The most abundant sporomorphs i n t h i s zone a r e spec i e s of b i s a c c a t e p o l l e n ( P i n u s p o l l e n i t e s , A l i s p o r i t e s ) and C l a s s o p o l l i s t o r o s u s ( R e i s s i n g e r ) , which t o g e t h e r comprise up t o 84 p e r c e n t of t h e sporomorph a s s i n b l a g e s .
E x e s i p o l l e n i t e s tumulus Balme, V i t r e i s p o r i t e s p a l l i d u s ( R e i s s i n g e r ) , E u c o m m i i d i t e s minor Groot and Penny, and
G i n k q o c y c a d o p h y t u s n i t i d u s ( B a l m e ) a r e common as w e l l .
2.
CLAVATIPOLLENITES Zone.
N e w zone d e f i n e d by t h e i n t e r v a l from t h e l o w e s t oc-
c u r r e n c e o f s p e c i e s i n C l a v a t i p o l l e n i t e s up t o t h e s t r a t i g r a p h i c appearance of R e t i t r i c o l p i t e s qeorqensis Brenner.
I t i s r e p r e s e n t e d a t s i t e 105 as t h e i n t e r v a l
from sample DSDP-105-18 c o r e c a t c h e r through sample 10516-2, 110-112 c m .
C l a v a t i p o l l e n i t e s sp.
,
C . huqhesii Couper, and C .
t e n e l l i s P a d e n - P h i l l i p s and F e l i x a l l o c c u r i n t h i s zone,
358
although only the first species was found in the lowest sample.
L i l i a c i d i t e s d i v i d u u s (Pierce), L . p e r o r e t i c u l a -
t u s (Brenner), and L. t r i c h o t o m s u l c a t u s Singh appear in
the upper half of the zone. C l a s s o p o l l i s t o r o s u s , P i n u s p o l l e n i t e s spp., and A l i s p o r i t e s b i l a t e r a l i s Rouse continue to dominate the spo-
romorph flora. 3.
RETITRICOLPITES GEORGENSIS Zone.
New zone defined by the interval from the first appearance of R e t i t r i c o l p i t e s g e o r g e n s i s up to the first appearance of C o m p l e x i o p o l l i s sp.
At site 105, the zone
is represented from the level of sample DSDP-105-16-2, 70-73 cm. to that of sample 105-9 core catcher.
zones are 2istinguished.
Two sub-
The lower subzone is defined
by the appearance of T r i c o l p i t e s m i n u t u s (Brenner), which appears in the lowest sample.
The higher subzone is de-
fined by the appearance of P s i l a t r i c o l p o r i t e s sp. (cf. "Tricolporopolleni t e s " t r i a n g u 1 u s Groot , Penny , and Groot)
in sample DSDP-105-13-2, 100-102 cm. In addition to R . g e o r g e n s i s and T . m i n u t u s , a number of tricolpate and tricolporoidate species appear at or very near the base of this zone.
These include R e t i t -
r i c o l p i tes s p h a e r o i d e s Pierce, R . m a g n i f i c u s Habib , T r i c o l p i t e s a u r i t u s (Bolkhovitina), T . micromunus (Groot and
Penny) , and P s i l a t r i c o l p i t e s p s i l a t u s Pierce.
These spe-
cies range through the RETITRICOLPITES GEORGENSIS Zcne into the next higher zone.
R u g u b i v e s i c u l i t e s rptluctus
Pierce (and R. r u g o s u s Pierce) first occur in the PSILATRICOLPORITES Subzone, along with S t r i a t o p o l l i s p a r a n e u s
359
‘Norris).Occurring
n e a r t h e t o p o f t h e PSILATRICOLPORITES
Subzone a r e t h e f i r s t t r i a n g u l a r o b l a t e t r i p o r a t e g r a i n s with s i m p l e , t h i c k e n e d p o r e s l o c a t e d a t t h e r a d i a l corners.
These p o l l e n a r e r e f e r r e d t o t h e genus Triporopol-
1eni tes
.
Classopollis torosus c o n t i n u e s t o be t h e s i n g l e most abundant sporomorph s p e c i e s n e a r t h e base of t h e RETITRICOLPITES GEORGENSIS Z m e .
However, w i t h i n t h e lower h a l f
of t h e zone, w i t h i n t h e TRICOLPITES MINUTUS Subzone, it d e c l i n e s markedly i n abundance and i s rare o r a b s e n t i n samples h i g h e r i n t h i s zone and i n t h e o v e r l y i n g zone. S p e c i e s of t r i c o l p a t e d i c o t y l e d o n o u s angiosperm p o l l e n become abundant i n i t s p l a c e and remain so through t h e RETITRICOLPITES GEORGENSIS Z m e .
4.
COMPLEXIOPOLLIS Zone.
New zone d e f i n e d by t h e appearance o f t h e Normapol-
l e s genus Complexiopollis s p . a t i t s b a s e .
I t i s repre-
s e n t e d a t s i t e 105 i n c o r e 9 , through t h e i n t e r v a l of samples DSDP-105-9-6, cm.
114-116 cm. through 105-9-1,
60-62
The t o p o f t h e zone i s unknown, as t h e cores immedi-
a t e l y above c o r e 9 a r e e s s e n t i a l l y devoid of sporomorphs. The COMPLEXIOPOLLIS Z m e r e f l e c t s t h e c o n t i n u i n g i n c r e a s e i n number of s p e c i e s and morphological d i v e r s i f i c a t i o n o f angiosperm p o l l e n i n t h e Middle Cretaceous of t t h e w e s t e r n North A t l a n t i c .
I n a d d i t i o n t o Complexiopol-
lis sp. a second genus of t h e Normapolles group, Atlantopollis s p . , a p p e a r s i n t h i s zone, as w e l l as p r o l a t e tric o l p o r a t e g r a i n s w i t h w e l l - d e f i n e d c i r c u l a r p o r e s (Tri-
colporites s p . ) and s p h e r i c a l t r i c o l p o r a t e g r a i n s oc-
360
curring in obligate tetrads (cf. D i c o t e t r a d i t e s sp.). CONCLUSIONS The individual zonations distinguished by dinoflagellates and sporomorphs respectively are valuable for relative age-dating in the western North Atlantic, as one zonation can complement the other and thereby afford greater precision.
Thus, dinoflagellates provide more precise
evidence for dating in the Neocomian, where four zones correspond to a single sporomorph zone (EPHEDRIPITES MULTICOSTATUS Zone).
On the other hand, in the late Hauter-
ivian-Cenomanian interval, both palynomorph groups appear to be of equal value f o r age-determination and correlation.
It is in this interval expecially that a multifos-
sil zonation incorporating the zonal taxa from both palynomorph groups might be useful.
To this end, a third
group of palynomorphs currently under investigation, small acritarchs of the M i c r y s t r i d i u m - V e r y h a c h i u m Group may contribute to such a zonation. Based on the biochronostratigraphic determinations proposed in this article the correlation of the palynomorph zonations against a geochronological scale is attempted (Figure 51, according to the scale proposed recently by van Hinte (1976). Such an attempt is useful for the interval represented by the BIORBIFERA JOHNEWINGIIDRUGGIDIUM RHABDORETICULATUM Zmes, as it is dated at site I
105 by the same nannofossil zonation (Thierstein, 1976)
which is presented in the geochronological scale (van Hinte, 1976).
Of equal value is the boundary between the
DEFLANDREA VESTITA and DEFLANDREA ECHINOIDEA zones, which is dated as Vraconian Albian on the basis of planktonic
M.Y. A.
DINOFL AGE L LAT E S
GEOCHRONOLOGY
SPOROMORPHS
- - - T R lTHY ROD1NlUM SUSPECTUM
CENOMANIAN
M
- - - -
COMPLEXIOPOLLIS
DEFLANDREA ECHINOIDEA ,
-
-
-
DEFLANDREA ALBIAN
VEST I TA
- -
M
SUBTlLl SPHAERA PERLUCIDA
- -
APTIAN E ~
PHOBEROCY STA
~~
NEOCOMICA
BARREMI AN
L I
HAUTERIVIAN
E1
VALANGINIAN
BERRIASIAN
1LAV A T I P 0 LL E NIT E$
E
- -DRUGTDIUMi HA B DORE T IC U LAT L - DRUGGEM- DEFLANDREI DRUGG IDIUM APICOPAUCICUM - -81ORB1-FERA -
E PHEDRlPlTES MULTlCO STAT US
- JOHNEWlNGll - .
Figlire 5. Pal;,,;:a-?cr;!~ z a n z t i o n s c o r r e l - ; t e d w i t h t h e g e o c h r o n o l o g i c a l s c a l e ( p a r t ) p u b l i s i i e d b y v a n H i n t c (137;).
W
B
362
foraminifers ( R o t a l i p o r a a p e n n i n i c a a p e n n i n i c a (Renz), Planomalina b u x t o r f i (Gandolfi), cf. Luterbacher, 1972, p. 561) at site 105, and which occur also in van Hinte's scale.
The ages of the remaining palynomorph zones are
not as well founded, as the evidence is provided by species of dinoflagellates, nannofossils, and ammonite aptychi which are not directly comparable to the geochronological scale. There is the potential for correlating the paleontologically dated sporomorph zonation in the western North Atlantic with the zonation established for the largely nonmarine coeval Cretaceous of the adjacent U.S. Atlantic Coastal Plain.
A zonation has been established in out-
crop and subsurface sections ranging from New Jersey to Virginia, based on sporomorph assemblages which indicate a range in age from Barremian (Zone I) to middle Turonian or younger (Zone V).
Although the sequence is largely
nonmarine, a major part of it has been compared with independently dated marine Aptian and Albian sporomorph assemblages of southern England (Kemp, 1970). The zonation has developed mainly through the comprehensive studies of Brenner (19631, Doyle (1969, 19731, Wolfe and Pakiser (19711, and Robbins, Perry, and Doyle (1975).
It is based to a large extent on the occurrence
of 'restricted' species, most of which are rare in the Coastal Plain (Brenner, 1963; Wolfe and Pakiser, 1971, p. 337) and therefore of limited value for correlation. These species occur in the North Atlantic as well, although they are equally rare.
However, the Coastal Plain
sequence contains the same succession of pollen morphotypes which occurs regularly in the sections studied from the North Atlantic.
Based mainly on the description of
363
the Coastal Plain stratigraphy summarized by Robbins, Perry, and Doyle (19751, a comparison is proposed for the North Atlantic zonation and Zones I-IV of the Atlantic Coastal Plain (Figure 6). Zone I, defined by Brenner (1963), can be correlated with the CLAVATIPOLLENITES Zone
in the western North Atlantic based on the occurrence of species of C l a v a t i p o l l e n i t e s .
The uppermost part of Zone
I and lower part of Zone I1 (subzones IIa and IIb) contain the first tricolpate and tricolporoidate species of reticulate dicotyledonous pollen, including Retitr(ico1p i t e s g e o r g e n s i s and T r i c o l p i t e s m i n u t u s , and can be cor-
related with the TRICOLPITES MINUTUS Subzone of this study.
Zones IIc and I11 (cf. Robbins, Perry, and Doyle,
1 9 7 5 ) contain R u g u b i v e s i c u l i t e s r u g o s u s and R . r e d u c t u s ,
as well as the first oblate triangular tricolporate grain grains, which suggests a correlation with the PSILATRICOLPORITES Subzone.
Zone IV is characterized by the first
Normapolles genera, specifically C o m p l e x i o p o l l i s and A t l a n t o p o l l i s , and prolate tricolporate and tetrad taxa,
all of which occur also for the first time in the COMPLEXIOPOLLIS Z x e . The EPHEDRIPITES MULTICOSTATUS Zone is not represented in the zonation published for the Coastal Plain. However, it is believed to represent the interval containing E p h e d r i p i t e s in the Bethards well (Robbins, Perry, and Doyle, 1 9 7 5 ) below the first specimens of C l a v a t i p o l lenites.
The close comparison of the two sporomorph flo-
ral successions permits the age-correlation of the North Atlantic sections with the Coastal Plain.
Zone I is
dated as late Hauterivian or Barremian to early Albian; IIa and IIb as middle to late Albian; IIc-I11 late Albian (including Vraconian) to Cenomanian; and Zone IV as Ceno-
364
C 0MPLEX I 0 POLLI!
vW
. I
I
t
PSILATRICOLPORI
-
I I C -ID
TRICOLPITES
--
MI NU TUS
C LAVAT IPOLLENI T E
I ----
EPHE DR IPITES
M ULTIC 0 STAT US
F i o u r e 6 . C o m p a r i s o n o f ldorth A t l a n t i c and a d j a c e n t U . S . A t l a n t i c Coastal P l a i n spororiorph z o n a t i o n s . Coastal P l a i n z o n a t i o n a f t c r Robbins, P e r r y and Doyle ( 1 9 7 5 ) .
365
manian. These age d e t e r m i n a t i o n s correspond r a t h e r c l o s e l y with t h a t i n d i c a t e d f o r t h e Coastal P l a i n zonation dated by t h e sporomorph f l o r a .
The c o r r e l a t i o n of t h e COM-
PLEXIOPOLLIS Zone w i t h Zone I V i s s t r e n g t h e n e d by t h e occ u r r e n c e of molluscs i n t h e marine Woodbridge Clay member of t h e lower p a r t o f t h e R a r i t a n Formation i n New J e r s e y ,
which d a t e s t h e upper p a r t of Zone I V as middle Cenomanian (Wolfe and P a k i s e r , 1971). ACKNOWLEDGMENTS
T h i s s t u d y w a s s u p p o r t e d by a g r a n t from t h e N a t i o n a l S c i e n c e Foundation, GA-39991. E.
I . Robbins reviewed t h e m a n u s c r i p t .
Discussion Dr. R. Aurisano: The high cingulum which is characteristic of Druggidium is rather unique. Do you have any ideas concerning the relationship of Druggidium to other dinoflagellates? Habib: Druggidium is similar in general morphology to Microdinium Cookson and Eisenack. However, it differs mainly by the position and type of archaepyle.
366
CITED REFEmNCES Brenner, G.J., 1963. The spores and pollen of the Potomac Group of Maryland: Marylanl. Dept. Geol., Mines, Water Res., Bull. 27, p. 1-215. Bukry, D., 1972. Coccolith stratigraphy, Leg XI, Deep Sea Drilling Project. In: Hollister, C.D., Ewing, J.I., and others, Initial Reports of the Deep Sea Drilling Project, Volume XI, p. 475-482. Couper, R.A., 1964. Spore-pollen correlation of the Cretaceous rocks of the northern and southern Hemispheres: SOC. Econ. Paleontol. Mineral., Spec. Publ. 11, p. 131142. Davey, R.J., and Verdier, J.P., 1971. An investigation of microplankton assemblages from the Albian of the Paris Basin: K. Nederl. W a d . Wetensch., Afd. Natuurk, Verh., v. 26, p. 5-58. , and I 1974. Dinoflagellate cysts from the Aptian type sections at Gargas and La Bedoule, France: Palaeontology, v. 17, pt. 3, p. 623-653. Doyle, J.A., 1969. Cretaceous angiosperm pollen of the Atlantic Coastal Plain and its evolutionary significance: Jour. Arnold Arboretum. v. 50, no. 1, p. 1-35. , 1973. The monocotyledons: their evolution and comparative biology. V. Fossil evidence on early evolution of the monocotyledons: Quart. Rev. Biol., v. 48, n. 3, p. 399-413. , Campo, M. Van, and Lugardon, B., 1975. Observations on exine structure of Eucommiidites and Lower Cretaceous angiospern pollen: Pollen et Spores, v. 17, n. 3, p. 429-486. Habib, D., 1976. Neocomian dinoflagellate zonation in the western North Atlantic: Micropaleontology, v. 21, n. 4, p. 373-392 (October, 1975). , in press. Palynostratigraphy of the Lower Cretaceous section at Deep Sea Drilling Project Site 391, Blake-Bahama Basin, and its correlation in the North Atlantic. Hinte, J.E. van, 1976. A Cretaceous time scale: Bull. Amer. Assoc. Petrol. Geol., v. 60, n. 4, p. 498-516. Kemp, E.M., 1970. Aptian and Albian miospores from southern England: Palaeontographica, v. 131, pt. B, p. 73-143. Lancelot, Y., Hathaway, J.C., and Hollister, C.D., 1972. Lithology of sediments from the western North Atlantic, Leg XI, Deep Sea Drilling Project, In: Hollister,C.D., Ewing, J.I., and others, Initial Reports of the Deep
367
Sea Drilling Project, Vol. XI, p. 659-666. Luterbacher, H.P., 1972. Foraminifera from the Lower Cretaceous and Upper Jurassic of the northwestern Atlantic. In: Hollister, C.D., Ewing, J.I., and others, Initial Reports of the Deep Sea Drilling Project, Volume XI, p. 561-593. Millioud, M.E., 1967. Palynological study of the type localities at Valangin and Hauterive: Rev. Palaeobot. Palynol., v. 5, p. 155-167. , 1969. Dinoflagellates and acritarchs from some western European Lower Cretaceous type localities: Internat. Conf. Planktonic Microfossils, First Geneva, 1967, Proc., v. 2, p. 420-434. Norris, G., 1969. Miospores from the Purbeck Beds and marine Upper Jurassic of southern England: Palaeontology, V. 12, pt. 4, p. 575-630. Renz, O., 1972. Aptychi (Ammonoidea) from the Upper Jurassic and Lower Cretaceous of the western North Atlantic (Site 105, Leg XI, Deep Sea Drilling Project). In: Hollister, C.D., Ewing, J.I., and others, Initial Reports of the Deep Sea Drilling Project, Vol. XI, P. 607-630. Robbins, E.I., Perry, W.J., and Doyle, J.A., 1975. Palynological and stratigraphic investigations of four deep wells in the Salisbury Embayment of the Atlantic Coastal Plain: U.S. Geol. Surv., Open File Report, No. 75-307, p. 1-120. Sverdlove, M . S . , and Habib, D., 1974. Stratigraphy and suggested phylogeny of D e f l a n d r e a v e s t i t a (Brideaux) comb. nov. and D e f l a n d r e a e c h i n o i d e a Cookson and Eisenack: Geoscience and Man, v. 9, p. 53-62. Thierstein, H.R., 1976. Calcareous nannoplankton biostratigraphy at the Jurassic-Cretaceous boundary: Bur. Rech. Geol. Min., Mem., v.86,~. 85-94. Verdier, J.P., 1975. Les Kystes de dinoflagellgs de la section de Wissant et leur distribution stratigraphique au Cre/tace/Moyen: Rev. Micropal6ontologieI v. 17, n. 4, p. 191-197. Wilcoxon, J.A., 1972. Upper Jurassic-Lower Cretaceous nannoplankton from the western North Atlantic Basin. In: ilollister, C.D., Ewing, J.I., and others, Initial Reports of the Deep Sea Drilling Project, Volume XI, P. 427-458. Wolfe, J.A., and Pakiser, H.M., 1971. Stratigraphic interpretations of some Cretaceous microfossil floras of the Middle Atlantic States: U.S. Geol. Surv., Prof. Paper 750-B, p. B35-B47.
This Page Intentionally Left Blank
369 Upper Cretaceous Dinoflagellate Zonation of the Subsurface Toms River Section Near Toms River, New Jersey Richard Aurisano and Daniel Habib
Rutgers U n i v e r s i t y , New Brunswick, N . J.;
Queens College, Flushing, New York
ABSTRACT
Thirteen peridinioid dinoflagellate species are used to zone the Upper Cretaceous subsurface section at the Tons River Chemical Well No. 84 site in the New Jersey Coastal Plain.
Seven of the thirteen species are con-
sidered to he restricted entirely to the Upper Cretaceous from their occurrence in the Toms River samples and also 2zrtially from their distribution in Upper Cretaceous sediments in North America, western Europe, the Soviet Union and Australia.
These are C h a t a n g i e l l a v n i g r i i , S u b t i l i s p h a e r a p o n t i s -
m a r i a e , C h a t a n g i e l l a manumii
, C.
new s p e c i e s
,
C . victoriensis, Deflan-
d r e a m i n o r , and D i c o n o d i n i o m f i r m m .
Six zones are distinguished in this section based on the stratigra>hic ranges of selected dinoflagellates.
In ascending stratigraphic order,
these ard the CHATANGIELLA VNIGYII zone, CHATANGIELLA MANUMII zone, CHATANGIELLA
NE\J SPECIES
zone, DICONODINIUY FIRh4UM zone, DEFLANDREA OEBISFEL-
DENSIS zone, and the DEFLANDREA BAKER11 z o n e .
One new species, C h a t a n g i e l l a
new s p e c i e s
,
is proposed.
Relative
percentage distribution data reveal an inverse relationship between Palaeoh y s t r i c h o p h o r a i n f u s o r i o i d e s and P . p a u c i s e t o s a .
The possibility is raised
that these species are morphotype variants of one species, dependent on some environmental factor ( s ) .
Ii4TRODUCTION
The purpose of this study is to propose a biostratigraphic zonation for Upper Cretaceous peridinioi?, dinofl3qellate cysts in the Xew Jersey Coastal Plain near Toms River.
The U?per Cretaceous section was recovered
370 from t h e Toms R i v e r Chemical Well N o . 84 (see Figure 1, Location Map). The samples from this s e c t i o n were made available by the U . S .
Geological
Survey.
41 I
11 00
40 I
I 0 00
39 0
19 00
I
15 00
10
0
I0
20
30
14 00
F i r j u r e 1 . S k e t c h map o f New J e r s e y s h o w i n ? a p p r o x i m a t e l ~ o s i t i o no f Toms R i v e r Chemical Well N o . 8 4 .
371 Investigations of Upper Cretaceous dinoflagellate cysts have been undertaken in Australia (Cookson, 1955, 1955; Cookson and Eisenack, 1958, 1960, 1961, 1962, 1968, and 1970), Europe (Alberti, 1959, 1961; Clarke and Verdier, 1967; Davey, 1970; Wilson, 1971), Africa (Davey, 19691, South America (Menendez, 1965), North America (Felix and Burbridge, 1973; Manum and Cookson, 1964; Harland, 1973; McIntyre, 1974; Drugg, 1967) and the USSR (Vozzhennikova, 1967).
These investigations show that many dinoflagellate
species have geographically wide ranging stratigraphic potential. The Toms River section is approximately 2,000 feet in thickness and ranges in age from Late Albian (Doyle, 1973
-
Pollen Zone IIC) to Tertiary.
The part of the section studied is that interval from subsurface depths of 1031 feet to 508 feet and ranges in age from Mid- to Upper Campanian to Tertiary.
The study was restricted to this interval because peridinioid
cysts are most abundant in the Upper Cretaceous and because little biostratigraphic work in this interval has been published.
The Lower and lower
Upper Cretaceous part of the Toms River section was zoned by Doyle (1973) utilizing pollen. Peridinoid dinoflagellate cysts, especially cavate species, were studied because of their abundance, ease of recognition and generally good preservation.
MATERIALS AND METHODS
Twenty samples of sediment, from the stratigraphic interval between 1031 feet and 508 feet depth, cored from the Toms River Chemical Well No. 84, were examined.
LITHOSTRATIGRAPHY
Based on data made available by W.J. Perry of the U.S. Geological Survey, the Tonis River section (TR1031 through TR508) consists of the f o l -
372 lowing lithostratigraphic units in ascending stratigraphic order: Woodbury Formation, Englishtown Formation, Marshalltown Formation, Wenonah Formation, Monmouth Group, Hornerstown Formation and the Vicentown Formation. These units have been identified on the basis of Gamma Ray and Self Potential Curves and lithological descriptions by the U . S .
Geological Survey.
BIOSTRATIGRAPHY
R.K. Olsson (pers. corn.) provided ages based on foraminifera1 data for the Toms River section studied.
The ages assigned are Campanian, Maes-
trichtian, and Tertiary (Figure 3).
The samples above the stratigraphic
level of sample TR599 (e.g., T'R591, T'R577, TR554, TR531, and TR508) are of Tertiary age.
The stratigraphic interval from level 1031 to somewhere be-
tween level 761 and 738 is Campanian.
The interval from somewhere between
levels 761 and 738 to level 599 is Maestrichtian. Dinoflagellate data suggest that the entire studied section (samples TR1031 to TR508) ranges in age from no older than Turonian to Paleocene. The Turonian lower limit is based upon the appearance of Chatangiella vnigrii (Lentin and Williams, 1975) in the lowest sample studied, TR1031. Caution is exercised in setting this age limit because C. vnigrii has been cite? in the literature only once at this writing (Vozzhennikova,1967), from the Turonian of Kazakhstan, USSR.
The geographic and time stratigra-
phl.c range of this species has not been firmly established.
Other species
occurrinq in sample TR1031 are Deflandrea echinoidea (Cookson and Eisenack, 1960), Palaeohystriciiopliora infusorioides (Deflandre, 1934) and P. paucisetosa
(Deflansre, 1943). Wi1liarc.s et a 1 (1974) reported the occurrence of
P. infusorioides in Early to Middle Albian material d t e d with foraminifera
from the Scotian Snelf of eastern Canada.
Deflandrea echinoidea has been
found in material as old as latest Albian (Vraconian) in the western North Atlantic (Sverdlove and Habib, 1974).
Palaeohystrichophora paucisetosa
373 however, has not been reported from material older t h m Coniacian (Sarjeant, 1967). Caution is exercised against assigninq a Coniacian lower age limit for the section because P. paucisetosa has not been frequently cited in the literature, and there is evidence from this study that P. infusorioides and
P. paucisetosa may be morphotype variants dependent upon some environmental factor(s). Doyle's work (1973) on angiosperm pollen types in the Atlantic Coastal Plain, including the lower section of the Toms River core (2000 feet to 1250 feet) places the Cenomanian/Turonian boundary at approximately 1250 feet, the approximate limit of Pollen Zone IV. The f:retaceous/Tertiary
boundary based upon dinoflagellates would be
placed somewhere between the levels of samples TR599 and TR623.
This is
reasonably close to the foraminifera1 boundary between samples TR591 and TR599.
The dinoflagellate species Deflandrea oebisfeldensis (Alberti, 1959)
makes its first appearance in the Upper Eocene of central Europe (Alberti, 1959), Paleocene through Lower Oligocene of the Volga region and the Eocene of the Baltic region (Vozzhennikova,1967).
In addition, the distinctive
outline of its periblast and archeopyle resembles those of other noted Tertiary species such as 2. phosphoritica (Eisenack, 1938) and D . andromedensis (Vozzhennikova, 1967).
Since dating with foraminifera is better established
and more reliable, the probability is greater that the Cretaceous/Tertiary boundary lies between samples TR599 and TR591.
This suggests the extension
of the stratigra.phic range of D. oebisfeldensis to Late Maestrichtian. A Campanian/Maestrichtian boundary is suggested between the levels of samples TR668 and TX646, based upon the stratigraphic disappearance of C h a tangiella manumii (Vozz:?ennikova, 1967) and Palaeohystrichophora infusorioides in sample TRG68, and the first appearance of Deflandrea diebelii (Alberti, 19591, in sample TR646.
C. manurnii has been found in Turonian to
Campanian material in Kazakhstan and Santonian material in western Siberia (Vozzhennikova, 1967).
Palaeohystrichophora infusorioides is found to
range from Early-Middle Albian to Campanian in material dated with foraminifera from the Scotian Shelf (Williams et a l , 19741, and the Grand Banks of New Foundland (Jenkins et a l , 1974). Deflandrea diebelii ranges from Lower Maestrichtian to lower Upper Maestrichtian In material dated with belemnites from Mons Klint, Denmark (Wilson, 1971), and is found in Danian stratotype material from Stevns Klint, Denmark (Wilson, 1971).
Jenkins et a 1
(1974), found D. diebelii to range from Maestrichtian to Paleocene in material dated with foraminifera from the Grand Banks, New Foundland.
Thus, it
appears that sample TR668 can be no younger than Campanian and sample TR646 no older than Maestrichtian.
Stage boundaries cannot be distinguished in
the interval between samples TR668 and TRld31, based on dinoflagellates. Dinoflagellate species found in this interval, which ranges in age from Campanian to no older than Turonian, are Deflandrea echinoidea, Chatangiel-
l a manumii, C. vnigrii, Subtilisphaera pontis-mariae, Chatangiella (new species) , Palaeohystrichophora infusorioides, P. paucisetosa, Chatanqiella victoriensis, and Diconodinium firmum. The published stratigraphic ranges of these species are as follows: Deflandrea echinoidea, latest Albian to Campanian (Davey, 1970; Harland, 1973; Sverdlove and Habib, 1974); Chatangiella manumii, Turonian to Campanian (Vozzhennikova,1967); Subtilisphaera pontis-mariae,Albian (Jenkins et
a l , 1974; Williams et a l , 1974) and Coniacian to Campanian (Sarjeant, 1967); Diconodinium firmum, Upper Campanian (Harland, 1973). Palaeohystrichophora infusorioides and P. paucisetosa The relative percentage distributions of selected peridinoid cysts, other cavate cysts, proximate and chorate cysts, were computed for each sample assemblage studied.
(See Table 2 ) .
The data show an inverse relationship between the re1at;ve percentage distributions of Palaeohystrichophora infusorioides and P. paucisetosa. For example, at the stratigraphic level of 1031 feet, the relative percen-
375
TABLE 2 Relative percenta7c d i s t r i b u t i o n o f ;elected o t h e r c a v a t e , p r o x i m a t e and c h o r a t e cyst;.
peridinioid cysts,
tage of P. infusorioides is 15.49 while for P . paucisetosa it is 24.34.
At
the level of 1009 feet, the relative percentage of P. infusorioides is 4.03 while for P. paucisetosa it is 55.24.
Thus, from level 1031 to level 1009,
the percentage distribution of P. infusorioides decreases while that of P. paucisetosa increases.
At the level of 907 feet, the relative percentage
of P. infusorioides is 56.20 while that of P . paucisetosa 13.57.
From le-
vel 1009 to level 987, P. infusorioides increases while P . paucisetosa decreases in relative percentage distribution.
This observation is made at
each level containing these species except 875, 852 and 761, 738 where the percentage of P . paucisetosa remains fairly constant (less than one percent change). It a,>?ears that when moving from one stratigraphic level to another, an increase in relative percentage distribution of one species is followed by a corresponding decrease in the other. P.
The inverse relationship between
paucisetosa and P. infusorioides apparently shows no corresponding re-
lationship to lit-hologicalchanges.
376 P. infusorioides and P. paucise'msa have similar morphologies.
They
are distinguished on the basis of periblast ornaLnentation, average size, and endoblast.
The periblast of P. paucisetosa has hair-like processes
(setae) which are longer than those of P. infusorioides and more sparsely distributed.
In addition, the periblast of P. paucisetosa, on the average,
is smaller than that of P. infusorioides.
The endoblast of P. paucisetosa
is rarely if ever appressed against the periblast. The question is raised: Are P. infusorioides and P. paucisetosa two separate species which have different specific environmental needs or are they morphotype variants of the same species, i.e., a species population whose average characteristics vary with respect to some environmental factor(s) such as salinity, temperature, trace elements, etc. This is an important consideration because P. infusorioides and P. paucisetosa have different stratigraphic ranges.
If P. infusorioides and
P. paucisetosa are morphotype variants of one species, any zonation utili-
zing the ranges of both species is invalid.
Further investigation into
this issue is warranted and until more information is available, it would be safer to consider the combined stratigraphic ranges of P. infusorioides and P. paucisetosa when making a zonation.
DINOFLAGELLATE ZONATION
Six zones are distinguished from the Upper Cretaceous/Tertiary section at Chemical Well No. 84 near Toms River, New Jersey, based on the stratigraphic ranges of selected peridinioid cysts.
In ascending strati-
graphic order, these are the CHATANGIELLA VNIGRII zone, CHATANGIELLA MANUMI1 zone, CHATANGIELLA
NEW SPECIES
zone, DICONODINIUM FIRYUM zone, DE-
FLANDREA OEBISFELDENSIS zone, and the DEFLANDREA BAKER.11 zone. Of the thirteen species whose ranges are shown in Figure 3, seven are considered to be restricted entirely to the Upper Cretaceous, from their
occurrence in the Toms River (TR) samples and also partially from their distribution in Upper Cretaceous sediments in North America, western Europe, and the Soviet Union.
These are Chatangiella vnigrii, Subtili-
sphaera pontis-mariae, Chatangiella manumii, C. ensis, Diconodinium firmum, and Deflandrea minor.
new species, C. victoriIn addition, one species
is considered to be restricted to the Lower Tertiary.
This species is De-
flandrea bakerii . CIIATANGIELLA VNIGRII ZONE Concurrent range zone defined by the stratigraphic interval from the lowest occurrence of the nominative species up to but not including the lowest occurrence of Chatangiella manumii. Other cavate cysts which appear in the CHATANGIELLA VNIGRII zone include Palaeohystrichophora infusorioides, P. paucisetosa, Deflandrea echinoidea, and Subtilisphaera pontis-mariae.
The most conspicuous feature of
this zone is the absence of C. manumii. The CHANTANGIELLA VNIGRII zone is the lowest zone in the Toms River samples studied.
Its lower limit is not known since samples below 1031
feet were not investigated.
Since its upper limit is approximately 829
feet, the CHATANGIELLA VNIGRII zone is at least 202 feet thick.
The zone
is considered to range in age from upper Lower Campanian to lower Upper Campanian on the basis of foraminifera1 data.
An age range of no older
than Turonian to Campanian is suggested by the published ranges of C. vnigrii and C. manumii. CHATANGIELLA MANUMII ZONE Concurrent range zone defined by the stratigraphic interval from the lowest occurrence of the nominative species up to but not including the lowest occurrence of Chatangiella new species Other cavate cysts which appear in the CHATANGIELLA IMANUMII zone include Palaeohystrichophora infusorioides, P. paucisetosa, and Deflandrea
378 echinoidea. The CHATANGIELLA MANUMII zone ranges in age from lower Upper Campanian to Uppermost Campanian or Lowermost Maestrichtian on the basis of foraminiferal data.
Dinoflagellates suggest an age range of no older than Turonian
to Campanian.
It occurs at the Toms River site from approximately 829 feet
to 738 feet and includes samples TR829, TR806, and TR761.
NEW SPECIES
CHATANGIELLA
ZONE
Concurrent range zone defined by the stratigraphic interval from the lowest occurrence of the nominative species up to but not including the lowest occurrence of Diconodinium firmum. Palaeohystrichophora infusorioides, P. paucisetosa, Deflandrea echinoidea, and Chatanqiella manumii appear stratigraphically within the CHATANGIELLA
NEW SPECIES
zone.
In addition, C. victoriensis and Deflandrea
minor make their first appearance in this zone. The CHATANGIELLA
NEW SPECIES
zone ranges in age from Upper Campa-
nian or Lower Maestrichtian to Maestrichtian on the basis of foraminifera1 data.
Dinoflagellates suggest a Campanian to Maestrichtian age.
It is
found at the Toms River site from approximately 738 feet to 668 feet and is approximately 70 feet thick.
Samples TR738 and TR715 are included in this
zone. DICONODINIUM FIRMUM ZONE Concurrent range zone defined by the stratigraphic interval from the lowest occurrence of the nominative species up to but not including the lowest occurrence of Deflandrea oebisfeldensis. Cavate cysts which appear stratigraphically in the DICCNODINIUM FIRMUM zone include Deflandrea echinoidea, Chatangiella victoriensis, and Deflandrea minor.
D. diebelii makes its first appearance within this zone, a
species whose published range is Maestrichtian to Danian (Alberti, 1959; Wilson, 1971).
The DICONODINIUM FIRMUM zone ranges in age from lower Upper Maestrichtian to upper Upper Maestrichtian on the basis of foraminifera1 data.
Di-
noflagellates suggest an age range of Campanian or Maestrichtian to Maestrichtian or Paleocene.
It is found at the Toms River site from approxi-
mately 668 feet to 599 feet and has a thickness approximating 69 feet. Samples TR668, TR646, and TR623 are included in this zone. DEFLANDREA OEBISFELDENSIS ZONE Concurrent range zone defined by the stratigraphic interval from the lowest occurrence of the nominative species up to but not including the lowest occurrence of Deflandrea bakerii. Deflandrea diebelii disappears stratigraphically within the DEFLANDREA OEBISFELDENSIS zone.
D. bakerii makes its first appearance within this
zone. The DEFLANDREA OEBISFELDENSIS zone has a lower age limit of Upper Maestrichtian to Paleocene based on foraminiferal data.
Dinoflagellate
data disagrees and places the lower age limit in the Paleocene.
The upper
boundary of this zone lies within the Early Tertiary. DEFLANDREA BAKERII ZONE Concurrent range zone defined by the stratigraphic interval from the lowest occurrence of the nominative species.
Because TOTE River samples
above 508 feet were not studied, the upper limit of this zone has not been determined. D. oebisfeldensis occurs in the DEFLANDREA BAKERII zone. The DEFLANDREA BAKERII zone is restricted to the Tertiary in age.
The
upper boundary of the zone is not known.
A COMPARISON OF LITHOSTRATIGRAPHY KITH BIOSTRATIGRAPHY
It is difficult to compare biostratigraphic boundaries with lithostratigraphic boundaries because the lithostratigraphic units are based on the
380 results of a contlnuous gamma log, whereas the biostratigraphic zones are based upon the stratigraphic ranges of dinoflagellates as represented by 20 samples (See Figure 2).
508 531 554 571 599 623 646 668
MONMOUTH GROUP
[,"",.""
,,.,"
FIRMUM ZONE
^..
715 738 761 806 829 852 875
1CHATANCIELLA
1
...__-----
964 987 009 031
WOOOBURV FORMATION
-
VNlGRll ZONE
Figure 2. Chart comparing dinoflagellate zonation with corresponding lithofacies.
Based upon the ages discussed in the beginning of the biostratigraphy section, the Tertiary/Cretaceous boundary lies somewhere at the base of the Hornerstown Formation and the Maestrichtian/Campanian boundary, based on foraminifera1 data, lies somewhere at the base of the Wenonah or top of the Marshalltown.
Dinoflagellate data place this boundary somewhere in the
middle of the Monmouth Group.
The rest of the section ranges in age from
no older than Turonian to Campanian based on selected dinoflagellates and Campanian, based on foraminifera.
ACKNOWLEDGMENTS
T h i s paper w a s made p o s s i b l e through t h e generous g i f t of t h e B. P. Alaska E x p l o r a t i o n Company i n s u p p o r t of R. A u r i s a n o ' s t h e s i s r e s e a r c h and by a g r a n t from t h e N a t i o n a l Science Foundation ( NSF a t Queens C o l l e g e .
-
GA
-
39991 )
im FORAMINIFERA OINOFLAGELLATE
%-
ZONATION
ZONATION
SPECIES LIST
531
55
I
TERTIARY 2
3 4 5 8 1
I J I1 11
I2
I3
FIGURE 3
382 REFERENCES Alberti, G., 1959. Zur Kenntnis der Gattung Deflandrea Eisenack (Dinoflag.) in der Kreide und im Alttertikr Nord--Mitte Deutschiands. Mitt. Geol . Stdatsinst, v. 28, p. 93-105. Clarke, R.F.A. and Verdier, J.P., 1967. An investigation of microplankton assemblages from the chalk of the Isle of Wight, England. vorh. K. Ned. Akad. Net., v. 24, n. 3, 96p. Cookson, I . C . , 1956. Additional microplankton from Australian Late Mesozoic and Tertiary sediments. Austral. J. Mar. Freshw. Res., v. 7, p. 183-191. and Eisenack, A., 1958. Microplankton from Australian and New Guinea upper Mesozoic sediments. ?roc. Roy. SOC. Victoria, v. 70, p . 19-79. , 1960. Microplankton from Australian Cretaceous sediments. Micropaleontology, v. 6, n. 1, p. 1-18. , 1961. Upper Cretaceous microplankton from the Belfast No. 4 Bore, southwestern Victoria. Proc. Roy. SOC. Victoria, V. 74, p. 69-76. , 1962. Additional microplankton from Australian Cretaceous sediments. Micropaleontology, v. 8, n. 4, p. 495-507. , 1968. Microplankton from two samples from Gengin Brook No. 4, Borehole, Western Australia. J. Roy. SOC. W. Austral., v. 51, p. 110-122. , 1970. Cretaceous microplankton from the Eucla Basin, Western Australia. ?roc. Roy. SOC. Victoria, v. 83, p. 137-157. and Manum, s., 1964. On Deflandrea victoriensis n. sp., D.. tripartita Cookson & Eisenack, and related species. ?roc. Roy. SOC. Victoria, v. 77, p. 521-524. Davey, R.J., 1970. Non-calcareous microplankton from the Cenomanian of England, northern France, and North America, part 11. Bull. Br. Mus. Nat. Hist. (Geol.) v. 18, n. 8, pp. 335-397. , 1969. Some dinoflagellate cysts from the Upper Cretaceous of northern Natal, South Africa. Paleontol. Afr., V. 12, pp. 1-23. Deflandre, G., 1934. Sur les microfossiles d'origine planctonique conserve's a ' I'ktat de mati'ere organique dans les silex de la craie. C.R. Acad. Sci. Paris, 199, 966-968. , 1935. Conside/rations biologiques sur les microorganismes d'origine planctonique conserve's dans les silex de la craie. Bull. Biol. Fr. Belg., v. 69, p. 213-244. , 1936. Microfossiles des silex cr&tace/s. Premikre partie. Generalities. Flagellgss; Ann. Pale'ont., v. 25, p. 151-191. , 1940. Microfossiles de quelques silex de la craie blanche de Vendame. Bull. SOC. Hist. Nat. Toulouse, 75, 155-159. _. , 1943. Sur quelques nouveaux Dinoflagellgs des silex crdtace's. Bull. SOC. Ggol. Fr., V. 13, p. 499-509. , 1966. Addendum 'a mon Me'moire: Microfossiles des silex cr6tac6s. Cah. Micropalebntol. (Arch. orig. Cen. Docum. C:N.R.S. no. 419) ser. 1, no. 2, p. 1-9. and Cookson, I.C., 1955. Fossil microplankton from Australian Late Mesozoic and Tertiary sediments. Austral. J. Mar. Freshw. Res., v. 6 , p. 242-313. and Courtville, H., 1939. Note prgliminaire sur les microfossiles des silex crGtac6s du Cambresis. Bull. SOC. Fr. Microsc. 8, 2 ~ 95-106. . et al, 1970. Re-issue of Deflandre and Cookson, 1955 (in French), by Laboratoire de Micropalebntologie de L'Ecole Practique des
383 Hautes Etudes. Institute de Palgontologie du Museum, Palls. avec addendum et post pace, pp. 1-70, 1-54. Douglas, J.G., 1960. Microplankton of the Deflandreidae group in western district sediments. Min. Geol. J., 6(4), pp. 17-32. Downie, C. and Sarjeant, W.A.S., 1966. The morphology, terminologi and classification of dinoflagellate cysts, in Davey, R.J., Downie, C., Sarjeant, W.A.S., and Williams, G.L., Studies on Mesozoic and Cain. ioA, dinoflagellate cysts: Bull. of the British Museum (Nat. Hist.) Geol. supplement 3, p. 10-17. , Evitt, W.R., and Sarjeant, W.A.S., 1963. Dinoflagellates, hystrichospheres and the classification of the Acritarchs.' Stanford Univ. Publ., Geol. Sciences, 7, 1-16. Doyle, J.A., 1973. The monocotyledons: their evolution and comparative biology, Part V. Fossil evidence on early evolution of the monocotyledons. Quart. Review of Biology, v. 48, n. 3, p. 399-413. Drugg, W.S., 1967. Palynology of the Upper Moreno Formation (Late Cretaceous-Paleocene) Escarpado Canyon, California. Palaeontographica, Abt. B, V. 120, p. 1-71. Eisenack, A., 1938. Die Phosphoritknollen der Bernsteinformation als uberlieferer Tertiaren Planktons. Sehr. Phys. b'kon. Ges., v. 70, p.181-188. and Cookson, I.C., 1960. Microplankton from Australian Lower Cretaceous sediments. Proc. Roy. SOC. Victoria, v . 72, p. 1-11. Evitt, W.R., 1961. Observations on the morphology of fossil dinoflagellates. Micropaleontology, v. 7, n. 4 , pp. 385-420. 1967. Dinofilagellate studies 11. The archeopyle. Stanford Univ. Publ., Geol. Sci., v. 10,n. 3, 84p. 1969. Dinoflagellates and other organisms in palynological preparation. Aspects of Palynology, R.H. Tschudy and R.A. Scott (Eds.) Wiley-Interscience, pp. 439-479. 1970. Dinoflagellates. A selective review. Geoscience and Man. v. 1,p. 29-45. and Davidson, S.E., 1964. Dinoflagellate studies I. Dinoflafellate cysts and thecae. Stanford Univ. Publ., Geol. Sci., v. x, n. 1, p. 3-12. and Wall, D., 1968. Dinoflagellate studies IV. 'Theca and cyst of recent freshwater P e r i d i n i u m l i m b a t i u m (Stokes) Lemmermann. Stanford Univ. Publ., Geol. Sci., v. xii, n. 2 , 15p. Felix, C.J. and Burbridge, P.P., 1973. A Maestrichtian age microflora from Arctic Canada. Geoscience and Man, v. vii, p. 1-29. Habib, D., 1972. Dinoflagellate stratigraphy, Leg XI, Deep Sea Drilling Project, in Hollister, C. Ewing, J. Hathaway, J., Paulus, F., Lancelot, Y., Habib, D., Poag, C.W., Luterbacher, H.P., Worstell, P. and Wilcoxon, J.A.. Initial Reports of the Deep Sea Drilling Project, v. X I , Washington ( U . S . Government Printing Office), xxii to 1077p. and Warren, J.S., 1973. Dinoflagellates near the Cretaceous/ Jurassic boundary. Nature, v. 241, n. 5386, p. 217-218. Harland, R., 1973. Dinoflagellate cysts and acritarchs from the Bearpaw Formation (Upper Campanian) of southern Alberta, Canada. Palaeontology, v. 16, part 4 , pp. 665-706. Jenkins, W.A.M. et a l , 1974. Stratigraphy of the Amoco 10E 4-1 puffin B-90 Well, Grand Banks of Newfoundland. Can. Geol. Survey Paper, pp. 74-61. Koch, R. and Olsson, R.K., 1974. Abt. Microfossil biostratigraphy of the uppermost Cretaceous beds of New Jersey. In Abstracts with Programs, v. 5, n. 1, GSA, p. 45. Lejeune - Carpentier, 1942. L'gtude microscopique des silex. Pgridiniens nouveaux ou peu connus. (Dixi'eme note). Ann. SOC. Ggol. Belg. v. 65, p. B181-Bl92. Lentin, J . K . and Williams, G.L., 1975. A monograph of fossil peridinioid
384 dinoflagellate cysts. Bedford Inst. of Oceanography,Rept.Ser.BI-R-75-16. Malloy, 1972. An Upper Cretaceous dinoflagellate cyst lineage from Gabon, West Africa. Geoscience and Man, v. 4, p. 57-65. Manum, S., 1963. Some new species of D e f l a n d r e a and their probable affinity with Peridinium. Arbok Norst Polar Inst. 1962, p. 55-67. and Cookson, I.C., 1964. Cretaceous microplankton in a sample from Graham Island, Arctic, Canada, collected during the second "Fram expedition" (1898-1902). Norske Vid - Akad., Skrifter, I, Mat.-Naturv., 17, 1-36. McIntyre, D.J., 1975. Morphologic Changes in D e f l a n d r e a from a Campanian section, District of Mackenzie, N.W.T., Canada. Geoscience and Man, v. xi, pp. 61-76. Menendez, 1965. Microplancton fosil de sedimentos Terciarios y cretacecos del nort de Tierra del Fuego (Argentina). Ameghiniana, v. 9 , p . 7-15. Minard, J.P. e t a l , 1974. Preliminary report on geology along Atlantic continental margin of northeastern United States. AAPG Bull., v. 58, n. 6, part I1 of 11, p. 1169-1178. Owens, J.P. and Sohl, N.G., 1969. Shelf and deltaic paleoenvironmects in the Cretaceous-Tertiary formations of the New Jersey Coastal Plain. Geology of Selected Areas in New Jersey and Eastern Pennsylvania and Guidebook of Excursions, Rutgers Univ. Press, pp. 235-278. , 1973. Glauconites from New Jersey - Maryland Coastal Plain: Their K-Ar ages and applications in stratigraphic studies. GSA Bull., v. 84, p. 2811-2838. Petters, Sunday, 1976. Upper Cretaceous subsurface stratigraphy of Atlantic Coastal Plain of New Jersey. AAPG Bull., v. 60, n. 1, p. 87-107. Sarjeant, W.A.S., 1967. The stratigraphic distribution of fossil dinoflagellates. Rev. Palaeobotan. Palynol., 1, pp. 323-343. , 1966. Further dinoflagellate cysts from the Speeton Clay: Studies on Mesozoic and Cainozoic dinoflagellate cysts. R.J. Davey e t a l , (eds.), Brit. Mus. (Nat. Hist.) Bull., Geol. Supp. 3, p. 199-214. Snead, X.G., 1969. Microfloral diagnosis of the Cretaceous - Tertiary boundary, central Alberta. Research Council Alberta, Bull. 25, 148p. Stover, L.E., 1973. Paleocene an:?. Eocene species of D e f l a n d r e a (Dinophyceae) in Victorian coastal ar3 offshore basins, Australia. Spec. Publ. Geol. SOC. Aust., 4: pp. 167-188. Sverdlove, M. and Habib, D., 1974. Stratigraphy and suggested phylogeny of D e f l a n d r e a v e s t i t a (Brideaux) comb. nov. and D e f l a n d r e a e c h i n o i d e a Cookson and Eisenack. Geoscience and Man, v. 9, p. 53-62. Upshaw, C.F., 1964. Palynological zonation of the Upper Cretaceous Frontier Formation near Dubois, Wyoming: Palynology in oil\exploration,A. T. Cross (ed.), SEPM Spec. Publ., n. 11, p. 153-168. Valensi, L., 1955. Etude micropalgontologique des silex du Magdalenien de Saint-Amand (Cher). Bull. Soc. Prghist. Fr., 52, p. 584-596. Vozzhennikova, T.F., 1965. Introduction to the study of fossil peridinean algae. Akad. Nauk. SSSR, Sib. Otd., Inst. Geol. Geofiz., ER., 156p. , 1967. Fossil peridinians of the Jurassic, Cretaceous and Paleogene deposits of the USSR. Akad. Nauk. SSSR, Sib. Otd., Inst. Geol. Geofiz., Tr., 347p. Wall, D., 1971. Biological problems concerning fossilizable dinoflagellates. Geoscience and Man, v. 3, p. 1-15. and Dale, B., 1967. The resting cysts of moderi! marine dinoflagellates and their paleontological significance. Rev. Paleobotan. Palynol., 2, pp. 349-354. , 1968. Modern dinoflagellate cysts and evolution of the Peridiniales. Micropaleontology, v. 14, n. 3, p. 265-304. ,1973. Paleosalinity relationships of dinoflagellates
385 on t h e L a t e Q u a t e r n a r y o f t h e Bl.ack Sea - A summary. Geoscience and Man, V . 7 , p . 95-102. Williams, G . L . and L e n t i n , J.K., 1973. F o s s i l d i n o f l a g e l l a t e s : i n d e x t o g e n e r a and s p e c i e s . Geol. S u r . Can., Dept. of Energy, Mines and Res o u r c e s , Paper 73-42, 176p. et al, 1 9 7 4 . S t a t i g r a p h y o f t h e S h e l l Naskapi N . 30 Well, S c o t i a n S h e l f , e a s t e r n Canada. Canadian Geol. Survey Paper 74-50,p.1-12. Wilson, G . J . , 1 9 7 1 . O b s e r v a t i o n s on European L a t e C r e t a c e o u s d i n o f l a g e l l a t e c y s t s . P r o c e e d i n g s o f t h e I1 P l a n k t o n i c C o n f e r e n c e , R o m a 1970. , 1967. Microplankton from t h e Garden Cove Formation, Campbell I s l a n d , N e w Zealand, J . B o t . , v . 5 , p . 223-240.
EXPLANATION OF PLATE 1 1. C h a t a n g i e l l a s p . , approx. 630x. Sample TR668, s l i d e no. RA-1, 7 . 7 ~105.5. V e n t r a l view. Note t h e b r o a d l y a r c h e d e p i t r a c t , two-walled e n d o b l a s t , and d i s t i n c t i v e cingulum. 2 . C h a t a n g i e l l a s p . . Same specimen a s i n f i g . 1 f o c u s i n g on t h e e n d o b l a s t . 3 . C h a t a n g i e l l a s p . . Same specimen a s i n f i g . 1 i n d o r s a l view. Note t h e p a r t i t e cingulum and p r e c i n g u l a r and p o s t c i n g u l a r p l a t e s . 4 . C h a t a n g i e l l a manumii ( Vozzhennikova ) L e n t i n and W i l l i a m s , 1975. Approx. 740x. Sample TR829, s l i d e n o . RA-14, 0 . 7 x 96.3. V e n t r a l view. Note weakly arched e p i t r a c t , cingulum o u t l i n e d by c o n i c a l p r o j e c t i o n s , and c o n i c a l p r o j e c t i o n s s c a t t e r e d about t h e p e r i b l a s t . 5. C h a t a n g i e l l a v i c t o r i e n s i s (Cookson and Manum) L e n t i n and W i l l i a m s , 1975. Approx. 630x. Sample TR623, s l i d e no. RA-15, 3 0 . 5 x 106.8. 6 . C h a t a n g i e l l a v n i g r i i (Vozzhennikova) L e n t i n and W i l l i a m s , 1975. Approx. 630x. Sample TR1031, s l i d e no. RA1 0 , 8.9 x 97.5. V e n t r a l view. Note d i s t i n c t i v e l y arched e p i t r a c t and c o n i c a l a p i c a l h o r n . A l s o , n o t e s u l c u s and c o n i c a l p r o j e c t i o n s on p e r i b l a s t . 7 . D e f l a n d r e a b a k e r i i D e f l a n d r e and Cookson. Approx. 630x. Sample TR554, s l i d e no. RA-9, 0 . 4 x 1 0 3 . 5 . Note s u l c u s and weakly i n d i c a t e d cingulum. Also, note the evenly scabrate ornamentation. 8 . Deflandrea echinoidea Cookson and E i s e n a c k . Approx. 500x. Sample TR1009, s l i d e no. RA-7, 6.4 x 105.3. 9. D e f l a n d r e a minor A l b e r t i . Approx. 630x. Sample TR646, s l i d e no. RA-21, 9 . 3 x 84.2. EXPLANATION OF PLATE 2
1. D e f l a n d r e a o e b i s f e l d e n s i s A l b e r t i . Approx. 400x. Sample TR591, s l i d e no. RA-8, 9.5 x 91.0. Note t h e l a r g e t r a p e z o i d a l a r c h e o p y l e . 2 . Diconodinium firmum H a r l a n d . Approx. 1OOOx. Sample TR668, s l i d e no. RA-6, 1 . 8 x 116.1. mariae Note b i f u r c a t i n g a p i c a l p r o c e s s . 3 . S u b t i l i s p h a e r a p o n t i s ( D e f l a n d r e ) L e n t i n and W i l l i a m s , 1975. Approx. 860x. Sample TR852, s l i d e no. RA-9, 0.4 x 8 7 . 5 . 4 . P a l a e o h y s t r i c h o p h o r a e a u c i s e t o s a D e f l a n d r e . Approx. 1 2 0 0 ~ .Sample TR829, s l i d e no. RA-14, 3 . 1 x 11C.O. 5. 2 p m c i s e t o s a . Approx. 1 2 0 0 ~ .Sample TR1031, s l i d e no. RA-8, 1 1 . 5 x 9 5 . 0 . 6 . P a l a e o h y s t r i chophora i n f u s o r i o i d e s D e f l a n d r e . Approx. 860x. Sample TR806, s l i d e no. RA-1, 6.0 x 8 2 . 1 . V e n t r a l view. Note s u l c u s . 7 . 2. i n f u s o r i o i d e s . Approx. 860x. Same specimen a s i n F i g u r e 1 w i t h f o c u s on e n d o b l a s t . 8. p. infuso r i o i d e s . Approx. 860x. Sane specimen a s i n F i g u r e 1 i n d o r s a l view. Note c o n t i n u o u s cingulum. A l s o , c o n t r a s t r e l a t i v e l e n g t h of s e t a w i t h t h o s e i n F i g u r e s 3 and 4 of p. p a u c i s e t o s a . 9. p. i n f u s o r i o i d e s . Approx. 860x. Same specimen a s i n F i g u r e 1 w i t h f o c u s on e n d o b l a s t i n d o r s a l view.
-
386
PLATE 1
387
PLATE 2
This Page Intentionally Left Blank
389
NORTH ATLANTIC CENOZOIC FORAMINIFERA by W. A. Berggren Department of Geology and Geophysics, Woods Hole Oceanographic Institution Woods Hole, Massachusetts 02543 and Department of Geological Sciences Brown University, Providence, Rhode Island 02912 Abstract Cenozoic foraminiferal biostratigraphy, biogeography and ecology in the North Atlantic and its marginal borderland areas is reviewed. Planktonic foraminiferal faunal diversity and provincialization exhibit fluctuating patterns which indicate a complex historical relationship between changing paleogeography, paleooceanography and paleoclimatology. Late Neogene climatic deterioration is reflected in increased planktonic foraminiferal provincialization and the development of separate zonations for high and low latitudes as well as a reduction in amphi-Atlantic cosmopolitanism among shallow water benthonic Foraminifera. In a similar manner stratigraphic and geographic distribution patterns among benthonic Foraminifera have been dependent upon the changing geometry of the Atlantic plate. The apparently slow rate of morphologic evolution in most shallow and deep-water benthonic foraminiferal phyletic lineages restricts their usefulness in fine-scaled biostratigraphic studies (the larger foraminifera are an obvious exception), but there has been a renewed interest in using them in paleobiogeographic, paleo-oceanographic and paleobathymetric studies. The evolution of the modern bathyal-abyssal benthonic foraminiferal fauna about 15 Ma allows paleobathymetric and paleocirculation interpretations by analogy with living faunas valid over most, if not all, of the Neogene, and in a general way for the Paleogene as well. RQsumd I1 s'agit de la biostratigraphie c6nozoique foraminifgrale, la bioggographie et l'ecologie dans l'iltlantique du Nord et les cbtes qui l'entourent , La diversite' faunale planctonique foraminifgrale et la provincialisation montrent des motifs fluctuants qui indiquent un rapport historique complique' parmi une pal&oge'ographie, une pal6oocdanographie et une palgoclimatologie changeantes. La dgterioration climatique de la Late Nkogkne est r6fle'chie dans la provincialidation foraminife'rale planctonique augment& et le dgveloppement des zonations sgpare'es des hautes et basses latitudes, ainsi qu'une r6duction dans le cosmopolitanisme amphi-Atlantique parmi les foraminifkres benthoniques des hauts-fonds. D'une faFon semblable, les motifs de distribution stratigraphique et ggographique parmi les foraminifhres benthoniques dgpendent de la g6ometrie changeante de la plate de 1'Atlantique. L'allure Lvidement lente de l'dvolution morphologique dans la plupart des lignages phylktiques foraminiferales benthoniques se trouvant dans les hauts-fonds et les eaux profondes limite leur utilite dans des 4tudes biostratigraphiques sur une petite e'chelle (les foraminifhres qui sont p l u s grands dvidemment font l'exception), mais il y a un renouvellement de l'inte'r8t dans leur utilisation dans des dtudes pale'obiog6ographiques,paldoocdanographiques et paldobathyme'triques. L'gvolution de la faune bathyal-abysaal-benthonique foraminife'rale moderne d'environ 15 Ma permet a des interpr6tation.s paldobathymetriques et palLocirculations par analogie avec des faunes vivantes. C e s interprktations sont valables pour la plupart, sinon toute, le Niog'ene et dans un sens gCngra1, Ie PalLoghe.
1;yTRODUCT I O N
C e n o z o i c p l a n k t o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y may b e s a i d t o h a v e matured, i f n o t been b o r n , i n t h e C a r i b b e a n and Gulf C o a s t . >fonographic i n v e s t i g a t i o n s i n t h e 1950's, p r i m a r i l y by o i l company m i c r o p a l e o n t o l o g i s t s , o f t h e P a l e o g e n e a n d Keogene o f t h e C a r i b b e a n - G u l f C o a s t r e g i o n d e m o n s t r a t ed t h e u t i l i t y o f planktonic Foraminifera i n l o c a l and r e g i o n a l s t r a t i graphic s t u d i e s t h e r e and around t h e margins of t h e North A t l a n t i c Basin. The a d v e n t o f t h e Deep Sea D r i l l i n g P r o j e c t i n t h e m i d - 1 9 6 0 ' s p r o v i d e d t h e m a t e r i a l a n d i m p e t u s f o r d e t a i l e d i n v e s t i g a t i o n s on t h e g e o g r a p h i c and 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 of t a x a a n d t h e f o r m u l a t i o n o f b i o s t r a t i g r a p h i c z o n a t i o n s o f i n c r e a s i n g r e s o l u t i o n . About 40-50 C e n o z o i c p l a n k t o n i c f o r a m i n i f e r a l zones have b e e n r e c o g n i z e d i n t r o p i c a l r e g i o n s . C o r r e l a t i o n o f t h e s e z o n e s w i t h t h e p a l e o m a g n e t i c ( t = 25-0 Y a ) , m a g n e t i c a n o m a l y ( t = 65-0 ?fa) a n d r a d i o m e t r i c ( t = 65-0 Ma) t i m e - s c a l e s i s p r o v i d i n g a g e o c h r o n o l o g i c framework i n which t h e Cenozoic e v o l u t i o n o f t h e A t l a n t i c Ocean c a n b e q u a n t i f i e d . A summary o f r e c e n t a d v a n c e s i n C e a o z o i c p l a n k t o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y a n d b i o g e o g r a p h y o f t h e A t l a n t i c Ocean h a s b e e n r e c e n t l y c o r n p l c t e d ( B e r g g r e n , in p r e s s ) , s o t h a t l e s s e m p h a s i s i s d e v o t e d t o t h i s p,:-oup t h a n t h e b e n t h o n i c F o r a m i n i f e r a i n t h i s summary. N o r t h A t l a n t i c Cenozoic p l a n k t o n i c f o r a m i n i f e r a l d a t a are c o n t a i n e d i n t h e P r e l i m i n a r y I i e p o r t s o f t h e Deep Sea D r i l l i n g P r o j e c t ( v o l s . 1-4, 1 0 - 1 5 ) a n d w i l l b e c o n t a i n e d i n t h e f o r t h c o m i n g v o l u m e s 3 7 , 3 8 , 44, a n d 45). S t u d i e s on C e n o z o i c ( e x c l u s i v e o f H o l o c e n e a n d l i v i n g ) d e e p - s e a b e n t h o n i c F o r a m i n i f e r a i n t h e North A t l a n t i c are s t i l l i n t h e i r i n f a n c y . P r i o r t o 1 9 7 0 t h e r e w e r e v i r t u a l l y no p r e - P l e i s t o c e n e d e e p sea c o r e s a v a i l a b l e . Between 1968-1975 t h e Glomar C h a l l e n g e r h a s o b t a i n e d n u m e r o u s c o r e s f r o m s e v e r a l areas o f t h e N o r t h A t l a n t i c ( l e g s 1, 2 , 11, 1 2 , 1 4 , 3 8 , 45 a n d t h e Gulf C o a s t l e g 10) a n d a g e n e r a l p i c t u r e o f t h e d i s t r i b u t i o n OF deep-sea Cerlozoic f a u n a s i s b e g i n n i n g t o e m e r g e . The u s e o f b e n t h o n i c F o r a m i n i f e r a i n b i o s t r a t i g r a p h y h a s b e e n hampe r e d b y l a c k o f c a r e f u l c o m p a r a t i v e taxonomic s t u d i e s which r e p r e s e n t t h e b a s i c d a t a b a s e f o r b i o s t r a t i g r a p h i c and b i o g e o g r a p h i c i n t e r p r e t a t i o n s . The a p p a r e n t l y s l o w r a t e s o f m o r p h o l o g i c e v o l u t i o n i n m o s t s h a l l o w a n d d e e p w a t e r p h v l e t i c l i n e a g e s has r e s t r i c t e d t h e i r usefulness i n fine-scaled b i o s t r a t i g r a p h i c s t u d i e s ( t h e l a r g e r F o r a m i n i f e r a are a n obvious exception), b u t t h e r e h a s b e e n a renewed i n t e r e s t , t r a c e a b l e t o t h e i r i n c r e a s e d r e c o v e r y f r o m t h e d e e p - s e a f l o o r b y t h e Deep S e a D r i l l i n g P r o j e c t , i n u s i n g them i n paleobiogeographic, paleooceanographic and paleobathymetric s t u d i e s . T h i s u s e f u l n e s s w i l l d e p e n d , i n t u r n , upon 1) t h e a b i l i t y t o e x t r a p o l a t e p r e s e n t d i s t r i b u t i o n p a t t e r n s i n t o t h e p a s t a n d 2 ) r e l i a b l e d a t a on (pa1eo)biogeography of benthonic Foraminifera. I t h a s b e e n found t h a t 1) a b o v e i s a p p l i c a b l e o v e r a p p r o x i m a t e l y t h e l a s t 15 my; b e y o n d t h a t i t i s p o s s i o l e t o e x t r i c a t e o n e s e l f f r o m t h e h o r n s o f t h e dilemma b y m a k i n g g e o p h y s i c a l l y - b a s e d s e a - f l o o r t o p o g r a p h i c r e c o n s t r u c t i o n s which t h e n s e r v e as a b a s i s f o r e v a l u a t i n g p a l e o b a t h y m e t r i c d i s t r i b u t i o n p a t t e r n s o f p r e l a t e Neogene t a x a . In t h e c a s e o f 2 ) a b o v e , p r e s e n t l y a v a i l a b l e d a t a s t r o n g l y s u g g e s t t h a t t h e b i o g e o g r a p h i c d i s t r i b u t i o n of b e n t h o n i c foramini f e r a i s s t r o n g l y l i n k e d w i t h p a s t o c e a n - c o n t i n e n t geometry and a t t e n d a n t p a l e o c l i m a t i c and p a l e o c i r c u l a t i o n h i s t o r y (Berggren and H o l l i s t e r , 1 9 7 4 ) . T h i s i s p a r t i c u l a r l y t r u e f o r t h e North A t l a n t i c Ocean and i t s circummarginal a r e a s . The d i s c u s s i o n o f b e n t h o n i c F o r a m i n i f e r a b e l o w r e p r e s e n t s a s y n t h e s i s o f a l a r g e amount o f l i t e r a t u r e i n a d d i t i o n t o c u r r e n t work b e i n g c o n d u c t e d a t o u r own l a b o r a t o r y . Most o f t h e b a s i c l i t e r a t u r e on A t l a n t i c - C a r i b b e a n C e n o z o i c b e n t h o n i c f o r a m i n i f e r a l f a u n a s up t o 1 9 6 9 WDS c i t e d i n B e r g g r e n
391 and P h i l l i p s ( 1 9 7 1 ) . The i n t e r e s t e d r e a d e r i s r e f e r r e d t o t h i s s o u r c e f o r details. Only t h e m o r e p e r t i n e n t r e f e r e n c e s c i t e d t h e r e a s w e l l as t h o s e from t h e i n t e r v a l s i n c e 1 9 7 0 w i l l b e s p e c i f i c a l l y c i t e d h e r e . A d i s c u s s i o n o f t h e t e c t o n i c - s t r a t i g r a p h i c setting o f t h e N o r t h A t l a n t i c ( i n c l u d i n g t h e N o r w e g i a n - G r e e n l a n d S e a ) , a n d t h e m a r g i n a l areas i n N o r t h America a n d E u r o p e p r o v i d e s t h e f r a m e w o r k f o r a summary o f t h e m a i n d i s t r i b u t i o n p a t t e r n s of f o r a m i n i f e r a 1 faunas i n t h e s e t h r e e a r e a s .
TECTONIC-STRATIGRAPHIC SETTING North A t l a n t i c The t e c t o n i c - s t r a t i g r a p h i c e v o l u t i o n o f t h e N o r t h A t l a n t i c Ocean h a s b e e n r e v i e w e d i n terms o f s e a - f l o o r s p r e a d i n g a n d p l a t e t e c t o n i c s by B e r g g r e n a n d P h i l l i p s ( 1 9 7 1 ) , P i t m a n a n d T a l w a n i ( 1 9 7 2 ) , Emery a n d Uchupi ( 1 9 7 2 ) , B e r g g r e n a n d H o l l i s t e r ( 1 9 7 4 ) , L a u g h t o n ( 1 9 7 5 ) , a n d i n Woodland ( e d . , 1 9 7 5 ) , among o t h e r s . The m a i n e v e n t s i n t h e C e n o z o i c e v o l u t i o n o f t h e N o r t h A t l a n t i c may b e e n u m e r a t e d a s f o l l o w s :
1)
I n t h e P a l e o c e n e a c h a n g e f r o m a two t o t h r e e p l a t e g e o m e t r y r e s u l t e d from a s h i f t i n a c t i v e s p r e a d i n g from t h e L a b r a d o r Sea (which c o n t i n u e d o p e n i n g u n t i l t h e Middle Eocene) t o t h e R e y k j a n e s Ridge which r e s u l t e d i n t h e s e p a r a t i o n of G r e e n l a n d a n d E u r a s i a a n d t h e o p e n i n g o f t h e N o r w e g i a n S e a . A t t h i s t i m e R o c k a l l Bank (DSDP S i t e 1 1 6 , 1 1 7 ) b e g a n s u b s i d i n g t o i t s p r e s e n t d e p t h (1200 m). C o n c o m i t a n t l y , b u t c a u s a l l y u n r e l a t e d , O r p h a n Knoll (DSDF S i t e 111) s u b s i d e d t o i t s p r e s e n t d e p t h ( 1 8 0 0 m) d u r i n g t h e l a t e s t C r e t a c e o u s - P a l e o c e n e t i m e . T h e Mesozoic t a p h r o g e n i c s t a g e of r i f t i n g o f t h e North Sea changed t o a n i n t r a c r a t o n i c d r i f t i n g s t a g e i n t h e E a r l y Cenozoic (mid-Paleocene). D u r i n g t h e l a s t 6 0 m . y . t h e N o r t h Sea B a s i n h a s b e e n c h a r a c t e r i z e d by r e g i o n a l s u b s i d e n c e a n d t h e f o r m a t i o n o f a n i n t r a c r a t o n i c b a s i n i n w h i c h t e r r i g e n o u s s e d i m e n t s up t o 3 . 5 km i n t h i c k n e s s h a v e b e e n deposited.
2)
I n t h e Middle Eocene s p r e a d i n g i n t h e L a b r a d o r S e a c e a s e d and G r e e n l a n d became p a r t of t h e N o r t h American p l a t e . The P y r e n e a n o r o g e n y r e s u l t e d i n t h e u p l i f t o f a c e n t r a l r i d g e i n t h e f l o o r o f t h e Bay o f B i s c a y (DSDP S i t e s 118 a n d 1 1 9 ) .
3)
During t h e E a r l y t o Middle Miocene I c e l a n d a p p e a r s t o have formed a s a s u b a e r i a l f e a t u r e , p e r h a p s r e l a t e d t o a m a n t l e plume. N o r t~ h A m e_ rica _ -
The e a s t e r n p a r t o f N o r t h A m e r i c a i s a c o a s t a l g e o s y n c l i n e c o n t a i n i n g g r e a t t h i c k n e s s e s o f ? l a t e P a l e o z o i c , Mesozoic and C e n o z o i c s e d i m e n t s which s l o p e g e n t l y away ( s e a w a r d ) f r o m t h e c e n t r a l s t a b l e ( c r a t o n i c ) p o r t i o n o f t h e c o n t i n e n t (Murray, 1 9 6 1 ) . Cenozoic s e d i m e n t s a r e r e l a t i v e l y u n d i s t u r k d a n d u n d e r l i e a l o w - r e l i e f c o a s t a l p l a i n w h i c h e x t e n d s some 6 0 0 0 km f r o m n o r t h o f Newfoundland t o Honduras i n C e n t r a l America. The c o m b i n e d a v e r a g e w i d t h o f t h e emerged a n d submerged p o r t i o n of t h e p r o v i n c e i n t h e A t l a n t i c r e g i o n i s 2 5 0 km; a r o u n d t h e G u l f o f Plexico t h e a v e r a g e w i d t h d e c r e a s e s f r o m 600 km i n t h e U n i t e d S t a t e s t o 1 6 0 km i n Mexico ( T a m p i c o ) . To t h e s o u t h i t c o n t i n u e s to, n a r r o w a n d t h e n e x p a n d s t o n e a r l y 900 km i n w i d t h (N-S) t h r o u g h E l P i t e n ( G u a t e m a l a ) a n d Y u c a t a n P e n i n s u l a . The s e a w a r d b o u n d a r y o f t h e c o a s t a l p r o v i n c e i s p l a c e d a t t h e seaward-
392 f a c i n g s c a r p o r s l o p e o f t h e c o n t i n e n t a l s l o p e a n d r i s e (2. &.; see a l s o Hedberg, 1 97 6 ) . The c l a s s i c p i c t u r e t h a t t h e r e g i o n a l M e s o z o i c a n d C e n o z o i c s t r u c t u r a l e x p r e s s i o n of t h e c o a s t a l p r o v i n c e h a s b e e n c o n t r o l l e d b y g r a v i t y d o m i n a t e d d e f o r m a t i o n a n d t h a t s e d i m e n t a r y a l i g n m e n t s are a c c o r d a n t w i t h p r e s e n t day r e g i o n a l s t r u c t u r e ( M u r r a y , 1 9 6 1 ) h a s r e c e n t l y b e e n q u e s t i o n e d (Brown 2.: z?., 1 9 7 2 ) . In t h e v i e w o f t h e l a t t e r p r i n c i p a l v e r t i c a l m o b i l i t y h a s b e e n c a u s e d by b l o c k f a u l t i n g o r f l e x i n g , a c c o m p a n i e d by r o t a t i o n a l a l i g n m e n t of t h e a x e s o f p o s i t i v e a n d n e g a t i v e r e g i o n a l s t r u c t u r e s , and o c c u r r i n g c o n c o m i t a n t l y w i t h d i f f e r e n t i a l r a t e s of r e l a t i v e s u b s i d e n c e f o r c o a s t a l s e g m e n t s t h a t a r e j u x t a p o s e d a l o n g e l e m e n t s of a n i n t e r s e c t i n g h i n g e b e l t s y s t e m , c o m p o n e n t s o f w h i c h h a v e o n e of t h r e e p r i n c i p a l a l i g n m e n t s - NE-SW, 1 J W S E o r N - S . F i r s t and second o r d e r s t r a t i graphic u n i t s a r e then delineated. Cenozoic s e d i m e n t s of t h e Gulf a n d A t l a n t i c C o a s t a l p r o v i n c e s a r e p a r t o f a l a r g e g e o s y n c l i n a l s e d i m e n t a r y c o m p l e x of c o n t i n e n t a l , e s t u a r i n e , d e l t a i c a n d m a r i n e d e p o s i t s . The i n t e g r a l r e l a t i o n s h i p b e t w e e n b i o - a n d l i t h o f a c i e s h a s p e r h a p s nowhere b e e n b e t t e r d e m o n s t r a t e d t h a n i n t h i s complex r e g i o n where t h e c y c l i c a l , s e q u e n t i a l r e p l a c e m e n t o f one l i t h o t o p e b y a n o t h e r , h a s b e e n d e l i n e a t e d o v e r two g e n e r a t i o n s b y m i c r o p a l e o n t o l o g i s t s engaged i n commercial ( p e t r o l e u m ) e x p l o r a t i o n and S t a t e and U . S . G e o l o g i c a l Survey e x p l o r a t i o n s . S o r e 12-15,000 m e t e r s o f !lesozoic-Cenozoic s e d i m e n t s h a v e b e e n e s t i m a t e d t o o c c u r i n t h e Gulf C o a s t a n d 6 0 0 0 m e t e r s f o r t h e A t l a n t i c C o a s t ( r o u g h l y a 2 : l r a t i o ) b y ! l u r r a y ( 1 9 6 1 ) , b u t r e c e n t e s t i m a t e s b a s e d on s e a ward e x t r a u o l a t i o n s of o n s h o r e t h i c k e n i n g t r e n d s f o r s t r a t i g r a p h i c u n i t s i n S o r t h C a r o l i n a w e l l s s u g g e s t s t h a t t h i c k n e s s e s i n t h e two a r e a s may b e c o m p a r a b l e , on t h e o r d e r of 12-15 km (Brown 2' c:., 1 9 7 2 ) . In t h e A t l a n t i c C o a s t a l P l a i n C e n o z o i c s e q u e n c e c a r b o n a t e s p r e d o m i n a t e i n t h e s o u t h e r n h a l f , w h e r e a s m a r l s and c o n t i n e n t a l c l a s t i c s predominate i n t h e n o r t h e r n h a l f ( X a h e r , 1 9 7 1 ) . O u t c r o p s o f m a r i n e P a l e o g e n e a n d Neogene h a v e b e e n d r e d g e d i n s u b m a r i n e c a n y o n s a l o n g t h e e a s t c o a s t a n d on t h e Subsurface stratiB l a k e P l a t e a u - Bahama Banks r e g i o n a n d G e o r g e s Bank. g r a p h i c c o r r e l a t i o n o f t h e A t l a n t i c C o a s t a l P l a i n h a s r e c e n t l v b e e n compiled by Maher ( 1 9 7 1 ) , b a s e d on a n i n v e s t i g a t i o n o f numerous s t a t e s u r v e y w a t e r wells a n d t h e J O I D E S P h a s e I w e l l s (51-56) o f f t h e F l o r i d a C o a s t . E
u
~
g
The C e n o z o i c t e c t o n i c e v o l u t i o n o f w e s t e r n E u r o p e h a s b e e n r e v i e w e d b y R u t t e n ( 1 9 6 9 ) a n d i t s ( b i 0 ) s t r a t i g r a p h i c e v o l u t i o n b v Brinkmann ( 1 9 6 0 ) a n d P o m e r o l (1972) on w h i c h w h o s e p u b l i c a t i o n s t h e f o l l o w i n g b r i e f summary is based. Because t h e p r e s e n t day c o a s t o f w e s t e r n Europe was e v o l v e d d u r i n g t h e Cenozoic, marine sediments a r e l i m i t e d l a r g e l y t o t h e v i c i n i t y of p r e s e n t c o a s t a l r e g i o n s , e x c e p t i n t h e A l p i n e r e g i o n (ex T e t h y s ) . Paleogene s e d i m e n t s c r o p o u t i n v a r i o u s b a s i n s e x t e n d i n g some 1500 km f r o m S c a n i a ( s o u t h e r n Sweden) t o t h e A q u i t a i n e B a s i n (SW F r a n c e ) a n d NW S p a i n , w h e r e a s Xeogene s e d i m e n t s o c c u r i n s c a t t e r e d s e q u e n c e s f r o m Denmark t o SW S p a i n . C e n t r a l E u r o p e may b e d i v i d e d i n t o two p r i m a r y z o n e s o f s u b s i d e n c e , a n €-I4 B a l t i c a n d a N-S R h e n i s h , a s t r u c t u r a l a l i g n m e n t w h i c h d e v e l o p e d i n the L a t e Permian. The N o r t h S e a a n d A n g l o - P a r i s - B e l g i a n b a s i n s w e r e i n e x i s t e n c e s i n c e t h e e a r l y M e s o z o i c ( J u r a s s i c ) . The l a t t e r f o r m e d a c o n t i n u o u s , i n t e r c o n n e c t e d zone of s u b s i d e n c e on t h e m a r g i n o f t h e F e n n o - S c a n d i a n s h i e l d t o t h e n o r t h a n d s e p a r a t e d f r o m t h e S o r t h German B a s i n by t h e A r d e n n e s - R h e n i s h M a s s i f a n d t h e Upper R h i n e R i d g e .
393 In P a l e o c e n e t i m e t h e f o r m e r l y u n i f i e d A n g l o - G a l l i c B a s i n w a s b r o k e n up i n t o s e v e r a l u n i t s - t h e London, H a m p s h i r e , P a r i s a n d B e l g i a n b a s i n s a n d become, p a l e o g e o g r a p h i c a l l y , a p a r t o f t h e same m a r i n e r e g i o n a s n o r t h e r n Germany. The S o r t h Sea B a s i n i s t h e m o s t p r o m i n e n t f e a t u r e i n t h e s e d i m e n t a r y h i s t o r y of w e s t e r n Europe n o r t h of t h e A l p i n e g e o s y n c l i n e . During i t s s e d i m e n t a r y h i s t o r y t r a n s v e r s e r i d g e s d i v i d e d i t i n t o s m a l l e r b a s i n s w h i c h were i n t e r m i t t e n t l y l i n k e d w i t h o n e a n o t h e r d u r i n g t h e Mesozoic and Cenozoic. S i g n i f i c a n t s u b s i d e n c e b e g a n i n t h e P a l e o c e n e i n t h e Nortll Sea Basin a s a r e s u l t o f t e n s i o n a l f o r c e s r e l a t e d t o t h e opening of t h e Xorwegian S e a . D u r i n g t h e Cenozoic t h e d e p o s i t i o n a l c e n t e r of t h e North Sea B a s i n was g r a d u a l l y d i s p l a c e d i n a n o r t h w a r d o r n o r t h w e s t e r l y d i r e c t i o n by t h e g r a d u a l r e t r e a t o f t h e sea f r o m t h e m a r g i n a l a r e a s o f n o r t h w e s t e r n E u r o p e . The g e n e r a l r e g r e s s i o n o f t h e s e a a n d d i f f e r e n t i a l t e c t o n i c s i n t h e d i f f e r e n t b a s i n s r e s u l t e d i n t h e termination of marine deposition i n t h e Anglo-Paris Basin a t t h e end of t h e Paleogene. The S o r t h S e a B a s i n i s t h e o n l y one i n which s e d i m e n t a t i o n h a s c o n t i n u e d t o t h e p r e s e n t t i m e . Linked i n t h e p a l e o g e o g r a p h i c s e n s e d u r i n g t h e P a l e o c e n e w i t h t h e North Sea B a s i n , t h e A n g l o - P a r i s - B e l g i a n b a s i n s h a v e b e e n e s s e n t i a l l y dormant f o r t h e p a s t 25 m i l l i o n y e a r s , a l t h o u g h p e r i o d i c s u b s i d e n c e d u r i n g t h e Neogene has r e s u l t e d i n s p o r a d i c , discontinuous sedimentary sequences i n these b a s i n s , p a r t i c u l a r l y d u r i n g t h e L a t e Miocene, P l i o c e n e and P l e i s t o c e n e . T h i c k n e s s e s o f C e n o z o i c s e d i m e n t a r y r o c k s r a n g e f r o m a b o u t 250 m i n t h e P a r i s B a s i n t o 800 m i n t h e H a m p s h i r e B a s i n t o o v e r 3000 m i n - t h e c e n t r a l p a r t of t h e North Sea. In SW E u r o p e m a r i n e C e n o z o i c s t r a t a o c c u r i n t h e A q u i t a i n e B a s i n o f SW F r a n c e , i n t h e t e c t o n i c a l l y a c t i v e c o a s t a l r e g i o n o f SW S p a i n a n d i n t h e Late Neogene G u a d a l q u i v i r B a s i n of SW S p a i n . S t r a t i g r a p h y was b o r n i n E u r o p e w i t h t h e i n v e s t i g a t i o n s of W i l l i a m Smith i n t h e e a r l y 1 9 t h c e n t u r y . The c l a s s i c s u b d i v i s i o n o f C e n o z o i c In p a r t i c u l a r , t h e m a r i n e s e d i m e n t s was o r i g i n a l l y d o n e i n w - s t e r n E u r o p e . c l a s s i c s u b d i v i s i o n o f t h e P a l e o g e n e i s b a s e d on m a r g i n a l m a r i n e s e q u e n c e s d e v e l o p e d i n t h e b a s i n s of NW E u r o p e ( L o n d o n , H a m p s h i r e , P a r i s , B e l g i a n , N o r t h G e r m a n ) , w h e r e i n d i v i d u a l s t a g e u n i t s a r e g e n e r a l l y i d e n t i f i e d with m a r i n e t r a n s g r e s s i o n s and c o n c o m i t a n t c h a n g e s i n b i o t a . Deeper w a t e r P a l e o g e n e s e d i m e n t s a r e e x p o s e d i n t h e A q u i t a i n e B a s i n (SW F r a n c e ) a n d a l o n g t h e Yh’ c o a s t o f S p a i n . Neogene d e p o s i t s a r e r a r e a n d p o o r l y e x p o s e d i n t h e m a r g i n a l a r e a s o f w z s t e r n E u r o p e e x c e p t i n t h e E-W s t r i k i n g l i n e a r r e e n t r a n t o f t h e A t l a n t i c Ucean i n t h e G u a d a l q u i v i r B a s i n - i n SW S p a i n . A s a r e s u l t , m o s t o f t h e Neogene s t r a t o t y p e s e c t i o n s a r e l o c a t e d i n t h e more compl et e s e q u e n c e s d e v e l o p e d i n t h e X e d i t e r r a n e a n r e g i o n . For a r e l a t i v e l y u p - t o - d a t e summary o f T e r t i a r y s t r a t i g r a p h y a n d r e l a t e d b o u n d a r y problems t h e i n t e r e s t e d r e a d e r i s r e f e r r e d t o aerggren ( 1 9 7 1 ) . PALEOGENE C e n o z o i c b i o g e o g r a p h y o f t h e N o r t h A t l a n t i c and m a r g i n a l ( i . e . a d j a c e n t ) l a n d areas i s i n t i m a t e l y r e l a t e d t o t h e c o n t i n u i n g p r o c e s s o f t h e fragmentation of L a u r a s i a . The o p e n i n g o f t h e X o r w e g i a n S e a d u r i n g t h e P a l e o g e n e a n d a t t e n d a n t s t r u c t u r a l r e o r g a n i z a t i o n o f t h e NE A t l a n t i c l e a d i n g t o t h e e v e n t u a l e s t a b l i s h m e n t o f an A r c t i c P r o v i n c e w i t h d i r e c t d e e p w a t e r connection t o t h e S o r t h A t l a n t i c r e s u l t e d i n d i s t i n c t f a u n a l changes a s a The o p e n i n g r e s u l t of t h e s e m o d i f i c a t i o n s t o l a n d and sea c o n f i g u r a t i o n s . o f t h e N o r t h A t l a n t i c t o t h e A r c t i c r e s u l t e d i n t h e f o r m a t i o n of North A t l a n t i c Deep Water, c o o l i n g o f s u r f a c e w a t e r s i n t h e N o r t h A t l a n t i c , enhanced l a t i t u d i n a l p r o v i n c i a l i z a t i o n of planktonic f o r a m i n i f e r a 1 faunas, s h a l l o w i n g o f t h e CCD ( a n d p r o b a b l e c o n c o m i t a n t c h a n g e s i n d i s t r i b u t i o n p a t t e r n s o f b a t h y a l b e n t h o n i c f o r a m i n i f e r a 1 f a u n a s ) and w i d e s p r e a d
394 d e v e l o p m e n t o f s h a l l o w , w a r m w a t e r . ' ~ , ~ : 4 ~ v ~ fLaiczieess d u r i n g t h e E o c e n e . S u p e r i m p o s e d upon t h e s e c h a n g e s w e r e t h o s e c a u s e d b y a g r a d u a l l o w e r i n g of t e m p e r a t u r e d u r i n g t h e Cenozoic and t h e g r a d u a l southward d i s p l a c e m e n t o f p a l e o l a t i t u d e s as t h e c r u s t moved n o r t h w a r d s o v e r t h e mantle. T h e r e h a s b e e n a n a p p r o x i m a t e l y 15"-20" s o u t h w a r d d i s p l a c e m e n t o f l a t i t u d e s i n t h e n o r t h e r n hemisphere d u r i n g t h e Cenozoic (Berggren and P h i l l i p s , 1971). One o f t h e m a j o r e v e n t s w h i c h a f f e c t e d P a l e o g e n e f o r a m i n i f e r a l e v o l u t i o n a n d b i o g e o g r a p h y w a s t h e s u d d e n a n d s i g n i f i c a n t 5°C c o o l i n g o f b o t t o m w a t e r s ( f r o m c?. 1 0 ° - 5 " C ) i n t h e S o u t h e r n Ocean ( K e n n e t t a n d S h a c k l e t o n , 1 9 7 6 ) n e a r t h e E o c e n e / O l i g o c e n e b o u n d a r y , c%. 38 ? l a , r e s u l t i n g i n t h e f o r m a t i o n o f t h e p s y c h r o s p h e r e ( s e e a l s o S a v i n e t aZ., 1 9 7 5 ) .
Planktonic Foraminifera The G u l f a n d A t l a n t i c C o a s t a l P l a i n P a l e o g e n e s u c c e s s i o n h a s b e e n p l a c e d w i t h i n a p l a n k t o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h i c framework by t h e s t u d i e s o f L o e b l i c h a n d T a p p a n ( 1 9 5 7 ) , O l s s o n ( 1 9 6 0 , 1 9 7 0 a , b ) , Eames eZ 9L. ( 1 9 6 2 ) , Blow ( 1 9 6 9 ) , a n d B a r k e r a n d Blow ( 1 9 7 6 ) , among o t h e r s . Similar s t u d i e s h a v e b e e n c o n d u c t e d on P a l e o g e n e s e d i m e n t s o f w e s t e r n E u r o p e b u t t h e f a u n a s t e n d t o b e p o o r e r owing t o t h e g e n e r a l l y s h a l l o w e r - w a t e r envirorrment o f d e p o s i t i o n . Paleogene p l a n k t o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y and biogeography of t h e N o r t h A t l a n t i c h a s r e c e n t l y b e e n r e v i e w e d b y B e r g g r e n ( 1 9 7 2 , i n p r e s s ) a n d B e r g g r e n a n d H o l l i s t e r ( 1 9 7 4 ) s o t h a t o n l y a b r i e f summary i s presented here. E a r l y P a l e o c e n e (Danian) f a u n a s e x h i b i t morphologic c o n s e r v a t i s m , low d i v e r s i t y a n d g e o g r a p h i c c o s m o p o l i t a n i s m . D i s t i n c t d i v e r s i f i c a t i o n and p r o v i n c i a l i z a t i o n o c c u r r e d by m i d - P a l e o c e n e time a n d b y t h e e n d o f t h e Paleocene t h r e e d i s t i n c t , l a t i t u d i n a l l y - d i s t r i b u t e d assemblages a r e recognizable. S u b b o t i n i d s and globular-chambered a c a r i n i n i d s c h a r a c t e r i z e t h e high l a t i t u d e assemblages; middle l a t i t u d e s are c h a r a c t e r i z e d by s u b b o t i n i d s , a c a r i n i n i d s a n d m o r o z o v e l l i d s , w h e r e a s low l a t i t u d e s a r e c h a r a c t e r i z e d by a c a r i n i n i d s a n d m o r o z o v e l l i d s . T e m p e r a t u r e - r e l a t e d f l u c t u a t i o n s i n f a u n a l a s s o c i a t i o n s a r e seer. t h r o u g h o u t t h e r e m a i n d e r o f t h e Paleogene. The m o s t n o t i c e a b l e c h a n g e o c c u r s a t t h e e n d o f t h e E o c e n e a n d d u r i n g t h e E a r l y O l i g o c e n e when a m a r k e d f a u n a l r e p l a c e m e n t o c c u r r e d a c c o m p a n i e d b y l o w e r e d s p e c i e s d i v e r s i t y a n d t h e d e v e l o p m e n t o f two g r o u p s of m o r p h o l o g i c a l l y c o n s e r v a t i v e g l o b i g e r i n i d s , t h e "small" and " l a r g e " g l o b i g e r i n i d s , w h i c h c h a r a c t e r i z e d h i g h a n d low l a t i t u d e s , r e s p e c t i v e l y . G l o b i g e r i n i t i d s ( i n i z a : a , d i s s i r ; i i < s ) a n d g l o b o q u a d r i n i d s a r e common e l e m e n t s i n Upper O l i g o c e n e s e d i m e n t s i n t h e N o r t h A t l a n t i c . B e c a u s e o f t h e s p a r s e n e s s a n d / o r i r r e g u l a r d i s t r i b u t i o n of t h e f a u n a , a s t r i c t b i o s t r a t i g r a p h i c z o n a t i o n was n o t f o u n d p o s s i b l e i n t h e h i g h l a t i t u d e N o r t h A t l a n t i c P a l e o g e n e . More u s e f u l was a m u l t i p l e z o n a t i o n b a s e d on t h e a s s o c i a t i o n o f v a r i o u s f a u n a l e l e m e n t s . Only a n a p p r o x i m a t e c o r r e l a t i o n with the t r o p i c a l A t l a n t i c zonation i s possible. P a l e o g e n e p l a n k t o n i c f o r a m i n i f e r a l a s s e m b l a g e s i n t h e N o r t h Sea a r e c h a r a c t e r i z e d by a c a r i n i n i d s - s u b b o t i n i d s i n t h e Paleocene-Eocene a n d by In s m a l l g l o b i g e r i n i d s and non-keeled g l o b o r o t a l i i d s i n t h e O l i g o c e n e . t h e Norwegian-Greenland S e a , p l a n k t o n i c F o r a m i n i f e r a a r e r e l a t i v e l y s p a r s e d u r i n g t h e P a l e o g e n e ; Eocene f a u n a s c o n s i s t a l m o s t s o l e l y o f s j b b (S. sa:agcnica, S. :ir.ase:pta) a n d a f e w , s m a l l a c a r i n i n i d s .
395 Benthonic Foraminifera P a l e o g e n e s m a l l e r b e n t h o n i c f o r a m i n i f e r a l f a u n a s e x h i b i t a marked c o s m o p o l i t a n i s m w h i c h may b e e x p l a i n e d a s a r e s u l t o f w i d e s p r e a d e q u i t a b l e c l i m a t i c c o n d i t i o n s a n d low p o l a r - e q u a t o r i a l t h e r m a l g r a d i e n t s . I n t h e P a l e o c e n e t h r e e main d e p t h - c o n t r o l l e d b e n t h o n i c f o r a m i n i f e r a l a continental s h e l f fauna, assemblages ( i . e . f a c i e s ) have been recognized: t e r m e d t h e "Midway-type f a u n a " (PIF) , a c o n t i n e n t a l s l o p e a n d a b y s s a l p l a i n f a u n a , t h e " V e l a s c o - t y p e f a u n a " (VF), a n d a s h a l l o w , warm-water c a r b o n a t e s h e l f f a u n a , t h e " T e t h y a n c a r b o n a t e f a u n a " (TCF) ( B e r g g r e n a n d A u b e r t , 1 9 7 5 ) . The Midway f a u n a a c h i e v e d e s s e n t i a l l y w o r l d w i d e d i s t r i b u t i o n a n d o c c u r s i n A t l a n t i c P a l e o c e n e s e d i m e n t s a s f a r n o r t h a s 70"N L a t . ( E a s t e r n
l i n i d s ) , polymorphinids and t e x t u l a r i i d s . The " T e t h y a n c a r b o n a t e f a u n a " (TCF) i s c h a r a c t e r i z e d by d i s t i n c t
a-.5c&szr/n,
and
7.1;
of b o t h a s s e m b l a g e s o c c u r i n t h e C a r i b b e a n and M e d i t e r r a n e a n ( T e t h y a n ) regions. Elements of assemblage b) occur i n Paleocene s t r a t a of western Europe ( N e t h e r l a n d s ) . P a l e o g e n e s e d i m e n t a t i o n on t h e m a r g i n o f t h e N o r t h A t l a n t i c o c c u r r e d i n a s e r i e s of c y c l e s of t r a n s g r e s s i o n s and r e g r e s s i o n s . T h e s e a r e nowhere b e t t e r demonstrated t h a n i n t h e s t u d i e s on t h e b e n t h o n i c f o r a m i n i f e r a l f a u n a s o f t h e A n g l o - P a r i s B a s i n s ( W r i g h t a n d M u r r a y , 1 9 7 2 ; Murray a n d W r i g h t , 1 9 7 4 ) . These a u t h o r s grouped similar a s s e m b l a g e s i n t o q u a n t i t a t i v e l y d e f i n e d f a u n u l e s w h i c h w e r e t h e n u s e d t o compare d e p o s i t i o n a l h i s t o r i e s on t h e two s i d e s o f t h e E n g l i s h C h a n n e l : marginal marine environments ( n e a r s h o r e s h e l f , d e l t a i c m a r i n e , l a g o o n a l , f l - u v i o m a r i n e , and marsh) i n t h e Hampshire B a s i n and a r e g i o n o f e n c l o s e d s h a l l o w s h e l f , l a g o o n a l , l i t t o r a l e n v i r o n m e n t s of normal o r h y p e r s a l i n i t y i n t h e P a r i s B a s i n a n d W e s t e r n A p p r o a c h e s . hlarked f a u n a l d i f f e r e n c e s o c c u r b e t w e e n t h e two b a s i n s :
-
1.
Textulariina
2.
M i l i o l i n a - c o s t a t e , a l v e o l i n e and a r e n a c e o u s i n d i v i d u a l s and s p e c i e s f a r more a b u n d a n t i n t h e P a r i s B a s i n a n d W e s t e r n A p p r o a c h e s .
3.
Rotali'ina
-
fei.7 s p e c i e s i n common t o t h e two a r e a s .
a ) t h e London C l a y h a s a d i s t i n c t a n d r e s t r i c t e d f a u n a of . .
.?c~alia, and
. .
?rrctrc
t h e P a r i s B a s i n a n d p r e s e n t o n l y i n t h e Upper B r a c k l e s h a m b e d s (Upper L u t e t i a n o f t h e H a m p s h i r e B a s i n ) ; c ) a l t h o u g h t h e E o c e n e b e d s o f t h e P a r i s B a s i n a n d W e s t e r n A p p r o a c h e s h a v e many s p e c i e s s markedly d i f f e r e n t , i n common, t h e c o m p o s i t i o n o f t h e f a u n e . g . P a r i s B a s i n : ijmis.n, Z L p k < & : m , 'cides, polymorphinids (even i n m i l i o l i d dominated a s s e m b l a g e s ) ; Western Approaches: Pnrsratnik, J o z a X a ( i n m i l i o l i d d o m i n a t e d a s s e m b l a g e s ) ; d ) some
396 s p e c i e s which o c c u r t y p i c a l l y i n t h e P a r i s B a s i n o c c u r b r i e f l y i n t h e H a m p s h i r e B a s i n i n Upper B r a c k l e s h a m b e d s ( F i s h e r b e d s X V I I /
21). On t h e o t h e r h a n d , ,r,:Ls::;as a n d ";sc<s:C?ee were snown t o h a v e s i m i l a r d i s t r i b u t i o n s t o t h e two a r e a s . A d i s t i n c t d e p e n d e n c e o f t h e a s s e m b l a g e s on l i t h o t o p e i s p o s t u l a t e d i n t h e t h r e e a r e a s . Thus, i n t h e H a m p s h i r e B a s i n r i v e r s d r a i n e d J u r a s s i c Cretaceous sediments with l a r g e c l a y content. In t h e P a r i s B a s i n r i v e r s drained massifs with l i t t l e clay content. Sea-grass and shallow water a t t a c h e d forms f l o u r i s h e d i n t h e r e s u l t i n g c l e a r w a t e r . I n t h e Western A p p r o a c h e s a n d C o n t e n t i n t h e low s e d i m e n t i n p u t a l s o r e s u l t e d i n c l e a r , s h a l l o w w a t e r i n which s e a - g r a s s , b r y o z o a n s , and c a l c a r e o u s a l g a e f l o u r i s h e d . The d i s t r i b u t i o n o f s e a - g r a s s was c o n s i d e r e d a m a j o r f a c t o r i n c o n t r o l l i n g t h e d i s t r i b u t i o n of benthonic Foraminifera. The g r e a t e r a m o u n t s o f f r e s h w a t e r e n t e r i n g t h e H a m p s h i r e B a s i n a r e r e f l e c t e d i n e s t i m a t e s o f h y p o s a l i n i t y (32-33 " / o o ) t h e r e v s . n o r m a l s a l i n i t y f o r t h e P a r i s B a s i n . During m i d d l e Eocene t i m e s a l i n i t i e s i n t h e (Cr 8 , peneP a r i s B a s i n - C o n t e n t i n r e g i o n may h a v e r e a c h e d 3 8 roplids). Summer t e m p e r a t u r e s i n e x c e s s o f 2 2 ° C a r e a l s o s u g g e s t e d b y t h e s e f a u n a s . A d d i t i o n a l i n f o r m a t i o n on t h e P a l e o g e n e b i o s t r a t i g r a p h y a n d biogeography of t h e Anglo-Paris Basin ( i n c l u d i n g f o r a m i n i f e r a l d a t a ) can be found i n t h e t h o r o u g h r e v i e w s by C u r r y ( 1 9 6 5 , 1 9 6 6 ) . A d d i t i o n a l d a t a on t h e Paleogene b e n t h o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y a n d p a l e o e c o l o g y o f NW E u r o p e may b e f o u n d in t h e s t u d i e s on t h e Eocene ( K a a s c h i e t e r , 1 9 6 1 ) a n d O l i g o c e n e ( B a t j e s , 1 9 5 8 ) o f B e l e i u m a n d t h e Mid-(Ulleberg, 1 9 7 4 ) and Late ( L a r s e n a n d D i n s e n , 1959) O l i g o c e n e of Denmark. I n g e n e r a l t h e l a g o o n a l - n e r i t i c n a t u r e o f t h e s e f a u n a l assemb l a g e s r e f l e c t t h e c o n t i n u i t y through t h e P a l e o g e n e of f l u c t u a t i n g , margina l d e p o s i t i o n a l c o n d i t i o n s a l o n g t h e Nh' E u r o p e a n c o a s t a l a r e a s . P a l e o c e n e d e e p w a t e r b e n t h o n i c f o r a m i n i f e r a l f a u n a s a r e known f r o m s e v e r a l t e c t o n i c a l l y a c t i v e b a s i n s on t h e p e r i m e t e r o f t h e A t l a n t i c i n E a s t e r n Mexico ( B a r k e r a n d B e r g g r e n , i n p r e s s ) , a n d t h e C a r i b b e a n ( T r i n i d a d , Venezuela, e t c . ) . T h e i r taxonomy, s t r a t i g r a p h i c a n d b i o g e o g r a p h i c d i s t r i b u t i o n i n t h e deep A t l a n t i c O c e a n a r e c u r r e n t l y b e i n g s t u d i e d by R . C . T j a l s m a ( W . H . O . I . ; s e e a l s o T j a l s m a a n d Lohmann, 1 9 7 5 , i n p r e s s ; Lohmann a n d T j a l s m a , 1 9 7 5 , i n p r e s s ) . F o u r m a j o r a s s e m b l a g e s were d i s t i n g u i s h e d b y a p r i n c i p a l c o m p o n e n t a n a l y s i s . T h r e e o f t h e m c o n t a i n a d i v e r s e , c a l c a r e o u s f a u n a a n d were f o u n d t o b e r e s t r i c t e d t o a p e l a g i c s e d i m e n t a r y e n v i r o n m e n t . They a r e :
8
~ y ~ < ~ si vi .. C - . f l v a- s ' ?s e m b l a g e
The f o u r t h , Smcoi.h
397 i t s e x t i n c t i o n during t h e l a t e s t Paleocene. Assemblage 1 c o n s i s t i n g o f long-range t a x a and c h a r a c t e r i z i n g deep s i t e s d u r i n g t h e Y i d d l e P a l e o c e n e , f o l l o w s t h i s t r e n d a n d o c c u r r e d a l s o a t i n t e r m e d i a t e d e p t n s d u r i n g t h e Late P a l e o c e n e . It i s t h e dominant assemblage a t a l l d e p t h s d u r i n g t h e e a r l i e s t Eocene a n d t e n d s t o b e more i m p o r t a n t i n t h e m i d - h i g h l a t i t u d e ( 3 0 " - 5 0 " ) s i t e s . A s s e m b l a g e 3 i s r e s t r i c t e d d u r i n g t h e P a l e o c e n e t o low-mid l a t i t u d e (0"-30") s i t e s . A d r a s t i c taxonomic r e p l a c e m e n t o c c u r r e d n e a r t h e Paleocene/Eocene boundary ( p r i m a r i l y a t t h e s p e c i e s l e v e l ) . The s i g n i f i c a n c e o f t h i s i s n o t yet clear. E o c e n e d e e p w a t e r s e d i m e n t s a r e known f r o m t h e C a r i b b e a n ( B a r b a d o s , Cuba, T r i n i d a d , V e n e z u e l a ) , a n d C e n t r a l A m e r i c a ( M e x i c o ) , b u t a l t h o u g h t h e i r b e n t h o n i c f o r a m i n i f e r a l f a u n a s h a v e be e n d e s c r i b e d by v a r i o u s i n v e s t i g a t o r s (Berggren and P h i l l i p s , 1 9 7 1 ) , t h e y have n o t been a n a l y z e d f o r c o m p a r a t i v e ( g e o g r a p h i c ) taxonomy o r q u a n t i t a t i v e b i o g e o g r a p h y . An examina t i o n o f v a r i o u s b a t h y a l a n d a b y s s a l f a u n a s from t h e Eocene and O l i g o c e n e suggests continued cosmopolitanism. It i s within the l a r g e r foraminifera that d i s t i n c t provincialism becomes n o t i c e a b l e . Adams ( 1 9 6 7 ) h a s a d m i r a b l y s u m m a r i z e d t h e g e o g r a p h i c d i s t r i b u t i o n o f T e r t i a r y l a r g e r F o r a m i n i f e r a i n t h e T e t h y a n , American and Indo-Pacific provinces. The T e t h y s ( o r M e d i t e r r a n e a n ) a n d / o r I n d o - P a c i f i c es, alveolinids, assilinids, a r e a s were t h e c e n t e r s o f d i s p e r s a l o f ES during t h e Paleogene. These forms o c c u r - P a r i s (where t h e y f i r s t a p p e a r e d i n l a t e F a r l y Eocene [ Y p r e s i a n ] t i m e ) a n d K o r t h German B a s i n s w h e r e t h e y a r e u s e d i n l o c a l and i n t e r - r e g i o n a l b i 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 s . These forms a r e a b s e n t i n t h e w e s t e r n A t l a n t i c a n d , i n f a c t , t h e r e i s n o warm-water c a r b o n a t e p r o v i n c e on t h e w e s t e r n s i d e o f t h e A t l a n t i c n o r t h o f t h e Caribbe a n a n d F l o r i d a - G u l f C o a s t a l P l a i n r e g i o n ( e . g . t h e .5e:ercs:a Late O l i g o c e n e a g e i n T e x a s ) . On t h e o t h e r h a n d l a r g e r F o r a m i n i f e r a common t o b o t h s i d e s o f t h e A t l a n t i c may h a v e h a d t h e i r o r i g i n d u r i n g t h e P a l e o g e n e i n t h e C a r i b b e a n (e.g. lepidocyclinids, miogypsinids). S u m e r o u s s t u d i e s h a v e b e e n made on t h e s e v a r i o u s g r o u p s , m o s t n o t a b l y by D r o o g e r a n d h i s s t u d e n t s a t U t r e c h t , C o l e a t C o r n e l l U n i v e r s i t y , C l a r k e a t B r i t i s h P e t r o l e u m i n London, a n d Blondeau and h i s c o l l e a g u e s i n P a r i s . The m i o g y p s i n i d s , i n p a r t i c u l a r , have been used i n b i o s t r a t i g r a p h i c s t u d i e s of t h e L a t e Paleogene-Early Neogene i n w e s t e r n E u r o p e , p a r t i c u l a r l y b e t w e e n t h e A q u i t a i n e a n d N o r t h German B a s i n s . I m p o r t a n t r e c e n t s t u d i e s on t h e b i o s t r a t i g r a p h y o f P a l e o g e n e l a r g e r F q r a m i n i f e r a o f t h e C a r i b b e a n i n c l u d e t h o s e by K u g l e r and C a u d r i (1975) and C a u d r i (1975) on t h e Paleocene-Eocene o f S o l d a d o Rock, T r i n i d a d and B a r k e r a n d Blow ( 1 9 7 6 ) o n t h e M i d d l e Eocene-Lower bliocene of t h e T a m p i c o - M i s a n t l a Embayment, M e x i c o . The p h y l o g e n y o f v a r i o u s o r b i t o i d a l l a r g e r F o r a m i n i f e r a i n t h e C a r i b b e a n E o c e n e w a s d i s c u s s e d by C a u d r i ( 1 9 7 5 ) . I n t h e l a t t e r p a p e r B a r k e r a n d Blow ( 1 9 7 6 ) h a v e p l a c e d X i d d l e E o c e n e t h r o u g h Lower Miocene f o r m a t i o n a l u n i t s i n t h e T a m p i c o - M i s a n t l a Embayment o f Mexico i n a p l a n k t o n i c f o r a m i n i f e r a l b i o s t r a t i p r a p h i c framework a n d c a l i b r a t e d t h e s t r a t i g r a p h i c r a n g e s o f t h e l a r g e r F o r a m i n i f e r a t o t h i s s c h e m e . The o c c u r r e n c e o f 135ioleptdzzxa. rs3Zeu.i i n t h e t y p e a r e a o f t h e T a n t o y u c a F o r m a t i o n ( i n l e v e l s r e f e r r a b l e t o l a t e s t Zone P 1 6 a n d / o r e a r l i e s t Zone P17 = Upper E o c e n e ) a n d e v e n a t l e v e l s r e f e r r a b l e t o Zone P 1 5 ( B a r k e r a n d Blow, 1 9 7 6 , p . 42, f i g . 2 ) s u p p o r t S t a i n f o r t h ' s (1965) arguments a ) t h a t i t i s a s t r i c t l y Upper E o c e n e f o r m b ) r e f u t i n g t h e a r g u m e n t p u t f o r t h b y Eames e2 T:. ( 1 9 6 2 , p . 41, 4 2 , 70, 71-81) i n s u p p o r t o f a m a j o r h i a t u s w i t h i n t h e San F e r n a n d o F o r m a t i o n o f T r i n i d a d a n d , b y e x t e n s i o n , of a m a j o r , v i r t u a l l y worldwide, intra-Oligocene h i a t u s . C i r c u m - A t l a n t i c and C a r i b b e a n s t r a t i g r a p h i c u n i t s t e m p o r a r i l y e x c l u d e d f r o m t h e O l i g o c e n e b y Eames 9; 7:. (1962)
398 h a v e now b e e n r e h a b i l i t a t e d , and t h e O l i g o c e n e a p p e a r s t o h a v e b e e n r e i n s t a t e d i n t h e h i e r a r c h y of L y e l l i a n Cenozoic c h r o n o s t r a t i g r a p h y a l t h o u g h i t s upper a n d l o w e r l i m i t s , i n terms o f f o r a m i n i f e r a l b i o s t r a t i g r a p h y , r e m a i n a matter of animated d i s c u s s i o n . The s e q u e n t i a l a p p e a r a n c e o f l e p - : d o e ; : cl <~ a( 3 ~ c e n e ( P 1 9 ) l e v e l s of t h e H o r c o n e s F o r m a t i o n is f r o m N2 = P 2 1 ( a n d q u e s t i o n a b l y as low a TZ f r o m t h e Upper Palma Real F o r m a t i o n (N2 = P21) a n d .'4 e s o n F o r m a t i o n (N3 = P 2 2 ) ( s e r s : ~B. a r k e r i n B a r k e r a n d B s i n i l a r t o t h a t which o c c u r s i n Europe and f a c i l i t a t e s L a t e O l i g o c e n e t r a n s - c o r r e l a t i o n b a s e d upon l a r g e r F o r a m i n i f e r a . The 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 o f P a l e o g e n e b e n t h o n i c F o r a m i n i f e r a i n t h e S o r t h A t l a n t i c h a s b e e n d i s c u s s e d i n r e c e n t s t u d i e s by B e r g g r e n (1972, 1974a) and Berggren and Aubert ( 1 9 7 6 a , b ) . In t h e s e s t u d i e s t h e d i s t r i b u t i o n o f b e n t h o n i c f a u n a s was u s e d i n d e l i n e a t i n g t h e s u b s i d e n c e h i s t o r y o f two c o n t i n e n t a l r e m n a n t s - O r p h a n Knoll i n t h e L a b r a d o r S e a a n d R o c k a i l Bank i n t h e NE A t l a n t i c - w h i c h w e r e f o r m e d d u r i n g e a r l y s t a g e s o f c o n t i n e n t a l r i f t i n g and sea-floor spreading. I n t h e c a s e o f R o c k a l l Bank t h r e e b e n t h o n i c f o r a m i n i f e r a l z o n u l e s s p a n a b o u t 5 m.y. o f Late P a l e o c e n e - E a r l y E o c e n e t i m e r e f l e c t i n g p r o g r e s s a b o u t 200 m . The n o r t h the b a s a l sediments j u s t ere recently described as e a n d Wong ( 1 9 7 4 ) . The L
s was p r o b a b l y d e p o s i t e d i n w a t e r d e p t h s o f l e s s t h a n 1 0 meters. The m i d d l e z o n u l e c o n t a i n e d a n e s s e n t i a l l y ?lidway f a u n a , w i t h n u m e r o u s e l e m e n t s t y p i c a l o f t h e G u l f C o a s t blidway, a s w e l l a s t h e P a l e o c e n e f a u n a s d e s c r i b e d f r o m G r e e n l a n d ( H a n s e n , 1 9 7 0 ) a n d Nw E u r o p e ( B r o t z e n , 1 9 4 6 ) . The u p p e r z o n u l e e x h i b i t e d s i m i l a r i t i e s w i t h t h e " l o w e r E o c e n e 3" f a u n a s o f NW E u r o p e ( S t a e s c h e a n d H i l t e r mann, 1 9 4 0 ; B e t t e n s t a e d t , 1 9 4 9 ) a n d r e p r e s e n t e d d e p t h s o f a b o u t 200 m, n e a r t h e s h e l f - s l o p e margin. By L a t e E o c e n e t i m e R o c k a i l Bank h a d s u n k t o m i d d l e b a t h y a l d e p t h s ( r ' - 700-900 m) h t t a Z Z i d e s trucmpyi a n d a n i n d e t e r m i n a t e o s a n g u l a r i i d c h a r a c t e r i z e t h e uppermost Eocene o f R o c k a l l Bank. T h e i r e x t i n c t i o n h e r e , synchronous w i t h v a r i o u s o t h e r b a t h y a l forms e l s e w h e r e c o i n c i d e s w i t h t h e Eocene/Oligocene boundary, a r e l a t i v e l y sudden r e d u c t i o n i n b o t t o m w a t e r ) and t h e e s t a b l i s h m e n t of t h e p s y c h r o t t a n d S h a c k l e t o n , 1 9 7 6 ) . The O l i g o c e n e a t R o c k a l l Bank i s c h a r a c t e r i z e d b y Jetero!e?a -exic?.ra ( C u s h m a n ) , SCp%c.ra Common a c c e s s o r y (Cushman) a n d 3. ~ d ; e ~(Cushman) forms i n c l u d e r e ; : z < Cushman and S t a i n f o r t h , 3. v % ! ~ * ~ a LH ~ a dml e y , z.n& ( R e u s s ) , ;r rs aSSs e c.is6.j .PE M R is ( Ga 11owa y a n d as:-rsls ( B e r m u d e z ) , f o r m s common i n b o t h M e d i t e r r a n e a n and C a r i b b e a n mid-Cenozoic b a t h y a l s e q u e n c e s . Sporadic, but p e r s i s t e n t , s t i l o s t o m e l l i d s , p l e u r o s t o m e l l i d s and u v i g e r i n i d s suggest middle b a t h y a l d e p o s i t i o n a l depth i n Late Paleogene t i m e . Xeritic faunal e l e m e n t s , c h a r a c t e r i s t i c o f t h e C h a t t i a n s t a g e o f NW E u r o p e , o c c u r a t c e r t a i n l e v e l s t o g e t h e r w i t h b a t h y a l e l e m e n t s , i n d i c a t i n g d i s p l a c e m e n t from m a r g i n a l a r e a s o f R o c k a l l Bank a n d p r o v i d i n g a n i n s i g h t i n t o t h e L a t e Oligocene shallow water faunas i n t h e a r e a . R o c k a l l Bank h a s r e m a i n e d e s s e n t i a l l y a t i t s p r e s e n t d e p t h (.^. 1100 m) t h r o u g h o u t t h e Neogene. A g r a d u a l taxonomic r e p l a c e m e n t c a n b e o b s e r v e d i n t h e e a r l y - m i d d l e Miocene w i t h t h e L a t e P a l e o g e n e f a u n a e v o l v i n g i n t o t h e p r e s e n t day f a u n a . There i s no e v i d e n c e o f any r e l a t e d t e c t o n i c e v i d e n c e a c c o m p a n y i n g t h e f a u n a l c h a n g e a t a b o u t 1 5 ?la.
.
.
399 Eocene b e n t h o n i c f o r a m i n i f e r a l f a u n a s a t Orphan K n o l l ( L a b r a d o r Sea) ides ( 6 s p e c i e s , ) towLia ( 4 species) a ( 3 s p e c i e s e a c h ) . Predominant
r'sdaims ( R e u s s ) , 3 ~ z : c a Cushman, 5'.
a:n
Cushman
(de W i t t Puyt) and a s s e m b l a g e s u g g e s t s d e p o s i t i o n a t l o w e r b a t h y a l t o a b y s s a l d e p t h s (10002000 m) a n d s u p p o r t s t h e i n t e r p r e t a t i o n made b y DSDP Leg 1 2 g e o l o g i s t s t h a t Orphan K n o l l s a n k t o i t s p r e s e n t d e p t h d u r i n g P a l e o c e n e t i m e . E s s e n t i a l l y c o n t e m p o r a n e o u s Eocene a s s e m b l a g e s from t h e c e n t r a l , d e e p e r p a r t of t h e L a b r a d o r S e a c o n s i s t a l m o s t w h o l e l y o f a g g l u t i n a t e d .s p e c i e s o f , f.0,. , CrCSrcsrow~,ses and p r o b a b l y i n d i c a t e d e p o s i t i o n n e a r o r below t h e l o c a l c a r b o n a t e compensation d e p t h i n d e e p c o l d waters ( B e r g g r e n , 1 9 7 2 , p . 4 7 4 ) . North Sea benthonic f o r a m i n i f e r a l faunas g e n e r a l l y e x h i b i t t h e following sequence during the Paleogene: &::&? E a r l y P a l e o c e n e ( D a n i a n ) - c a l c a r e o u s a s s e m b l a g e s (%ie a n d v a r i o u s c i b i c i d i d s , a n o m a l i n i d s ) - whose a n c e s t r y may b e t r a c e d i n t o t h e underlying Cretaceous carbonate sequences - i n d i c a t i n g d e p o s i t i o n i n a n e x t e n s i v e c a r b o n a t e s h e l f p r o v i n c e a t d e p t h s of l e s s t h a n 200 m , f o l l o w e d a b r u p t l y by
P a l e o c e n e ( L a t e D a n i a n - T h a n e t i a n ) - a g g l u t i n a t e d a s s e m b l a g e s (. '?a, 7r<,!;r e s , .3ec2motdes, Spir;p!azto sCs, <.aZ.) r e f l e c t i n g r a p i d s u b s i d e n c e , t u r b i d i t i c f a c i e s a n d d e p o s i t i o n a l d e p t h s i n e x c e s s o f 1000 m , f o l l o w e d by Eocene (Ypresian)-Oligocene ( C h a t t i a n ) - Gradual b a l a n c e between c a l c a r e o u s a n d a g g l u t i n a t e d f a u n a l components r e f l e c t i n g d i f f e r e n t i a l rates of b a s i n f i l l i n g . P a l e o g e n e b e n t h o n i c f o r a m i n i f e r a l f a u n a s o f t h e Norwegian G r e e n l a n d S e a a r e known f r o m t h e I c e l a n d - F a e r o e s R i d g e (DSDP S i t e 3 3 6 , N o r t h F l a n k ; 3 5 2 , S o u t h F l a n k ) , I c e l a n d P l a t e a u (DSDP S i t e 3 4 8 ) a n d t h e J a n Hayen R i d g e (DSDP S i t e s 3 4 6 , 3 4 7 , 3 4 9 ) ( T a l w a n i , U d i n t s e v e t aZ., 1 9 7 5 ) . A g g l u t i n a t e d a s s e m b l a g e s c h a r a c t e r i z e t h e L a t e P a l e o g e n e ( O l i g o c e n e ) of t h e I c e l a n d P l a t e a u ( 3 4 8 ) a n d J a n Hayen R i d g e ( 3 4 6 , 3 4 7 , 3 4 9 ) . Two d i f f e r e n t s e q u e n c e s were f o u n d o n t h e n > r t h a n d s o u t h f l a n k s o f t h e I c e l a n d - F a e r o e s R i d g e . On
Oligocene assemblages i d s , polymorphinids, f i s s u r i n i d s , i n d i c a t i v e of o u t e r n e r i t i c - u p p e r b a t h y a l d e p t h s o f w a t e r . The t e n t a t i v e c o n c l u s i o n was d r a w n t h a t t h e n o r t h f l a n k o f t h e I c e l a n d - F a e r o e s R i d g e h a d no c o n n e c t i o n w i t h t h e N o r t h A t l a n t i c t o t h e s o u t h e s s e n t i a l l y d u r i n g most o f t h e P a l e o g e n e and t h a t i t s e v e n t u a l s u b s i d e n c e was a p o s t - l l i d d l e O l i g o c e n e phenomenon ( T a l w a n i , U d i n t s e v e : ,?l.,
1975).
400 The P a l e o g e n e b e n t h o n i c f o r a m i n i f e r a l f a u n a s i n d i c a t e s i g n i f i c a n t v e r t i c a l movement o f t h e s e a f l o o r i n t h e N o r w e g i a n - G r e e n l a n d S e a a n d a r e c u r r e n t l y u n d e r g o i n g i n v e s t i g a t i o n i n a n a t t e m p t t o u n r a v e l t h e complex Cenozoic g e o t e c t o n i c h i s t o r y of t h i s a r e a .
SEOGENE
The m a j o r p a l e o g e o g r a p h i c a n d / o r p a l e o c l i m a t i c e v e n t s w h i c h a f f e c t e d c i r c u l a t i o n ( a n d , c o n c o m i t a n t l y f o r a m i n i f e r a l biogeography) i n t h e North . \ t l a n t i c Ocean d u r i n g t h e p a s t 2 5 >la i n c l u d e : S e p a r a t i o n o f t h e e a s t e r n and w e s t e r n T e t h y s by t h e j u n c t i o n of A f r i c a a n d E u r a s i a a b o u t 18 !la ( B u r d i g a l i a n Age) Growth a n d e x p a n s i o n o f t h e A n t a r c t i c I c e Cap d u r i n g t h e ? f i d d l e - L a t e Miocene (12-6 ?la) J u n c t i o n of Europe and S o r t h A f r i c a , and g r a d u a l c l o s u e and e v e n t u a l s e q a r a t i o n o f w e s t e r n T e t h y s a n d A t l a n t i c Ocean b a s i n s d u r i n g t h e L a t e >lioci,ne (10-5 bIa) R e c o n n e c t i o n o f A t l a n t i c a n d M e d i t e r r a n e a n i n E a r l y F l i o c e n e ( 5 ?la) E a r l y P l i o c e n e ( 3 . 5 - 4 . 0 ?la) e l e v a t i o n o f t h e I s t h m u s o f Panama s e v e r i n g t h e marine c o n n e c t i o n a n d f a u n a l i n t e r c h a n g e between t h e A t l a n t i c and P a c i f i c Oceans ? l i d - P l i o c e n e ( 3 ?la) i n i t i a t i o n o f p o l a r g l a c i a t i o n i n t h e N o r t h e r n Hemisphere, tlir ( p r o b a b l e ) f o r m a t i o n of t h e Labrador c u r r e n t a s a s i g n i f i c a n t water mass a n d t h e d e v e l o p m e n t o f a P o l a r ( A r c t i c ) f a u n a l province.
Planktonic Foraminifera ._____ The g r a d u a l c o o l i n g o f t h e e a r t h d u r i n g t h e Neogene r e s u l t e d i n s t r o n g e r l a t i t u d i n a l t h e r m a l g r a d i e n t s a n d , c o n c o m i t a n t l y , i n a more pronounced p r o v i n c i a l i z a t i o n of p l a n k t o n i c f o r a m i n i f e r a l f a u n a s i n t h e N o r t h A t l a n t i c . A t t h e same t i m e t h e c o n t i n u e d p r e s e n c e o f t h e G u l f S t r e a m as f a r n o r t h a s t h e L a b r a d o r S e a u n t i l m i d - P l i o c e n e t i m e (%. 3 ?la) b r o u g h t t r o p i c a l f a u n a s as f a r n o r t h as 54"N L a t . i n t h e w e s t e r n A t l a n t i c , w h e r e a s c o n t e m p o r a n e o u s f a u n a s i n t h e c e n t r a l a n d e a s t e r n X o r t h A t l a n t i c w e r e of cool t e m p e r a t e n a t u r e . The i n c r e a s i n g p r o v i n c i a 1i za t i on o f A t 1a n t i c p 1a n k t on i c f o ram i n i f e r a 1 f a u n a s i n t h e L a t e Neogene - d u e t o t h e c o m b i n e d e f f e c t s o f c h a n g i n g c o n t i n e n t - o c e a n geometry and c l i m a t i c d e t e r i o r a t i o n r e n d e r s t h e a p p l i c a t i o n o f a s t a n d a r d m o n d i a l t r o p i c a l z o n a t i o n scheme i n o p e r a b l e i n t h e A t l a n t i c O c e a n , p a r t i c u l a r l y f r o m t h e L a t e M i o c e n e o n . A s a r e s u l t , a s e p a r a t e T-ate !!iocene-Pliocene l o w - l a t i t u d e ( B e r g g r e n , 1 9 7 3 ; i n p r e s s ) and h i g h l a t i t u d e ( B e r g g r e n , 1972; Poore a n d B e r g g r e n , 1975) z o n a t i o n s have b e e n developed f o r t h e N o r t h A t l a n t i c which r e f l e c t t h e i n c r e a s i n g p r o v i n c i a l i t y o f A t l a n t i c faunas during the Late Seogene. During t h e E a r l y Xiocene t h e dominant f a u n a l e l e m e n t s i n t h e North A t l a n t i r ( e x c l u s i v e of t h e l o w - l a t i t u d e , t r o p i c a l r e g i o n and t h e w e s t e r n A t l a n t i c - i n f l u e n c e d Gulf S t r e a m ) a r e t h e l a r g e , r o b u s t g l o b o q u a d r i n i d s and - ^LO,. c", , globigerinitids. The n-,9
i_
-
401 l i n e a g e s c a n b e used i n b i o s t r a t i g r a p h i c s u b d i v i s i o n d u r i n g t h e Middle-Late ' e n e a n d Ea e ne, respectively. The a p p e a r a n c e o f . . a;; a n d ,-ie ~ . < . / npanr.?ser-a c o i n c i d e s w i t h t h e f i ted i g n i f i e s t h e development of a p o l a r f a u n a l r e a l m i n itus (at 3 the circum-Arctic region. T h i s r e p r e s e n t s t h e c o m p l e t i o n of t h e g r a d u a l p r o c e s s of f a u n a l p r o v i n c i a l i z a t i o n which had been t a k i n g p l a c e d u r i n g t h e Cenozoic. A f t e r t h i s t h r e e main f a u n a l p r o v i n c e s c a n b e s e e n i n t h e Xorth Atlantic: tropical, boreal, polar In t h e N o r t h S e a p l a n k t o n i c F o r a m i n i f e r a became s p a r s e r d u r i n g t h e l a t e r Neogene a s a r e s u l t o f b a s i n f i l l i n g a n d c o n c o m i t a n t s h a l l o w i n g o f facies. I n t h e Norwegian-Greenland Sea p l a n k t o n i c F o r a m i n i f e r a a r e v i r t u a l l y a b s e n t d u r i n g t h e Neogene u n t i l t h e m i d - P l i o c e n e ( 3 Ma) w h e n , a r g l a c i a t i o n , a p o l a r f a u n a a p p e a r s d o m i n a t e d by with t h e advent - . s i n i s t r a l ..,Y L , ; z sncq$aerT;. c t o r r e g r e s s i o n ) i n v e s t i g a t i o n s o f p l a n k t o n i c forami n i f e r a l a s s e m b l a g e s b y t h e CLIllAP g r o u p h a v e r e s u l t e d i n t h e d e l i n e a t i o n of o c e a n i c p a l e o c l i m a t i c t r e n d s o v e r t h e p a s t 1 Ma. The more i n p o r t a n t r e s u l t s of t h i s major program i n c l u d e t h e e l u c i d a t i o n o f :
1)
R e p e a t e d ( p e r h a p s a s many a s 30 o r m o r e ) g l a c i a t i o n s a t h i g h a n d midl a t i t u d e s i n t h e n o r t h e r n h e m i s p h e r e o v e r t h e p a s t 1 . 5 Ma.
2)
R e p e a t e d a n d d r a s t i c l a t i t u d i n a l d i s p l a c e m e n t o f c l i m a t i c z o n e s by a s much a s 2 0 " - 3 0 " .
3)
Dramatic c h a n g e s i n t h e North A t l a n t i c c i r c u l a t i o n and e v i d e n c e t h a t most o f t h e Sorwegian-Greenland Sea h a s b e e n i c e - c o v e r e d f o r a b o u t 100,000 o f t h e p a s t 1 2 7 , 0 0 0 y e a r s .
The i n t e g r a t e d g e o p h y s i c a l , g e o c h e m i c a l a n d p a l e o n t o l o g i c s t u d i e s on t h e d e e p - s e a Late P l e i s t o c e n e s t r a t i g r a p h i c r e c o r d o f t h e X o r t h A t l a n t i c a r e l e a d i n g t o a b e t t e r understanding of g l o b a l c l i m a t e over t h e p a s t m i l l i o n y e a r s . The g o a l o f t h i s p r o g r a m i s t h e f o r m u l a t i o n o f q u a n t i t a t i v e s y n o p t i c p a l e o t e m p e r a t u r e maps f o r s e l e c t e d s p e c i f i c "moments" i n t i m e ( N c I n t y r e , Moore e z a;., 1 9 7 6 ) w h i c h c a n , i n t u r n , s e r v e a s b o u n d a r y c o n d i t i o n s f o r modeling g e n e r a l atmospheric c i r c u l a t i o n p a t t e r n s i n the P l e i s t o c e n e ( G a t e s , 1 9 7 6 ) a n d s e r v e as a r e l i a b l e means t o p r e d i c t i n g future global climatic trends. The Neogene s t r a t i g r a p h i c s u c c e s s i o n i n t h e A t l a n t i c C o a s t a l P l a i n h a s b e e n p u t i n a p l a n k t o n i c h i o s t r a t i g r a p h i c f r a m e w o r k by t h e i n v e s t i g a t i o n s o f A k e r s ( 1 9 7 2 ) , a n d i n t h e C a r i b b e a n b y B o l l i ( 1 9 5 7 1 , Blow ( 1 9 6 9 1 , a n d Lamb a n d B e a r d ( 1 9 7 2 ) . Of p a r t i c u l a r s i g n i f i c a n c e f o r A t l a n t i c Coastal P l a i n s t r a t i g r a p h y has been t h e demonstration t h a t t h e Chipola F o r m a t i o n o f F l o r i d a i s o f B u r d i g a l i a n Age (Zone S 7 ) a n d t h e Yorktown Formation - l o n g c o n s i d e r e d Late Xiocene - i s a c t u a l l y of P l i o c e n e (pred o m i n a n t l y Z a n c l e a n ) a g e ( A k e r s , 1 9 7 2 ) . S u b s e q u e n t work h a s s h o w n , i n f a c t , t h a t t h e "Yorktown" as u s e d i n t h e SE A t l a n t i c C o a s t a l P r o v i n c e e s s e n t i a l l y spans the e n t i r e Pliocene (Hazel, i n p r e s s ) . In w e s t e r n E u r o p e t h e N i o c e n e o f n o r t h e r n B e l g i u m ( H o o y b e r g h s a n d De X e u t e r , 1 9 7 2 ) , a n d t h e Upper 4 l i o c e n e o f SW S p a i n ( V e r d e n i u s , 1 9 7 0 ; T j a l s m a , 1 9 7 1 ; B e r g g r e n a n d Haq, 1 9 7 6 ) h a v e b e e n p u t i n t o p l a n k t o n i c forami n i f e r a l zonal perspective. I n t h e c a s e o f t h e B e l g i a n Miocene p l a n k t o n i c f o r a m i n i f e r a 1 faunas r e v e a l e d s i g n i f i c a n t temporal gaps between t h e v a r i o u s l i t h i c u n i t s which had p r e v i o u s l y b e e n c o n s i d e r e d t o have b e e n b r i e f o r non-existent.
402 Benthonic Foraminifera Neogene s h a l l o w w a t e r b e n t h o n i c f o r a m i n i f e r a l f a u n a s e x h i b i t a g r a d u a l d e c l i n e i n a m p h i - A t l a n t i c c o s m o p o l i t a n i s m as a r e s u l t o f more p r o n o u n ced c l i m a t i c g r a d i e n t s o v e r t h e g r e a t e r t r a n s - o c e a n i c and i n t e r c o n t i n e n t a l d i s t a n c e s c a u s e d by t h e combined e f f e c t s o f c o n t i n e n t a l d r i f t a n d s e a - f l o o r spreading. S h a l l o w w a t e r Neogene m a r i n e d e p o s i t s a r e e x t r e m e l y s p a r s e i n t h e U.S. A t l a n t i c a n d G u l f C o a s t a l p r o v i n c e s , a s w e l l as i n W e s t e r n E u r o p e . Those t h a t a r e p r e s e n t c o n t a i n f a u n a s w i t h p r e d o m i n a n t l y c i b i c i d i d s , nonionids, e l p h i d i i d s , r o t a l i i d s . Neogene b e n t h o n i c f o r a m i n i f e r a l a s s e m b l a g e s o f t h e A t l a n t i c C o a s t a l P l a i n a r e t y p i f i e d b y t h o s e d e s c r i b e d b y Cushman ( 1 9 3 0 , 1 9 3 3 ) f r o m t h e C h o c t a w h a t c h e e S t a g e o f F l o r i d a , P u r i ( 1 9 5 3 ) f r o m t h e Alum B l u f f S t a g e o f F l o r i d a a n d by Gibson (1962) from M a r y l a n d - V i r g i n i a ( s e e a l s o B e r g g r e n a n d P h i l l i p s , 1 9 7 1 , f o r a d d i t i o n a l r e f e r e n c e s ) . S i m i l a r s t u d i e s h a v e b e e n made i n Western Europe by t e n Dam a n d R e i n h o l d (1941, 1942) and v a n Voorthuysen ( 1 9 5 8 ) on t h e Neogene f a u n a s o f H o l l a n d a n d B e l g i u m , r e s p e c t i v e l y , and b y M a r g e r e l ( 1 9 6 8 ) on t h e P l i o c e n e o f B r i t t a n y . The d i s r u p t i o n o f t h e t r o p i c a l T e t h y a n s e a w a y i n t h e E a r l y M i o c e n e l e d t o s e p a r a t e a n d d i s t i n c t p h y l e t i c t r e n d s among t h e l a r g e r F o r a m i n i f e r a . Thus t h e m i o g y p s i n i d s a n d l e p i d o c y c l i n i d s h a d e s s e n t i a l l y r u n t h e i r c o u r s e by e a r l i e s t H i d d l e Miocene t i m e i n w e s t e r n Europe and t h e C a r i b b e a n , whereas they continued t o evolve i n t h e e a s t e r n Tethys and Indo-Pacific r e g i o n u n t i l n e a r t h e M i d d l e / L a t e Miocene b o u n d a r y . Recent (unpublished) e v i d e n c e i n d i c a t e s t h a t some f o r m s may h a v e s u r v i v e d e v e n l o n g e r . Neogene b a t h y a l b e n t h o n i c f o r a m i n i f e r a l f a u n a s are known i n t h e C a r i b b e a n (Cuba, P u e r t o R i c o , V e n e z u e l a , Dominican R e p u b l i c , e t c . ) and from t h e E-W r e e n t r a n t o f t h e A t l a n t i c O c e a n i n SW S p a i n , t h e G u a d a l q u i v i r B a s i n , among o t h e r p l a c e s . R e c e n t s t u d i e s on Late M i o c e n e b e n t h o n i c f o r a m i n i f e r a l f a u n a s i n two s e c t i o n s o f t h e w e s t e r n G u a d a l q u i v i r B a s i n ( B e r g g r e n a n d Haq, 1 9 7 6 ; B e r g g r e n e t al., i n p r e s s ) h a v e a l l o w e d t h e d e l i n e a t i o n o f i n t r a b a s i n a l e v o l u t i o n f r o m m i d d l e b a t h y a l ( 1 5 0 0 m) t o i n n e r - m i d d l e n e r i t i c < 5 0 m) d e p t h s o v e r a n i n t e r v a l o f a b o u t 5 Ma. The o l d e r s e c t i o n , E l C u e r v o (T = 5 11-10 :"la) c o n t a i n s a f a u n a d o m i n a t e d b y L?ikicidoides me&:+ P i s ( F i n l a y ) ( = C. pse2doxnger Cushman, 1 9 3 1 , ? I G ~Cushman, 1 9 2 2 ) ) , ,lon?k scZdar.ii ( d ' o r b i g n y ) , F a : ~ u e Z . Z @ m t f i ~( S~cfh~w a g e r ) a n d s u c h a (Brady) , Grid;rsnli,s x h n s t u s a c c e s s o r y f o r m s a s EpisroF4neL7, ( B r a d y ) , a n d v a r i o u s s t i l o s t o m e l l i d s a n d was i n t e r p r e t e d as h a v i n g b e e n Thc o t h e r s e c t i o n , d e p o s i t e d a t l o w e r b a t h y a l (ca. 1.500 m) w a t e r d e p t h s . Carmona-Dos H e r m a n o s , s t r a t o t y p e o f t h e A n d a l u s i a n S t a g e , c o n t a i n s a f a u n a l s e q u e n c e w h i c h i n d i c a t e s upward s h a l l o w i n g f r o m m i d d l e b a t h y a l ( c a . 8 0 0 m) d e p t h s a t t h e b o t t o m t o i n n e r n e r i t i c ( < 5 0 m) a t t h e t o p o v e r a 4-5 my interval. A p a r t i c u l a r l y s h a r p c h a n g e i n w a t e r d e p t h (ca. 50-70 m) w i t h i n t h e u p p e r p a r t o f t h e A n d a l u s i a n ( c a . 5 . 5 Ma) was l i m i t e d t o a n a p p a r e n t l y c o n t e m p o r a n e o u s s i g n i f i c a n t e u s t a t i c f a l l in s e a - l e v e l ( b a s e d on o x y g e n i s o t o p e d a t a i n SW P a c i f i c d e e p - s e a c o r e s ) r e l a t e d t o a m a j o r e x p a n s i o n o f t h e A n t a r c t i c I c e Cap. A s i m u l t a n e o u s a n d similar f a u n a l c h a n g e o c c u r s i n t h e Mediterranean Basin. Thus t h e c o n t e m p o r a n e o u s e l i m i n a t i o n of b e n t h o n i c f o r a m i n i f e r a l faunas r e f l e c t a sudden e u s t a t i c s e a - l e v e l f a l l , t h e former i n a b a s i n which m a i n t a i n e d o p e n c o n n e c t i o n w i t h t h e A t l a n t i c , t h e l a t t e r i n a b a s i n w h i c h t e m p o r a r i l y l o s t c o n n e c t i o n w i t h t h e A t l a n t i c a n d became a b a s i n o f i n t e r i o r d r a i n a g e d u r i n g t h e M e s s i n i a n Age. The i m p l i c a t i o n s o f t h i s A t l a n t i c f a u n a f r o m SW S p a i n a r e o f m a j o r i m p o r t a n c e i n u n d e r s t a n d i n g t h e Late M i o c e n e g e o l o g i c a l h i s t o r y o f t h e X e d i t e r r a n e a n B a s i n (Van C o u v e r i n g e t sl., i n p r e s s ) . Neogene b a t h y a l a n d a b y s s a l b e n t h o n i c f o r a m i n i f e r a l f a u n a s h a v e b e e n r e c o r d e d f r o m R o c k a l l Bank (DSDP S i t e 1 1 6 ) i n t h e N o r t h A t l a n t i c a n d t h e
Bay o f B i s c a y (DSDP S i t e s 1 1 8 , 1 1 9 ) ( B e r g g r e n , 1972), r e s p e c t i v e l y . E a r l y Miocene a s s e m b l a g e s a t R o c k a l l a n d B i s c a y a r e e s s e n t i a l l y t h e same a s t h o s e which o c c u r i n t h e O l i g o c e n e . In t h e M i d d l e Y i o c e n e a c h a n g e i s s e e n i n b o t h a r e a s . P r e l i m i n a r y a n a l y s i s of t h i s m a t e r i a l ( B e r g g r e n , 1 9 7 2 ) s u g g e s t s t h a t modern b a t h y a l - a b y s s a l b e n t h o n i c f o r a m i n i f e r a l f a u n a s ( B a r k e r , 1 9 6 0 ) d e v e l o p e d i n t h e M i d d l e M i o c e n e a b o u t 1 5 Ma. The e s s e n t i a l l y conservative evolution i n t h e s e faunas enhances the p o s s i b i l i t y of paleoe c o l o g i c , paleobathymetric and paleohydrologic i n t e r p r e t a t i o n s over t h e p a s t 1 5 Ma ( s e e a b o v e ) . In t h e N o r t h S e a b e n t h o n i c F a r a m i n i f e r a r e f l e c t g r a d u a l l y s h o a l i n g c o n d i t i o n s . P l i o c e n e - P l e i s t o c e n e a s s e m b l a g e s a r e d o m i n a t e d by a n 5:pk< L.T- C i k ctse s- m i 1i o 1i d a s s emb 1a g e s , i n many i n s t a n c e s In t h e N o r w e g i a n - G r e e n l a n d S e a a g g l u t i n a t e d a s s e m b l a g e s c h a r a c t e r i z e t h e p r e - g l a c i a l Neogene s e q u e n c e s o f t h e I c e l a n d a n d V d r i n g P l a t e a u s , whereas c a l c a r e o u s assemblages c h a r a c t e r i z e t h e g l a c i a l l a t e PlioceneP l e i s t o c e n e s e q u e n c e s t h e r e a n d o n t h e J a n Mayen R i d g e a n d I c e l a n d - F a e r o e s Ridge. Dominant FJeogene f a u n a l e l e m e n t s i n t h e N o r w e g i a n - G r e e n l a n d S e a s i t e s a r e shown i n t a b u l a r f o r m b e l o w ( t a b l e 1 ) .
?c
<
.
SUMMARY 1.
The s t r a t i g r a p h i c a n d g e o g r a p h i c d i s t r i b u t i o n o f C e n o z o i c p l a n k t o n i c a n d b e n t h o n i c f o r a m i n i f e r a l f a u n a s o f t h e A t l a n t i c Ocean a n d a d j a c e n t a r e a s a r e b e s t u n d e r s t o o d w i t h i n t h e framework o f d y n a m i c a l l y c h a n g i n g r e l a t i o n s h i p s between o c e a n and c o n t i n e n t s and t h e r e s u l t i n g changes i n c i r c u l a t i o n a n d c l i m a t e which t h e y h a v e c a u s e d .
2. P l a n k t o n i c f o r a m i n i f e r a l f a u n a s e x h i b i t a n i n c r e a s i n g d e g r e e o f l a t i t u d i n a l p r o v i n c i a l i z a t i o n d u r i n g t h e C e n o z o i c . D u r i n g t h e L a t e Neogene (ma. l a s t 15 my) t h i s p r o v i n c i a l i z a t i o n h a s become s o p r o n o u n c e d t h a t m u l t i p l e z o n a t i o n s f o r low and h i g h l a t i t u d e s must b e s u b s t i t u t e d f o r ( " . e . r e p l a c e ) t h e " s t a n d a r d " m o n d i a l , low l a t i t u d e z o n a t i o n ( s ) The p r o c e s s o f f a u n a l a p p l i c a b l e d u r i n g p r e - L a t e Neogene t i m e . p r o v i n c i a l i z a t i o n , which b e g a n i n t h e E a r l y Eocene w i t h t h e development o f a d i s t i n c t b o r e a l f a u n a , was c o m p l e t e d i n t h e M i d - P l i o c e n e w i t h t h e development o f a d i s t i n c t p o l a r f a u n a which w a s r e l a t e d t o t h e i n i t i a t i o n of n o r t h e r n hemisphere p o l a r g l a c i a t i o n 3 Ha.
3.
P l a n k t o n i c Foraminifera have been u s e f u l i n p l a c i n g v a r i o u s Paleogene a n d Neogene f o r m a t i o n s i n t h e m a r g i n a l a r e a s o f N o r t h A m e r i c a a n d Europe i n a t i m e - s t r a t i g r a p h i c framework. In s e v e r a l i n s t a n c e s t h e p l a n k t o n i c F o r a m i n i f e r a have r e s o l v e d long s t a n d i n g d i f f e r e n c e s i n age i n t e r p r e t a t i o n s as w e l l a s demonstrated t h e temporal e x t e n t of h i a t u s ( e s ) i n supposedly continuous o r quasi-continuous sequences.
4.
Paleogene benthonic f o r a m i n i f e r a l assemblages i n t h e marginal a r e a s of t h e circum-North A t l a n t i c e x h i b i t a h i g h d e g r e e of cosmopolitanism w h i c h i s somewhat r e d u c e d d u r i n g t h e Neogene a s a r e s u l t o f t h e combined e f f e c t s o f c h a n g i n g p a l e o g e o g r a p h y a n d l o w e r e d w o r l d w i d e temperatures. However, i n t r a - p r o v i n c i a l m i g r a t i o n o v e r l o n g d i s t a n c e s i s s e e n , s u c h as t h e t r a n s - A r c t i c m i g r a t i o n i n t o n o r t h e r n E u r o p e o f Arctic-Pacific faunal elements during l a t e Pleistocene i n t e r g l a c i a l intervals.
5. E v o l u t i o n a r y c o n s e r v a t i s m among b a t h y a l a n d a b y s s a l b e n t h o n i c Forami n i f e r a a p p e a r t o make p a l e o b a t h y m e t r i c a n d p a l e o c i r c u l a t i o n
404 i n t e r p r e t a t i o n s by a n a l o g y w i t h l i v i n g f a u n a s v a l i d o v e r t h e p a s t 15 my a t l e a s t . T h e s e s t u d i e s are i n t h e i r i n f a n c y a n d h o l d g r e a t promise i n paleooceanographic s t u d i e s . Bathymetric and environmental d a t a on s h a l l o w water l i v i n g b e n t h o n i c F 3 r a m i n i f e r a a r e g e n e r a l l y a p p l i c a b l e t o p a l e o e c o l o g i c s t u d i e s throughout t h e Cenozoic.
5.
X a j o r t a x o n o m i c r e p l a c e m e n t among b a t h y a l b e n t h o n i c F o r a m i n i f e r a h a s been o b s e r v e d n e a r t h e Paleocene/Eocene, Eocene/Oligocene and F a r l y / M i d d l e M i o c e n e b o u n d a r i e s , c7. 54, 38 a n d 15 Pla a g o . The c h a n g e a t 3 8 Ma c o i n c i d e s w i t h t h e e s t a b l i s h m e n t o f t h e p s y c h r o s p h e r e .
AC&:NOWLEDGE?IENTS T h i s summary was p r e p a r e d a t t h e r e q u e s t o f P r o f e s s o r F.?",.Swain f o r p r e s e n t a t i o n a t a Symposium on t h e S t r a t i g r a p h i c b l i c r o p a l e o n t o l o g y o f A t l a n t i c B a s i n a n d B o r d e r l a n d s h e l d a t t h e U n i v e r s i t y o f D e l a w a r e , 14-15 J u n e , 1 9 7 6 . I t forms p a r t of d c o n t i n u i n g r e s e a r c h program b e i n g c o n d u c t e d j o i n t l y on C e n o z o i c b e n t h o n i c f o r a m i n i f e r a by t h e Woods H o l e O c e a n o g r a p h i c I n s t i t u t i o n and t h e S o c i e t e Nationale d e s P e t r o l e d'Aquitaine (Pau, F r a n c e ) . R e s e a r c h a t Woods H o l e i s s u p p o r t e d b y a j o i , i t g r a n t f r o m t h e C h e v r o n , I am i n d e b t e d t o Drs. R . C . Marathon, N o b i l a n d S h e l l O i l Companies. T j a l s m a ( W . H . O . I . ) a n d C . W y l i e P o a g ( U . S . G e o l o g i c a l S u r v e y , Koods H o l e ) f o r a c r i t i c a l r e v i e w o f a n e a r l y d r a f t o f t h i s p a p e r a n d t o D r . J . E . van H i n t e ( C X X O X , EPR-C, B o r d e a u x ) f o r h i s comments a n d g e n e r a l i n f o r m a t i o n on ( a s y e t u n p u b l i s h e d ) f o r a m i n i f e r a 1 f a u n a s from t h e Norwegian-Greenland Sea (DSDP Leg 3 8 ) . T h i s i s Woods H o l e O c e a n o g r a p h i c I n s t i t u t i o n C o n t r i b u t i o n Number
3796.
Discussion D r . F . G r a d s t e i n : The d i s t i n c t i o n of Paleogene Midway v e r s u s Velasco f a u n a s , a s I u n d e r s t a n d i t , would r e f l e c t s h a l l o w e r v e r s u s d e e p e r water cond i t i o n s . I wonder i f s u c h a n e c o l o g i c d i s t i n c t i o n i s i n p a r t r e l a t e d t o a c e r t a i n broad l a t i t u d i n a l e f f e c t on d e p t h d i s t r i b u t i o n of s p e c i e s . Is i t p o s s i b l e t h a t Velasco f a u n a s i n h i g h e r l a t i t u d e s , t h a n where most o f your work i s d e r i v e d from, o c c u r a t a s h a l l o w e r d e p t h ? D r . W. A. Berggren: A t t h e p r e s e n t t i m e w e c a n n o t answer t h a t w i t h c e r t a i n t y . A t h i g h l a t i t u d e s i n t h e North A t l a n t i c I am n o t f a m i l i a r w i t h Velasco-type b e n t h o n i c f o r a m i n i f e r a l f a u n a s . I n t h e c i r c u m - A t l a n t i c a r e a P a l e o c e n e d e p o s i t s a r e e s s e n t i a l l y s h a l l o w w a t e r i n n a t u r e w i t h Midway t y p e f a u n a s i n them. I n t h e few i n s t a n c e s of d e e p e r w a t e r P a l e o c e n e f a u n a s t h a t I a m f a m i l i a r w i t h , t h e y a r e of a n a g g l u t i n a t e d n a t u r e (North Sea B a s i n , Norwegian S e a , e t c . ) . I would, i n p r i n c i p l e , doubt t h a t Velasco e l e m e n t s a p p e a r a t s h a l l o w e r d e p t h s i n h i g h e r l a t i t u d e s b e c a u s e I have n o t found t h e s e e l e m e n t s i n Midway f a u n a s a t h i g h l a t i t u d e s .
405 REFERENCES Adams, C . G . , 1 9 6 7 . T e r t i a r y F s r a m i n i f e r a i n t h e T e t h y a n , A m e r i c a n , a n d Adams, C . G . a n d A g e r , D . B . ( e d s . ) , A s p e c t s Indo-Pacific provinces: of Tethyan b i o g e o g r a p h y : System. A s s o c . , P u b l . 7 , p . 195-217.
I .
A k e r s , W . H . , 1 9 7 2 . F l a n k t o n i c F o r a m i n i f e r a a n d b i o s t r a t i r r a p h y o f some Neogene f o r m a t i o n s , n o r t h e r n F l o r i d a a n d A t l a n t i c C o a s t a l P l a i n : T u l a n e S t u d i e s G e o l . , P a l e o n t o l o g y , V . 9 , n o s . 1 - 4 , 1 3 9 p . . 60 p l s . B a r k e r , R.W. a n d B e r g g r e n , W . A . , i n p r e s s . P a l e o c e n e a n d E a r l y Eocene o f t h e R i o G r a n d e a n d Tampico e m b a y m e n t s : foraminiferal biostratigraphy and paleoecology: J o u r . Foram. R e s . B a r k e r , R . W . a n d Blow, W . H . , 1 9 7 6 . B i o s t r a t i g r a p h y o f some T e r t i a r y f o r m a t i o n s i n t h e T a m p i c o - M i s a n t l a embayment, M e x i c o : J o u r . Foram. R e s . , v . 6 , n o . 1, p . 3 9 - 5 8 . B a t j e s , D . A . J . , 1 9 5 8 . F o r a m i n i f e r a o f t h e O l i g o c e n e o f Belglum: Roy. S c i . N a t . B e l g . , Mem. No. 1 4 3 , 188 p . , 1 3 p l s .
Inst.
Berggren, W.A., 1971. T e r t i a r y boundaries: Micropaleontology of the O c e a n s , B.F. F u n n e l 1 a n d W . R . R i e d e l ( e d s . ) , Cambridge U n i v . P r e s s , p . 693-809. B e r g g r e n , W . A . , 1 9 7 2 . C e n o z o i c b i o s t r a t i g r a p h y and p a l e o b i o g e o g r a p h y of t h e N o r t h A t l a n t i c : & L a u g h t o n , A . S . , P , e r g g r e n , W . A . e ; a:., 1 9 7 2 , I n i t i a l R e p o r t s o f t h e Deep Sea D r i l l i n g P r o j e c t , v o l . 1 2 , W a s h i n g t o n , p . 965-1001, 1 3 p l s . B e r g g r e n , W . A . , 1 9 7 3 . The P l i o c e n e t i m e - s c a l e : c a l i b r a t i o n of p l a n k t o n i c f o r a m i n i f e r a l a n d c a l c a r e o u s n a n n o f o s s i l z o n e s : N a t u r e , v . 243. n o . 5407, p . 391-397. B e r g g r e n , W . A . , 1 9 7 4 . L a t e P a l e o c e n e - E a r l y Eocene b e n t h o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y a n d p a l e o e c o l o g y o f R o c k a l l Bank: M i c r o p a l e o n t o l o g y , v . 2 0 , n o . 4 , p . 426-448. Berggren, W.A., i n p r e s s . R e c e n t a d v a n c e s i n C e n o z o i c p l a n k t o n i c forami n i f e r a l b i o s t r a t i g r a p h y , b i o c h r o n o l o g y a n d biogeo:raptiy: Atlantic O c e a n : & S a i t o , T . a n d R i e d e l , W . R . ( e d s . ) , Ocean P l a n k t o n a n d Sediments, Micropaleontology, Spec. Publ. Berggren, W.A. and Aubert, J . , 1975. Paleocene benthonic f o r a m i n i f e r a l b i o s t r a t i g r a p h y , p a l e o b i o g e o g r a p h y and p a l e o e c o l o g y of t h e A t l a n t i c Tethyan regions: ?lidway-type f a u n a : P a l a e o p e o g r . , P a l a e o c l i m a t o l . , P a l a e o e c o l . , v . 1 8 , p . 73-192, 1 9 p l s . B e r g g r e n , W.A. and A u b e r t , J . , 1 9 7 6 a . Eocene b e n t h o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y a n d p a l e o b a t h y m e t r y o f Orphan K n o l l ( L a b r a d o r S e a ) : M i c r o p a l e o n t o l o g y , V. 2 2 , n o . 2 . B e r g g r e n , W . A . a n d A u b e r t , J . , 1976b. L a t e P a l e o g e n e ( L a t e Eocene and O l i g o c e n e ) b e n t h o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y and paleobathymetry o f R o c k a l l Bank a n d H a t t o n - R o c k a l l B a s i n : P l i c r o p a l e o n t o l o g y , v . 2 2 , no. 2 .
406 B e r g g r e n , W . A . a n d Hnq, B . , 1 9 7 6 . The A n d a l u s i a n S t a g e ( L a t e M i o c e n e ) : B i o s t r a t i g r a p h y , biochronology and palaeoecology: Falaeopeogr., P a l a e o c l i m a t o l . , P a l a e o e c o l . , v . 20. B e r g g r e n , W.A. a n d H o l l i s t e r , C . D . , 1974. P a l e o g e o g r a p h y , p a l e o b i o g e o g r a p l y a n d t h e h i s t o r y of c i r c u l a t i o n o f t h e A t l a n t i c Ocean: Hay, W . W . ( e d . ) , Sympos. on G e o l o g i c H i s t o r y of t h e O c e a n s , S O C . E c o n . P a l . M i n . , S p e c . M e m . 2 0 , p . 126-186. B e r g g r e n , W.A. a n d P h i l l i p s , J . D . , 1 9 7 1 . The i n f l u e n c e o f c o n t i n e n t a l d r i f t on t h e d i s t r i b u t i o n of Cenozoic b e n t h o n i c F o r a m i n i f e r a i n t h e C a r i b b e a n a n d i 4 e d i t e r r a n e a n r e g i o n s : 3 G r a y , C . ( e d . ) , The G e o l o g y of' L i b y a , p . 265-299, C a t h o l i c P r e s s , B e i r u t . R e r g g r e n , W . A . , E e n s o n , R . H . , Haq, B . , R i e d e l , W . R . , S a n f i l i p p o , A . , S c h r a d e r , H . J . a n d Tjalsma, R . C . , i n p r e s s . The E l C u e r v o s e c t i o n ( A n d a l u s i a , S p a i n ) : M i c r o p a l e o n t o l o g i c anatomy o f a n e a r l y L a t e Xiocene l o w e r b a t h y a l d e p o s i t . M a r i n e M i c r o p a l e o n t o l o g y , v . 1, n o . 3. d e s n o g d w e s t d e u t s c h e n Terti:r B e t t e n s t a e d t , F . , 1 9 $ 9 . Pa1:ogeographik besonderer Berucksichtigung d e r Mikropalaontologie: Erdgl u. T e k t o n i k i n N o r d w e s t d e u t s c h l a n d , p . 143-172, 1 p l .
mit
B l o w , W . H . , 1 9 6 9 . L a t e m i d d l e Eocene t o R e c e n t p l a n k t o n i c f o r a m i n i f e r a 1 b i o s t r a t i g r a p h y : I n Bronnimann, P. a n d Renz, H . H . ( e d s . ) , 1st I n t e r n a t . C o n f . P l a n k t o n i c X i c r o f o s s i l s , P r o c . , v . 1, p . 1 9 9 - 4 2 2 , 54 p l s .
B o l l i , H . M . , 1 9 5 7 . P l a n k t o n i c F o r a m i n i f e r a from t h e Oligo-Miocene C i p e r o a n d Lengua f o r m a t i o n s o f T r i n i d a d , B . W . I . : U . S . N a t . h s . B u l l . 215, p . 97-113. a r i n k m a n n , R . , 1 9 6 0 . G e o l o g i c e v o l u t i o n o f E u r o p e , H a f n e r P u b l . Co.. N . Y . , 1 6 1 p . , 46 f i g s . , 1 9 p l s . ( t r a n s l . f r o m G e r m a n ) . B r o t z e n , F . , 1 9 4 8 . The S w e d i s h P a l e o c e n e a n d i t s f o r a m i n i f e r a l f a u n a : S v e r . G e o l . U n d e r s . , S e r . C . , n o . 1, 4 9 3 (3rsb. 4 2 , n o . 2 ) , 1 4 0 p . , 19 p l s . Brown, P.M., M i l l e r , J . A . a n d S w a i n , F . M . , 1 9 7 2 . S t r u c t u r a l a n d s t r a t i g r a p h i c framework, and s p a t i a l d i s t r i b u t i o n of p e r m e a b i l i t y of t h e A t l a n t i c C o a s t a l P l a i n , N o r t h C a r o l i n a t o New Y o r k . U.S. G e o l . S u r v . P r o f . P a p . 7 9 6 , p . i i i - i v + 1 - 7 9 , 59 p l a t e s , 1 3 f i g s . 5 t a b s . C a u d r i , C . M . B . , 1 9 7 5 . G e o l o g y a n d p a l e o n t o l o g y o f S o l d a d o Rock, T r i n i d a d The L a r g e r F o r a m i n i f e r a : Ecolog. Geol. (West I n d i e s ) . P a r t 2 : H e l v . , v . 6 8 ( 3 ) , p . 5 3 3 - 5 8 9 , 30 p l s . C o u v e r i n g , J . A . Van, B e r g g r e n , W . A . , D r a k e , R . E . , A g u i r r e , E . , a n d C u r t i s , G.H., i n press. The t e r m i n a l M i o c e n e e v e n t . Mar. M i c r o p a l . , v . 1, n o . 3. Curry, D . , ASS.,
1 9 6 5 . The P a l a e o g e n e b e d s o f s o u t h - e a s t V . 76, p . 151-174.
England:
P r o c . Geol.
C u r r y , D . , 1966. Problems o f c o r r e l a t i o n i n t h e Anglo-Paris-Eelgian P r o c . G e o l . Ass., v . 7 7 , p . 437-467.
Basin:
407 Cushman, J . A . , 1 9 3 0 . The F o r a m i n i f e r a o f t h e C h o c t a w h a t c h e e F o r m a t i o n o f Florida: F l o r i d a S t a t e G e o l . S u r v . , B u l l . No. 4 . Cushman, J . A . , 1 9 3 3 . The F o r a m i n i f e r a of t h e C h o c t a w h a t c h e e F o r m a t i o n o f F l o r i d a . F l o r i d a S t a t e G e o l . S u r v . , Bull. No. 6 . Dam, A . t e n a n d R e i n h o l d , T . , 1 9 4 1 . , , Die s t r a t i g r a p h i s c h e G l i e d e r u n g d e s niederlzndischen Plio-Plistozans nach Foraminiferen: Meded. G e o l . S t i c h t . , S e r . C , V . n o . 1, 6 6 p . , 6 p l s .
D a m , A . t e n a n d R e i n h o l d , T . , 194;. Die s t r a t i g r a p h i s c h e G l i e d e r u n g d e s n i e d e r l z n d i s c h e n O l i g o - M i o z a n s n a c h F o r a m i n i f e r e n . Meded. G e o l . S t i c h t . , S e r . C , V , n o . 2 , 1 0 6 p . , 10 p l s . E a m e s , F . E . , B a n n e r , F . T . , Blow, W . H . , a n d C l a r k e , W . J . , 1 9 6 2 . F u n d a m e n t a l s o f m i d - T e r t i a r y s t r a t i g r a p h i c a l c o r r e l a t i o n , Cambridge Univ. P r e s s , 163 p . , 1 7 p l s .
Emery, K . O . a n d U c h u p i , E . , 1 9 7 2 . W e s t e r n N o r t h A t l a n t i c O c e a n : Topog r a p h y , R o c k s , S t r u c t u r e , W a t e r , L i f e a n d S e d i m e n t s . Am. A s s o c . P e t r o l . G e o l . Mem. 1 7 : 532 p . Gates, W . L . , 1976. p 1138-1144.
.
M o d e l i n g t h e Ice-Age
climate:
S c i e n c e , v . 1 9 1 , n o . 4232,
Hansen, H . J . , 1970. D a n i a n F o r a m i n i f e r a f r o m N u g s s u a q , West G r e e n l a n d . X e d d e l . G r d n l a n d , v . 1 9 3 , n o . 2 , 132 p . , 33 p l s . H e d b e r g , H . , 1 9 7 6 . Ocean b o u n d a r i e s a n d p e t r o l e u m r e s o u r c e s . V . 1 9 1 , p . 1009-1018.
Science,
Hazel, J . E . , i n press. D i s t r i b u t i o n o f some b i o s t r a t i g r a p h i c a l l y d i a g n o s t i c o s t r a c o d e s i n t h e P l i o c e n e a n d P l e i s t o c e n e of V i r g i n i a a n d n o r t h e r n N o r t h C a r o l i n a . U.S. G e o l . S u r v e y , J o u r n . R e s . , V . 5 . H i n t e , J . E . v a n a n d Wong, T . , 1 9 7 5 , . ' h w m L ~ % e srccka/ZensI;s n . s p . f r o m t h e Upper P a l e o c e n e o f R o c k a l l P l a t e a u ( N o r t h A t l a n t i c ) . J o u r . Foram. R e s . , v. 5 , n o . 2, p . 90-101, 3 p l s . Hooyberghs, H . J . F . a n d de N e u t e r , F . J . C . , 19 7 2 . B i o s t r a t i g r a p h y and i n t e r r e g i o n a l c o r r e l a t i o n of t h e "Miocene" d e p o s i t s o f n o r t h e r n B e l g i u m b a s e d o n p l a n k t o n i c f o r a m i n i f e r a ; t h e O l i g o c e n e - M i o c e n e b o u n d a r y on t h e s o u t h e r n e d g e o f t h e N o r t h S e a B a s i n : Pieded. K o n i n k l . Acad. W e t e r s c h . , L e t t . S c h o n e K u n s t e n Bel.g., K1. W e t e n s c h . , v . 3 4 , n o . 3 , 4 7 p . , 11 pls. K a a s c h i e t e r , J . P . H . , 1 9 6 1 . F o r a m i n i f e r a o f t h e Eocene o f Belgium: Roy. S c i . Nat. B e l g . , M e m . No. 1 4 7 , 2 7 1 p . , 1 6 p l s .
Inst.
K e n n e t t , J . P . a n d S h a c k l e t o n , N . J . , 1 9 7 6 . Oxygen i s o t o p e e v i d e n c e f o r t h e d e v e l o p m e n t o f t h e p s y c h r o s p h e r e 3 8 Myr a g o : N a t u r e , v . 2 6 0 , p . 513-515. Kugler, H.G. and Caudri, C.M.B., 1975. Geology and p a l e o n t o l o g y o f S o l d a d o Rock, T r i n i d a d (West I n d i e s ) . P a r t 1: G e o l o g y a n d b i o s t r a t i g r a p h y : E c l o g . G e o l . H e l v . , V . 6 8 ( 2 ) , p . 365-430, 2 p l s .
408 L a r s e n , G . and D i n e s e n , A . , 1959. V e j l e F j o r d Formationen ved B r e j n i n g : s e d i m e n t e r n e og f o r a m i n i f e r f a u n a e n (Oligocaen-Miocaen): Danm. G e o l . U n d e r s . , v . 2 , no. 8 2 , 114 p . , 9 p l s . L a u g h t o n , A . S . , 1 9 7 5 . T e c t o n i c e v o l u t i o n of t h e N o r t h e a s t A t l a n t i c Ocean: a r e v i e w : Y o r g e s . G e o l . U n d e r s . , V . 316, p . 169-193. L o e b l i c h , A . R . , J r . a n d Tappan, H . , 1 9 5 7 . P l a n k t o n i c f o r a m i n i f e r a of P a l e o c e n e a n d E a r l y E o c e n e a g e s f r o m t h e G u l f and A t l a n t i c C o a s t a l B u l l . U.S. Nat. X u s . 2 1 5 , p . 1 7 3 - 1 9 8 , p l s . 40-64. Plains: Lohmann, G.P. a n d T j a l s m a , R . C . , 1 9 7 5 . G e o g r a p h i c a n d b a t h y m e t r i c d i s t r i b u t i o n of P a l e o c e n e deep-water b e n t h i c f o r a m i n i f e r a from t h e w e s t e r n A t l a n t i c Ocean: A b s t r a c t s , B e n t h o n i c s ' 7 5 , p . 27, H a l i f a x . Lohmann, G . P . a n d T j a l s m a , R . C . , i n p r e s s . P a l e o c e n e d e e p w a t e r b e n t h o n i c f o r a m i n i f e r a f r o m t h e w e s t e r n A t l a n t i c O c e a n . P a r t 11: G e o g r a p h i c and b a t h y m e t r i c d i s t r i b u t i o n . Maher, J.C., 1 9 7 1 . G e o l o g i c f r a m e w o r k a n d p e t r o l e u m p o t e n t i a l o f t h e A t l a n t i c Coastal P l a i n and C o n t i n e n t a l S h e l f . U.S. Geol. Surv. P r o f . Pap. 659, p . i i i - i v + 1-98, 1 7 p l s . , 8 f i g s . 4 t a b s . P I a r g e r e l , J . P . , 1 9 6 8 . Les F o r a m i n i f g r e s du R e d o n i e n . N a n t e s , 207 p . , 44 p l s .
Thgse d o c t . , Univ.
E l c I n t y r e , A . , N o o r e , T . C . 2 ; J:., 1 9 7 5 . The s u r f a c e o f t h e Ice-Age S c i e n c e , V . 1 9 1 , n o . 4232, p . 1131-1137.
earth:
Murray, G . E . , 1 9 6 1 . Geology o f t h e A t l a n t i c and Gulf C o a s t a l P r o v i n c e o f S o r t h A m e r i c a , H a r p e r a n d B r o t h e r s , New Y o r k , p . v - x v i i + 1-692. M u r r a y , J.W. a n d W r i g h t , C . A . , 1 9 7 4 . P a l a e o g e n e F o r a m i n i f e r i d a a n d p a l a e o e c o l o g y , Hampshire and P a r i s B a s i n s a n d t h e E n g l i s h Channel: S p e c . P a p . P a l a e o n t o l . No. 1 4 , P a l a e o n t o l . A s s o c . L o n d o n , 1 7 1 p . , 20 p l s . 1 9 6 0 . F o r a m i n i f e r a o f l a t e s t C r e t a c e o u s and e a r l i e s t T e r t i a q a g e i n t h e New J e r s e y C o a s t a l P l a i n : J o u r . P a l e o n t o l o g y , v . 34, p . 1-59.
Olsson, R . K . ,
Olsson, R . K . , 1970a. P a l e o c e n e p l a n k t o n i c f o r a m i n i f e r a 1 b i o s t r a t i g r a p h y a n d p a l e o z o o g e o g r a p h y o f PJew J e r s e y : J o u r . P a l e o n t o l o g y , v. 44, p . 589-597. Olsson, R . K . , 1 9 7 0 b . P l a n k t o n i c F o r a m i n i f e r a f r o m t h e b a s e o f t h e T e r t i a r y , ? l i l l e r ' s F e r r y , Alabama: J o u r . P a l e o n t o l o g y . v . 4 4 , p . 598-604. P i t m a n , K . C . a n d T a l w a n i , M., 1 9 7 2 . S e a - f l o o r s p r e a d i n g i n t h e N o r t h Atlantic: G e o l . S O C . America B u l l . , v . 8 3 , p . 619-646. Pomerol, C . , 1973. S t r a t i g r a p h i e e t P a l g o g g o g r a p h i e : ( T e r t i a r e e t Q u a t e r n a i r e ) , D o i n , P a r i s , 269 p .
;re
CGnozoique
P o o r e , R . Z . and B e r g g r e n , W . A . , 1975. L a t e Cenozoic p l a n k t o n i c f o r a m i n i f e r a l b i o s t r a t i g r a p h y and p a l e o c l i m a t o l o g y o f t h e n o r t h e a s t e r n Atlantic: DSDP S i t e 1 1 6 : J o u r . Foram. R e s . , v . 5 . n o . 4 , p . 270-293.
409 P u r i , H.S., 1 9 5 3 . C o n t r i b u t i o n t o t h e s t u d y o f t h e X i o c e n e o f t h e F l o r i d a Foraminifera, Panhandle: F l o r i d a Geol. Surv. B u l l . KO. 36, P t . 2 : p . 71-213, 30 pis. R u t t e n , M . G . , 1 9 6 9 . The g e o l o g y o f W e s t e r n E u r o p e , Amsterdam, 520 p .
E l s e v i e r Publ. Co.,
Savin, S . M . , Douglas, R . G . , and S t e h l i , F.G., 1975. T e r t i a r y marine paleotemperatures: Geol. Soc. America, v . 8 6 , p . 1499-1510. S t a e s c h e , K . a n d H i l t e r m a n n , H . , 1 9 4 0 . M i k r o f a u n e n aus dem T e r t i ' i r Nordwestdeutschlands: Abh. R e i c h s a n s t . B o d e n f o r s c h . , N.F. B e r l i n , n o . 2 0 1 , 26 p . , 51 p l s . S t a i n f o r t h , R . X . , 1965. ?iid-Tertiary diastrophism i n n o r t h e r n South America: F o u r t h C a r i b b e a n G e o l . Conf, T r i n i d a d , p . 159-174. l a l w a n i , Y., U d i n t s e v , G . t?: a:., 1 9 7 5 . G e o t i m e s , v . 2 0 , n o . 2 , p . 24-26.
Leg 38 - Deep S e a D r i l l i n g P r o j e c t :
Tjalsma, R . C . , 1 9 7 1 . S t r a t i g r a p h y a n d F o r a m i n i f e r a o f t h e S e o g e n e o f t h e Eastern Guadalquivir Basin (southern Spain): Utrecht Nicropaleontol. B u l l . , V . 4 , p . 1-161. Tjalsma, R . C . a n d Lohmann, G . P . , 1 9 7 5 . Taxonomy a n d d i v e r s i t y o f P a l e o c e n e deep water b e n t h o n i c F o r a m i n i f e r a from t h e W . A t l a n t i c : A b s t r a c t s , Benthonics '75, p . 48, H a l i f a x .
T j a l s n ? , R . C . a n d Lohmann, G . P . , i n p r e s s . P a l e o c e n e d e e p w a t e r b e n t h o n i c F a r a m i n i f e r a f r o m t h e w e s t e r n A t l a n t i c O c e a n . P a r t I : Taxonomy. U l l e b e r g , K . , 1 9 7 4 . F o r a m i n i f e r a and s t r a t i g r a p h y o f t h e Viborg Formation i n S o f i e n l u n d , Denmark: B u l l . G e o l . S o c . Denmark, v . 2 3 , p . 2 6 9 - 2 8 2 . V e r d e n i u s , J . G . , 1 9 7 0 . S e o g e n e s t r a t i g r a p h y of t h e w e s t e r n G u a d a l q u i v i r Basin (southern Spain): U t r e c h t X i c r o p a l a e o n t o l . B u l l . , v . 3, p . 1-109. V o o r t h u y s e n , J . H . v a n , 1 9 5 8 . Les F o r a m i n i f z r e s M i o - P l i o c s n e s a i r e s du K r u i s s c h a n s : I n s t . Roy. S c i . S a t . B e l g . , N 6 m . 34 p . , 10 p l s .
e t QuaternNo. 1 4 2 ,
Woodland, A . W . ( e d . ) , 1 9 7 5 . P e t r o l e m a n d t h e c o n t i n e n t a l s h e l f o f n o r t t i w e s t E u r o p e , v . 1, G e o l o g y , H a l . s t e d P r e s s , J . W i l e y a n d S o n s , I n c . , New Y o r k , 5 0 1 p . W r i g h t , C . A . a n d M u r r a y , J.W., 1 9 7 2 . C o m p a r i s o n s o f modern a n d P a l e o g e n e f o r a m i n i f e r a 1 d i s t r i b u t i o n s and t h e i r environmental i m p l i c a t i o n s . M6m. B u r . R e c h . G g o l . ? ; i n . , v . 7 9 , p . 8 7 - 9 6 .
ICELAND-FAEROES (336, 352)
I C E L A N D PLATEAU (348)
J A N MAYEN R I D G E
V @ R I N C PLATERL
(346, 3 4 7 , 3 4 9 )
(344)
I s l a n d i e l l a spp. E b i c i d e s spp . Pullenia bulloides -~ Pullenia zinqueloba Nonion z a n d a m a e E l p h i d i m incertum rniliolids
Planulina w u e l l e r st o r f i
Planulina w u e l l e r s torfi Bu 1i m i n a a c u 1e a t a Islandiella
RIDGE
__
L
P l a n u l i n_ ~ a wuellerstof i I s 1a n d i e 11a
n o r crossi
L Z ------
E __
L
B A R R E N
lar ti n o t i e 1l a -
communis
M
__ S p i r 0 si g m o i 1-
inella _-
Splr o l o c ammi n i
E
_ C_y _ clammi __ n aP s amos p l i a e r a C r i b r o s torno i de s
B A R R E N
Table 9
D o m i n a n t b e n t h o n i c f o r a m i n i f e r a 1 e l e m e n t s i n Neogene a s s e m b l a g e s of t h e Norwegian-Greenland S e a .
_
411
PALEOGENE FORAMINIFERA - SOUTH ATLANTIC Alwine Bertels Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires , Rephblica Argentina. Consejo Nacional de Investigaciones Cientif icas y Tkcnicas , RepAblica Argentina Resumen En el presente trabajo de efectda el anAlisis de las asociaciones foraminiferolhgicas que prevalecieron durante el Paleogeno en el Atlintico Sur. Se compendian 10s resultados, hasta ahora obtenidos, sobre 10s conjuntos faunisticos haliados en las distintas cuencas sedimentarias, tanto en el continente sudamericano como en el africano. I I Los conjuntos planctonicos y bentonicos del Paleoceno alcanzan una gran distribuci6n areal en el AtlLntico Sur. Durante el Eoceno se observa una diferenciaci6n faun:stica latitudinal mas restricta que durante el Paleoceno; por otra parte, en latitudes mayores a 1 0 s 3OP Sur las influencias australoI asiaticas son marcadas, l a s cuales persisten durante el Oligoceno. Abstract The several recorded foraminifera1 assemblages from the South American and African continents and those of the South Atlantic Ocean are summarized in the present work. Paleocene planktonic and benthonic assemblages have a wide Atlantic areal distribution. During Eocene times foraminifera1 assemblages show more restricted latitudinal differentiations than in the Paleocene and, at latitudes higher than 30' S Australasian influences are evident, which persist throughout the Oligocene. Introduction The purpose of the present work is to show in an approximate integral form the foraminiferal assemblage which prevailed in the South Atlantic Ocean during Paieogene times. To accomplish this purpose all sedimentary basins adjacent to the South Atlantic Ocean are analyzed for the Paleogene Period; chronostratigraphic and geographic order are followed; this analysis is made from northernto southern latitudes in both the South American and African continents.
412
The a v a i l a b l e d a t a o b t a i n e d from s e v e r a l o c e a n o g r a p h i c exp e d i t i o n s , p r o v i d e d a l s o a g r e a t amount of i n f o r m a t i o n concerned with t h e marine sedimentary sequences i n t h e A t l a n t i c Ocean; t h e s e d a t a a r e s y n t h e s i z e 6 and t h e p r i n c i p a l p l a n k t o n i c Zones a r e m e n t i o n e d . The d a t a f o r t h i s work were t a k e n from a l l t h e a v a i l a b l e l i t e r a t u r e and a l s o from u n p u b l i s h e d t h e s e s d e a l i n g w i t h t h e same t o p i c . I t i s a c c e p t e d i n t h i s work t h a t t h e P a l e o g e n e b e g a n
a p p r o x i r r a t e l y 6 5 m.y.
a g o , and t h a t a t t h i s t i m e t h e A t l a n t i c
Ocean was d e f i n i t i v e l y o u t l i n e d .
Throughout t h e T e r t i a r y , s e a
f l o o r s p r e a d i n g , accompanied by t h e d r i v e of t h e S o u t h American and A f r i c a n t e c t o n i c p l a t e s c o n t i n u e d t o o p e r a t e . Acknowledgments. The a u t h o r w i s h e s t o e x p r e s s g r a t i t u d e t o t h e A r g e n t i n a N a t i o n a l C o u n c i l f o r S c i e n t i f i c and T e c h n i c a l R e s e a r c h , f o r e c o n o m i c a l a i d i n t h e r e a l i z a t i o n of t h e p r e s e n t w o r k , and , f o r g e n e r o u s l y p r o v i d i n g u s e o f t h e J e o l c o J.S.M.-U/3 s c a n n i n g e l e c t r o n m i c r o s c o p e w i t h w h i c h t h e m i c r o g r a p h s were t a k e n . The w r i t e r i s a l s o g r a t e f u l t o t h e Buenos A i r e s U n i v e r s i t y , Department of Geology, f o r t h e provided f a c i l i t i e s f o r a c h i e v i n g t h e c o m p l e t i o n of t h e work. P a r t i c u l a r thanks are a l s o expressed t o D r . C a r l o s A. R i n a l d i and t o M r .
P a t r i c i o G a n d u g l i a , who k i n d l y and g e n e r o u s -
l y h e l p e d i n t h e c o n s t r u c t i o n o f t h e s t r a t i g r a p h i c c h a r t s and r a n g e c h a r t s which i l l u s t r a t e t h e p a p e r . The i l l u s t r a t e d m a t e r i a l i s d e p o s i t e d i n t h e L a b o r a t o r y of Micropaleontology, Facultad d e Ciencias Exactas y N a t u r a l e s , U n i v e r s i d a d d e Buenos A i r e s , A r g e n t i n a . P a l e o g e n e of S o u t h America Paleocene Along t h e p r e s e n t S o u t h A t l a n t i c S o u t h American b o r d e r l a n d s and t h e i r c o n t i n e n t a l margins Paleocene s t r a t a a r e d e v e l oped i n t h e Foz d e Amazonas, S ~ O Luiz/Barreirinhas, B r a z i l and i n t h e R e c i f e / J o $ o
Pessoa
c a v o and E s p i r i t o S a n t o b a s i n s ,
,
Segipe/Alagoas
northern
,
Rocon-
l o c a t e d i n n o r t h e a s t e r n and
413
eastern Brazil. In Argentina, Paleocene sequences have been recognized in the Colorado, NeuquAn, San Jorge and Austral basins; it is necessary to remark that the Austral Basin, in a northwest-southeast trend, extends into the southern region of the Chilean Republic. In the Brazilian literature there is controversy about the interpretation of the extent, limits and nomenclature of several of the sedimentary basins; this lack of agreement is the result of many difficulties in interpreting the results of the tectonism which has been active along the present Atlantic coast of Brazil since the late Jurassic. For instance, AmapA State, Maraj; Island and Par; State, which were once considered by several authors to lie in independent basins, are included now in the single Foz de Amazonas or Amazonas Basin. From the sedimentary Foz de Amazonas Basin Schaller, Vasconcelos and Castro (1971) recognized the Limoeiro, Amap; and Maraj; Formations of Paleocene age. Based on palynological data the age of the Limoeiro Formation ranges from late Cretaceous to Paleocene. It rests upon the Precambrian crystalline basement or on Paleozoic remnants. The sequence is composed of sandstones and several conglomeratic levels; the sandstones contain intercalated claystones. Its origin is partly fluviatile and partly marine. No Foraminifera havebeen recorded from this formation. The Limoeiro Formation is overlain in this basin by the I Marajo Formation which on lithological features was subdivided by Schaller, Vasconcelos and Castro (op. cit.) into four members (Text Fig. 1 ) . The formation lacks foraminifera1 faunas. I I I The Amapa Formation crops out in the Para and Amapa States and occurs in their continental margins; it is mainly composed of limestones with intercalated claystones and siltstones, often glauconitic and pyritic. The age of the Amap; Formation ranges from Paleocene to the Miocene. Although some Forariiinifera are present, they are poorly represented and not clearly indicative of Paleocene age. The S%o Luiz/Barreirinhas Basin contains marine sedimentary sequences which range in age from the Aptian to the Tertiary. Although a palynological zonation was made (Lima, 19731, there
414
are no foraminifera1 records from this part of the basin. From the continental margin of the Szo Luiz/Barreirinhas Basin the Globorotalia pseudobulloides Zone was recognized by Noguti (1975). Paleocene sections become more complete in northeastern Brazil whereas in southeastern regions the Paleocene is absent. In the Recife/Jo$o Pessoa Basin Tinoco (1967) and Mabesone, Tinoco and Coutinho (1968) mentioned the Paleocene Maria Farinha Formation which transitionally overlies the Upper Cretaceous Gramame Formation. The exposed section of the Maria Farinha Formation may represent B regressive facies; it is composed of clastic limestones, finer at the base and coarser to the top. A disconformity separates the Maria Farinha Formation from the overlying continental sequence in the region of the Barreiras Group of Pliocene or younger age. From the cited fauna and the absence of Globoconusa daubjergensis (Bronnimann) the authors (Mabesone &. , 1968) inferred that part of the Danian is not well represented. Taking into account the cited planktonics, although
e-
=-
boconusa daubjergensis is absent as well as Globigerina large part of the Danian and Montian stages may be recognized. In northeastern Brazil Schaller (1969) studied sedimentary sequences of the Sergipe/Alagoas Basin and the Reconcavo/Tucano r and Jatoba basins. These basins are separated by a horst over which undifferentiated Cretaceous exists. The basement of these basins consist of Precambrian and Paleozoic rocks, represented by gneisses, migmatites, metasedimentites, intrusive, extrusive and hypabyssal rocks. In the Sergipe/Alagoas Basin continuous marine sections of Aptian to Lower Eocene age are present. The Paleocene Muribeca Formation was studied by Schaller (1969); this author recognized two zones: Z 117 and Z 118. Z 117 is based on Chiloguembelina morsei (Kline), Globoconusa daubjergensis (Bronnimann) Globorotalia compressa (Plummer) Globorotalia pseudobulloides (Plummer) and Globorotalia perclara Loeblich and Tappan; Zone 118 is proposed on the content
bins, a
415 m
of Globorotalia anqulata (White), Globorotalia velascoensis
- 0
.
N60LA
I-
~~
JULIANO
LEOYIAWO
ISUBSURFACEJ
-7-p-
LNTRERRIANO AND/OR PARANIAND
OLIGOCENE
BENIN
Fm
1
AKOSSO
WIN? S T I S U
ARGENTINA
o
(Cushman) and Globorotalia acuta (Toulmin)
Text Fig. 1. Tertiary stratigraphic units from South America and Africa.
416
Noguti and Fontana dos Santos (1972) and Noguti (1975) recognized in the Sergipe/Alagoas Basin and in its prolongation on the continental margin the Paleocene zones of: Globoconusa daubjergensis and Globorotalia pseudobulloides in descending order of age, whereas in the extension of the basin into the continental margin the same authors proposed additionally: the pseudomenardii Globorotalia pusilla pusilla, Globorotalia (5.) and Globorotalia ( G . ) velascoensis zones in descending order. In the continental margin of the Reconcavo Basin, Noguti (1975) recognized the Paleocene Globorotalia pusilla pusilla Zone. The Espirito Santo Basin yielded diagnostic planktonics; Noguti and Fontana dos Santos (1972) and Noguti (1975) proposed the Globorotalia pseudomenardii and Globorotalia velascoensis zones in ascending order. In Argentina the oldest Tertiary sediments were deposited in the NeuquAn, Colorado, San Jorge and Austral basins. Danian strata were recorded from subsurface of the Colorado Basin by Kaasschieter (1963) and Malumian (1970b). -After the deposition of a thick terrestrial sequence of the Upper Cretaceous Neuquhn Group, the Danian sea invaded the basin and deposited the Pedro Luro Formation, This formation is only recognized in subsurface; it is mainly composed of clayey to marly sediments. The sediments yielded planktonic and benthonic microfauna. Based on the presence of Globorotalia pseudobulloides (Plummer) and Globorotalia compressa (Plummer) a Danian age was assigned to these strata (Kaasschieter, 1963). Bertels (1964; 1969) studied the foraminifera1 content of the Roca Formation and recognized the Cretaceous-Tertiary boundary over a large area in northern Patagonia, i.e. the Neuqu&n and Colorado basins, The Lower Tertiary strata are represented by the Roca Formation which in the type area (NeuquAn Basin) is of early Danian age based on the occurrence of Globoconusa daubjergensis (Bronnimann), Globorotalia pseudobulloides (Plummer) and Subbotina triloculinoides (Plummer) Benthonic Foraminifera are also abundant. The Roca Formation is composed Of clays at
.
417
the base grading into marls and limestones to the top indicating a regressive facies (Bertels, 1970). Bertels (1975) based on the occurrence of Globorotalia pseudobulloides (Plummer) recognized that Zone in the Roca. In the San Jorge Basin Danian sedimentary sequences extend over large areas which shows lateral facial changes; the strata are exposed from the coast to the central region of the Chubut Province. These strata constitute the Salamanquiano Stage. The known formations are the Salamanca and Boror6 formations. The first micropaleontological studies in this basin were carried out by Camacho (1949) with subsurface samples; subsequent studies of this formation were undertaken by Mendez (1966), Masiuk (1967), and Bertels (1973, 1976a) based on surfaced sections. Mendez (1966) stLdied outcrop samples from the Salamanca Formation at Punta Peligro locality (Chubut Province). The assigned age was Middle to Upper Danian. Masiuk (1967) analyzed outcrop samples located at Puesto Alvarez, on the lower course of the Rio Chico River; the assigned age was Middle to late Upper Danian for this sequence. Bertels (1973) I studied the microfaunal assemblage of the Cerro Bororo Formatiofi which crops out in the central area of the Chubut Province. Based on the presence of Globorotalia compressa (Plummer) a late Upper Danian age was suggested for the Cerro BororA Formation. Bertels (1976) studied the microfaunal content of the sedimentary sections of the Salamanca Formation at the type section of Bajada de Hansen locality (Andreis, Mazzoni and Spalletti, 1975). The micropaleontological content comprises planktonic and benthonic Foraminifera. The most conspicuous planktonic is Globorotalia, (Turborotalia) compressa (Plummer), and Bertels (1975) proposed the zone of Globorotalia (Turborotalia) compressa for strata of the upper or Hansen Member of the Salamanca Formation. Benthonic Foraminifera constitute rich and diagnostic assemblages. The Austral Basin is the southernmost basin and extends in a northwest-southwest trend comprising the southern regions of
418
Chile and Argentina. In subsurface samples of the Austral Basin Malumian, Masiuk and Riggi (1974) found a more or less continuous sequence of strata which range in age from Valanginian to Miocene. Danian assemblages were recorded by those authors. The species found in the Austral Basin are basically the same as those recorded from northern areas. In the Chilean part of the Austral Basin Brunswick Peninsula,Paleocene strata were studied by Herm (1966) and Charrier and Lahsen ( 1 9 6 9 ) . From the southern Chile Brunswick Peninsula area, Charrier and Lahsen (1969) mentioned Upper Cretaceous and Lower Tertiary sediments. These authors included in the Paleocene the upper part of the Chorrillo Chico Formation, the San Jorge Formation and the lower part of the Agua Fresca Formation. The Chorrillo Chico Formation consists of very hard, light-brown, silty,partly glauconitic argillite of tuffaceous appearance. Large lenticular limestone concretions are characteristic of the formation which yielded species of the genera gclammina and Allomorphina, etc. Charrier and Lahsen concluded an Upper Cretaceous-Lower Tertiary age for this formation. The San Jorge Formation consists of poorly fossiliferous, glauconitic, Drown argillite containing many limestone concretions. Charrier and Lahsen (1969) suggested a Paleocene age for these strata. Eocene The Eocene is represented in the Foz de Amazonas, Sergipe/ Alagoas, Reconcave, Espirito Santo, Campos (Rio de Janeiro) and Paran: Basin in Brazil, and in the Argentina Colorado, San Jorge and Austral Basins. The northernmost basin is the Foz de Amazonas Basin. This basin comprises an emerged area (=sedimentary MarajA Basin) and a second situated on the continental margin; the basin is limited by the Guayanas Shield at the northeast and the Central 1 Brazilian Shield at the southwest. The Gurupa arc separates this basin from the Paleozoic basin in the "Lower" Amazonas and the Tocantins arc separates it from the Maranhao sedimentary basin (Schaller, Vasconcelos and Castro, 1971). Forming part of the Foz de Amazonas Basin are the AmapA and Par; regions.
419
In the Amapi Formation several sequences have been recognized in which the following planktonic foraminifera1 zones were recorded: Globorotalia wilcoxensis Zone below and Truncorotaloides rohri Zone above from the lower Candiru sequence. The Jacundi sequence (Upper Eocene-Oligocene) yielded the zonal species: Globorotalia cerroazulensis and Globorotalia opima opima. Schaller ( 1 9 6 9 ) in the Sergipe/Alagoas Basin -found the Eocene planktonics Globanomalina aff. pseudoiota Hornibrook and Globorotalia aff. 5. wilcoxensis Cushman and Parker, Noguti and Fontana dos Santos ( 1 9 7 2 ) and Noguti ( 1 9 7 5 ) recorded from this basin the planktonic zones of: Globorotalia wilcoxensis, Truncorotaloides rhori, and Globorotalia cerroazulensis, in descending order of age. Noguti and Fontana dos Santos ( 1 9 7 2 ) and Noguti ( 1 9 7 5 ) based on studies of sedimentary sequences of the Sergipe/ Alagoas Basin continental margin recognized in ascending
--
2rder of age the Zones of: Globorotalia wilcoxensis, Globorotalia quetra, Globorotalia palmerae, Globigerinoides Orbulinoides beckmanni, Truncorotaloides rohri, Globigerapsis semiinvoluta, and Globorotalia cerroazulensis. From the Reconcavo Basin (South of Bahia) Noguti ( 1 9 7 5 ) recorded the planktonic Zones of: Globorotalia wilcoxensis, Globorotalia guetra, Globigerinoides higginsi, Truncorotaloides rohri, and Globorotalia cerroazulensis, in ascending order. In the Espirito Santo Basin Noguti and Fontana dos Santos ( 1 9 7 2 ) and posteriorly Noguti ( 1 9 7 5 ) recognized the following Zones: Globorotalia wilcoxensis, Globorotalia quetra, G l o r bigerinoides higginsi, Orbulinoides beckmanni, Truncorotaloides rohri, and Globorotalia cerroazulensis, in ascending order. In the Campos Basin (Rio de Janeiro) Noguti ( 1 9 7 5 ) recognized in ascending order: Globorotalia wilcoxensis Zone, Gborotalia guetra Zone, Orbulinoides beckmanni Zone, Truncorotaloides rohri Zone. In the continental margin of the Parani Basin Noguti ( 1 9 7 5 ) recognized the Globorotalia cerroazulensis Zone. From subsurface sequences of the Austral Basin, Santa Cruz Province, Argentina, Malumian (19ES) found an Eocene fauna with marked affinities with that of the Aqua Fresca Formation in which foraminiferal content was previously studied from the
--
m,
420
C h i l e a n r e g i o n o f t h i s b a s i n by Todd and. Kniker ( 1 9 5 2 ) . Todd and Kniker (1952) r e c o r d e d Eocene F o r a m i n i f e r a from t h e Agua F r e s c a F o r m a t i o n of M a g a l l a n e s P r o v i n c e , S o u t h e r n Chile.
N e v e r t h e l e s s , t h e a g e i s P a l e o c e n e t o e a r l y Eocene
b a s e d on t h e p l a n k t o n i c s found i n s u r f a c e and s u b s u r f a c e samp l e s such a s :
G l o b i g e r i n a t r i l o c u l i n o i d e s Plummer, G l o b o r o t a l -
-
i a compressa (Plummer), G l o b i g e r i n a a q u i e n s i s L o e b l i c h and
Tappan, G l o r o t a l i a membranacea ( E h r e n b e r g ) , A c a r i n i n a t r i p l e x S u b b o t i n a , Globanomalina p s e u d o i o t a Hornibrook and Pseudoh a s t i g e r i n a w i l c o x e n s i s Cushman and P o n t o n . S u b s e q u e n t s u b s u r f a c e s t u d i e s i n t h e A u s t r a l B a s i n were c a r r i e d o u t by Malumian, Masiuk, and R i g g i ( 1 9 7 4 ) ; t h e s e aut h o r s r e c o r d e d T r u n c o r o t a l o g c o l l a c t e a ( F i n l a y ) and a s s i g n e d a l a t e Middle Eocene t o e a r l y l a t e Eocene a g e t o t h i s i n t e r v a l . O u t c r o p s samples of t h e San J u l i i n F o r m a t i o n ( A u s t r a l B a s i n ) y i e l d e d a f o r a m i n i f e r a 1 a s s e m b l a g e i n which t h e p l a n k t o n i c zones o f :
G l o b i g e r i n a eocaena ( l a t e Eocene) and
%-
b i g e r i n a o f f i c i n a l i s ( e a r l y O l i g o c e n e ) were p r o p o s e d ( B e r t e l s , 1975).
I
The San J u l i a n F o r m a t i o n i s of l a t e Eocene t o e a r l y
O l i g o c e n e a g e ( B e r t e l s 1975; 1 9 7 6 ~ ) . Oligocene The O l i g o c e n e i s m a i n l y r e c o r d e d from s u b s u r f a c e samples from t h e c o n t i n e n t a l m a r g i n of B r a z i l . I t i s r e p r e s e n t e d i n t h e Foz d e Amazonas B a s i n and i t s
c o n t i n e n t a l margin.
From t h e Amap; F o r m a t i o n , i n i t s Tambaqu: Sequence, S c h a l l e r , V a s c o n c e l o s and C a s t r o ( 1 9 7 1 ) r e c o r d e d t h e G l o b i g e r i n a c i p e r o e n s i s c i p e r o e n s i s Zone.
I n the continental
m a r g i n from t h e Foz d e Amazonas B a s i n Noguti and F o n t a n a dos S a n t o s ( 1 9 7 2 3 and N o g u t i ( 1 9 7 5 ) r e c o r d e d t h e :
Globigerina
a m p l i a p e r t u r a Zone, G l o b i g e r i n a opima opima Zone, and t h e SGlobigerina c i p e r o e n s i s c i p e r o e n s i s Zone, i n a s c e n d i n g o r d e r of age, Noguti and F o n t a n a d o s S a n t o s ( 1 9 7 2 ) and Noguti (1975) r e c o r d e d from t h e c o n t i n e n t a l m a r q i n of t h e S e r g i p e / A l a g o a s B a s i n t h e O l i g o c e n e zones o f :
--
Globigerina ampliapertura
G l o b o r o t a l i a opima opima, and G l o b i g e r i n a c i p e r o e n s i s c i p e r o e n s i s , i n ascending order.
-
N o g u t i and F o n t a n a d o s S a n t o s
421
(1972) and Noguti (1975) from the continental margin of the Reconcavo and Espirito Santo basins recorded, in ascending order, the zones of: Globigerina ampliapertura, Globorotalia opima opima, Globigerina ciperoensis ciperoensis, and Globorotalia kugleri. In the Campos Basin (Rio de Janeiro) the zones of Globigerina ampliapertura and Globorotalia were recognized by Noguti (1975). The continental margin of the P a r a d Basin (Noguti, 1975) yielded fossils of the Oligocene Globigerina ampliapertura Zone. In Argentina, subsurface samples from the Colorado Basin yielded planktonic Foraminifera that are of Oligocene age (Malumian, 1972). Benthonic Foraminifera are also abundant in the Oligocene section. From the Austral Basin Bertels (1975, 1976b) mentioned planktonic and Oenthonic assemblages from the Oligocene type I Monte Leon Formation and the Leoniano Stage. Lithologically the Monte Le& Formation is composed of siltstones with a large pyroclastic content; at the top arenaceous sediments intergrade gradually with the non-marine Santa Cruz Formation. The Leonian Stage is also represented in subsurface in this basin and in the San Jorge (surface) and Colorado (subsurface) basins. The microfaunal assemblage yielded planktonic and benthonic Foraminifera; for the Leonian Stage Bertels (1975) proposed three planktonic foraminifera1 zones in ascending order: i) Zone of Globigerina anguliofficinalis, ii) Zone of Globiger-
ina ciperoensis,
iii) Zone of Globigerina angulisuturales. The benthonic Foraminifera are also well represented. Subsurface Oligocene sequences of the Austral Basin and their foraminiferal assemblages were studied by Malumian, Masiuk and Riggi (1971).
422
Africa Paleocene The Togo/Dahomey s e d i m e n t a r y b a s i n is t h e w e s t e r n p r o l o n g a t i o n of t h e l a r g e N i g e r i a n s e d i m e n t a r y b a s i n ( S l a n s k y , 1 9 6 3 ) .
&.) During t h e Danian ( S l a n s k y , %, c
regressive facies
a r e r e g i s t e r e d which p e r s i s t t h r c u g h o u t t h e P a l e o c e n e .
The
l i t h o l o g i c a l s e q u e n c e of t h e e a r l y P a l e o c e n e i s composed of s e v e r a l s e d i m e n t a r y f a c i e s a t t h e b a s e and a t t h e t o p z o o g e n i c calcareous s t r a t a . The P a l e o c e n e f o r a m i n i f e r a 1 a s s e m b l a g e w a s r e c o r d e d by Slansky (1963). I n s e v e r a l r e g i o n s of N i g e r i a Paleogene s t r a t a have been d e p o s i t e d ; t h e s e s e q u e n c e s a r e found i n s u r f a c e e x p o s u r e s and i n subsurface samples.
I n t h e p r e s e n t work t h e l i t h o s t r a t i -
g r a p h i c scheme o f Adegoke ( 1 9 6 6 ) i s f o l l o w e d and t h e p r i n c i p a l r e f e r e n c e s a r e made t o t h e w e s t e r n and e a s t e r n N i g e r i a format i o n s which m o s t l y were d e p o s i t e d i n s o u t h e r n N i g e r i a . The S o u t h e r n N i g e r i a B a s i n i s l i m i t e d t o t h e n o r t h and t h e e a s t by t h e c r y s t a l l i n e complex of t h e A f r i c a n B o u c l i e r b a s e ment; t o t h e w e s t and s o u t h i t e x t e n d s t o t h e c o n t i n e n t a l margin.
The Lower T e r t i a r y s e d i m e n t a r y o u t c r o p s form a n a r c b o r -
d e r i n g t h e a n c i e n t complex.
The s o u t h e r n b a s i n i s s u b d i v i d e d
i n t h e w e s t e r n and e a s t e r n b a s i n s ; t h e y were p a r t i a l l y s e p a r a t ed by a s u b m a r i n e basement r i d g e ; t h i s s t r u c t u r e may have b e e n p a r t i a l l y r e s p o n s i b l e f o r t h e minor l i t h o f a c i e s d i f f e r e n c e s known t o o c c u r between e a s t e r n and w e s t e r n N i g e r i a (Adegoke, 1966). During t h e P a l e o c e n e m a r i n e s e d i m e n t s were d e p o s i t e d i n both r e g i o n s ; t h e p r i n c i p a l l i t h o s t r a t i g r a p h i c u n i t s are t h e Ewekoro F o r m a t i o n , exposed i n w e s t e r n N i g e r i a and t h e Imo S h a l e which c r o p s o u t i n w e s t e r n and e a s t e r n N i g e r i a
( T e x t F i g . 1).
Paleocene s t r a t a are a l s o w e l l represented i n subsurface s t r a t a . The b a s a l Ewekoro F o r m a t i o n i s composed of b i o g e n i c l i m e stones.
The Imo S h a l e d i s c o n f o r m a b l y o v e r l i e s t h i s f o r m a t i o n ;
i t s l i t h o l o g i c compositioii a r e g r e e n i s h - g r a y ,
f i n e l y laminated
s h a l e s c o n t a i n i n g i n t e r b e d d e d s a n d s t o n e s and g l a u c o n i t e , c u l m i n a t i n g i n d u l l r e d s i l i c e o u s and s a n d y s h a l e s .
According t o
Reyment ( 1 9 6 5 ) t h e Imo S h a l e h a s t h r e e a r e n a c e o u s e q u i v a l e n t s i n e a s t e r n N i g e r i a ; I g a b a b u S a n d s t o n e , Ebenebe S a n d s t o n e and Omuma S a n d s t o n e , The P a l e o c e n e Ewekoro F o r m a t i o n a t i t s t y p e
423
l o c a l i t y c o n t a i n F o r a m i n i f e r a r e c o r d e d by Reyment
(1965).
Typ-
i c a l D a n i a n p l a n k t o n i c F o r a m i n i f e r a were r e c o r d e d from s o u t h e a s t e r n N i g e r i a from t h e Nsukka F o r m a t i o n (Reyment, 1 9 6 5 ) and from s o u t h e r n N i g e r i a by S t o l k ( 1 9 6 3 ) . The D a n i a n S t a g e i s a l s o w e l l documented i n c o a s t a l w e s t e r n N i g e r i a ; s u b s u r f a c e s a m p l e s a t A r a r o m i and I l a r o I and I l a r o I1 b o r e h o l e s a r e composed o f d i a g n o s t i c m i c r o f a u n a l a s s o ciations
( B e r g g r e n , 1 9 6 0 ; Reyment,
1960, 1 9 6 6 ) .
Younger P a l e o -
c e n e i s w i d e l y d i s t r i b u t e d i n w e s t e r n N i g e r i a and t h e p l a n k t o n -
i c f o r a m i n i f e r a 1 c o n t e n t compares w i t h t h a t o f C e n t r a l A m e r i c a (Reyment, 1 9 6 6 ) . I n a d d i t i o n , i n n o r t h e a s t e r n N i g e r i a t h e Kalambaina F o r m a t i o n c o n t a i n s t h e l a r g e f o r a m i n i f e r :
Operculi-
---
na c a n a l i f e r a d ' A r c h i a c ; o n t h e o t h e r hand s p e c i e s of Raniko-
t a l i a and -
D i s c o c y c l i n a were f o u n d i n w e s t e r n and n o r t h w e s t e r n
Nigeria.
A c c o r d i n g t o Reyment ( 1 9 6 6 ) t h e P a l e o c e n e zone of
G l o b o r o t a l i a t r i n i d a d e n s i s i s w e l l d e v e l o p e d and s e v e r a l s u b s e q u e n t zones are c l e a r l y d e l i n e a t e d .
The d i a g n o s t i c t a x a h a v e
b e e n r e c o r d e d by s e v e r a l a u t h o r s from t h e P a l e o c e n e o f Nigeria
( B e r g g r e n , 1960;
Reyment, 1 9 6 0 , 1 9 6 5 , 1 9 6 6 ;
Stolk, 1963). The P a l e o g e n e of Cameroon ( B e l m o n t e , 1 9 6 6 ) c o m p r i s e s P a l e o c e n e and e a r l y t o m i d d l e Eocene d e p o s i t s , r e p r e s e n t e d i n w e s t e r n Cameroon, w h i c h i s t h e c o n t i n u a t i o n o f t h e e a s t e r n Nigeria Basin. The m a r i n e P a l e o c e n e i s known from b o r e h o l e s i n t h e Duala B a s i n c h a r a c t e r i z e d by p l a n k t o n i c and b e n t h o n i c F o r a m i n i f e r a ( d e K l a s z , L e C a l v e z and R & r a t , 1 9 6 4 a ; B e l m o n t e , 1 9 6 6 ) and l a r g e Foraminifera such as Discocyclina g r . a g u e r r e v e r s i ( C a u d i ) and s m a l l Nummulites
(Blondeau, 1 9 6 6 ) .
f a c i e s i n t h e D u a l a B a s i n become more l i t t o r a l , F o r a m i n i f e r a b e i n g found.
To t h e e a s t t h e
only Benthonic
Reyment ( 1 9 6 5 ) r e c o r d e d t h e Mpundo F o r m a t i o n and t h e Logbaba b e d s i n Cameroon a s b e i n g of P a l e o g e n e t o Neogene a g e . They c o n t a i n p l a n k t o n i c , b e n t h o r i c and l a r g e F o r a m i n i f e r a . I n Gabon t h e s e d i m e n t a r y b a s i n was s u b d i v i d e d by d e K l a s z and G a g e o n n e t ( 1 9 6 3 ) i n t o a w e s t e r n and a n e a s t e r n p a r t . Paleocene i s represented i n both b a s i n s ; post-Paleocene
The sedi-
m e n t s were d e p o s i t e d o n l y i n t h e w e s t e r n p a r t o f t h e b a s i n .
424
The P a l e o c e n e c o m p r i s e s :he I k a n d o and l o w e r O z o u r i S t a g e s . The Lower P a l e o c e n e i s r e p r e s e n t e d by t h e I k a n d o F o r m a t i o n and t h e Ikandoian Stage.
The I k a n d o F o r m a t i o n i s l a r g e l y c l a y e y
w i t h a r e e f a l c a r b o n a t e l e v e l ; i t c o n t a i n s p l a n k t o n i c and benthonic Foraminifera
r e c o r d e d by d e K l a s z , Marie
and Meijer ( 1 9 6 0 ) , d e K l a s z and G a g e o n n e t ( 1 9 6 3 ) , d e K l a s z , L e C a l v e z and R L r a t , ( 1 9 6 4 b ) and L e C a l v e z , d e K l a s z and Brun (1974).
The O z u r i F o r m a t i o n i s composed o f s i l i c i f i e d s e d i m e n t s The m i c r o f a u n a l assem-
w i t h c h e r t i n t e r c a l a t e d by c l a y s t o n e s .
b l a g e o f t h e O z u r i a n S t a g e was l i s t e d by d e K l a s z and G a g e o n e t ( 1 9 6 3 ) , d e K l a s z , L e C a l v e z and R&at
,
( 1 9 6 4 b , c ) a n d Graham,
d e K l a s z and R e r a t ( 1 9 6 5 ) . I n Angola f o u r s e d i m e n t a r y b a s i n s a r e d i s t i n g u i s h e d (Antunez, 1 9 6 4 ) :
t h e Congo B a s i n , d e Cuanza B a s i n , t h e Bengue-
l a B a s i n and t h e Moqamedes B a s i n . From t h e s e b a s j . n s o n l y i n t h e Cuanza B a s i n d i d t h e s e d i mentary f o s s i l i f e r o u s sequence which began d u r i n g E a r l y C r e t a The ceous t i m e , c o n t i n u e d u r i n g l a t e Senonian-Paleocene t i m e . c e n t r a l p a r t of t h e b a s i n c o n t a i n s s t r a t a o f more o r l e s s s i l t y s h a l e s whereas i n o t h e r p a r t s of t h e b a s i n t h e Paleocene i s a t h i c k u n f o s s i l i f e r o u s s e q u e n c e o f y e l l o w t o t a n s a n d s t o n e and r e d t o t a n a r g i l l i t e w h i c h may r e p r e s e n t a f l u v i o - d e l t a i c e n v i r o n m e n t ( B r o g n o n and V e r r i e r , 1 9 6 6 ) ; t o w a r d t h e n o r t h , a t Cac u a c o a n d a l s o i n Luanda t h e w e l l s e n c o u n t e r b l a c k u n f o s s i l i f e r o u s s h a l e , a r g i l l i t e and f i n e l y l a m i n a t e d i n t e r b e d s o f d a r k m a r l , c o n t a i n i n g numerous F o r a m i n i f e r a , e s p e c i a l l y c a r i n a t e p l a n k t o n i c s (Broghon and V e r r i e r , 1 9 6 6 )
.
I n t h e Cuanza b a s i n t h e P a l e o g e n e ( A n t u n e z , 1 9 6 4 ) o v e r l i e s c o n c o r d a n t l y t h e M a a s t r i c h t i a n , t h e i r boundary b e i n g d i f f i c u l t t o determine.
A c c o r d i n g t o Happener ( 1 9 5 8 , i n A n t u n e z , 1 9 6 4 )
t h e r e e x i s t s a r i c h m i c r o f a u n a o f G l o b o r o t a l i a and G l o b i g e r i n a d a t e d a s e a r l y Eocene w i t h q u e s t i o n a b l e P a l e o c e n e a t t h e b a s e . D e K l a s z , M a r i e and M e i j e r
( 1 9 6 0 ) made r e f e r e n c e t o t h e
p r e s e n c e o f G a b o n e l l a g i g a n t e a d e K l a s z and lzleijer i n t h e DanoM o n t i a n of A n g o l a , a s p e c i e s w h i c h was a l s o f o u n d i n Gabon i n s t r a t a of t h e same a g e . Eocene The Eocene s e d i m e n t a r y s e q u e n c e of Togo and Dahomey
425
( S l a n s k y , 1 9 6 3 ) i s marked by c l a y e y s e d i m e n t a t t h e i r b a s e beYpresian s t r a t a are
coming g l a u c o n i t i c and m a r l y a t t h e t o p .
developed i n c l a y e y f a c i e s a t t h e b a s e w i t h g l a u c o n i t i c marly clay a t the top. Sequences a t t r i b u t e d t o t h e L u t e t i a n v a r y between 1 0 and 1 7 0 meters o f l i m e s t o n e s w i t h a p h o s p h a t i c l a y e r a t t h e b a s e and becoming l i g h t g r a y m a r l y c l a y w i t h t h i n gypsum b e d s a t t h e top.
The L u t e t i a n seems t o r e p r e s e n t a r e g r e s s i v e f a C i e S
(Slansky, 1963). I n Dahomey t h e Oshosum F o r m a t i o n , which i s a l s o d e v e l p e d i n N i g e r i a n e a r I f o , y e i l d e d many p l a n k t o n i c F o r a m i n i f e r a s u c h a s G l o b o r o t a l i a a e q u a Cushman and Renz, G l o b o r o t a l i a p s e u d o t o p i l e n s i s Bronnimann and G l o b i g e r i n a s a l d a d o e n s i s Bronnimann, w h i c h a r e c o r r e l a t e d w i t h t h e s e q u e n c e o f t h e b o r e h o l e a t O t t a , W e s t e r n N i g e r i a , by Reyment ( 1 9 6 5 ) . O t h e r Eocene m i c r o f a u n a s a r e l o c a l l y a b u n d a n t i n Togo and Dahomey.
T h e r e a r e a l s o t h e l a r g e F o r a m i n i f e r a s u c h a s Disco-
cyclina,
Nummulites,
and C l a v i g e r i n e l l a .
The p l a n k t o n i c ,
b e n t h o n i c and l a r g e F o r a m i n i f e r a were r e c o r d e d by S l a n s k y ( 1 9 6 3 ) and Reyment ( 1 9 6 5 ) . The Eocene of N i g e r i a and a d j a c e n t a r e a s was a p e r i o d of regression.
The s e d i m e n t s a r e l o c a l l y f o s s i l i f e r o u s and p r e -
dominantly f i n e g r a i n e d
(Adegoke, 1 9 6 6 )
.
D i a g n o s t i c s p e c i e s were r e c o r d e d from b o r e h o l e s a t I j u and Otta
(Oshosun F o r m a t i o n ) by B e r g g r e n ( 1 9 6 0 ) , Reyment ( 1 9 5 9 ) and
t h o s e r e c o r d e d by d e K l a s z , L e C a l v e z a n d R&at
(196433, c ) and
Adegoke ( 1 9 6 6 ) . Berggren
( 1 9 6 0 ) p o i n t e d o u t t h e a b s e n c e i n t h i s a r e a of
t h e g e n u s H a n t k e n i n a a n d o t h e r g e n e r a t y p i c a l o f t h e Eocene i n T r i n i d a d s u c h a s G l o b i g e r a p s i s and P o r t i c u l a s p h a e r a .
Reyment
( 1 9 6 5 ) s u g g e s t e d a n Y p r e s i a n t o Lower L u t e t i a n a g e f o r t h e Oshosum F o r m a t i o n w h e r e a s Adegoke ( 1 9 6 6 ) a c c e p t s a m i d d l e Eocene ( L u t e t i a n ) a g e .
I n w e s t e r n N i g e r i a t h e Oshosun Forma-
t i o n i s o v e r l a i n by t h e Ameki F o r m a t i o n ;
i n eastern Nigeria the
Ameki F o r m a t i o n i s t h e o n l y r e c o g n i z e d l i t h o s t r a t i g r a p h i c u n i t . The t y p e area o f t h e Ameki F o r m a t i o n i s i n e a s t e r n N i g e r i a , w h e r e i t i s composed p r e d o m i n a n t l y o f g r e e n i s h - g r a y ,
clayey
s a n d s t o n e and s a n d y c l a y s t o n e , w i t h r a p i d l a t e r a l f a c i e s chang-
es.
I n w e s t e r n N i g e r i a t h e Ameki F o r m a t i o n form a t h i n v e n e e r
426
above the phosphate-bearing beds of the Oshosun Formation (Adegoke, 1966). The Ameki Formation in eastern Nigeria is richly fossiliferous. Outcrops of the Ameki Formation in western Nigeria are non-fossiliferous and only few species were recovered from boreholes at Otta and Iju (Berggren, 1960; Reyment, 1960). The microfaunal assemblage of the Ameki Formation were recorded by Berggren (1960), Reyment (1960; 1965), Stolk (1963) and Adegoke (1966). They are listed in Text Fig. 3. In addition Reyment (1966) mentioned the large foraminifer Discocyclina sp. at some localities, i.e. Lakolabo. The Ameki Formation is considered to be of Lutetian to Lower Bartonian age by Adegoke (1966); Reyment (1965) assigned an early and middle Eocene (Ypresian-Lutetian) age for this formation. Stolk (1963) stated that during the early, middle and late Eocene it is not possible here to subdivide the microfaunal succession into the zones proposed by Bolli for the Caribbean area. Locally,Stolk (1963) found the following biostratigraphic units: Late Eocene - Chiloguembelina martini cubensis Zone, Middle Eocene - Cassigerinelloita amekiensis Zone, Early Eocene - Globorotalia formosa Zone. According to Reyment (1966) the late Eocene Was not well defined in Nigeria and evidences of the presence of the Oligocene Stage is not extensive, On the other hand Stolk (1963) mentioned the following late Eocene or early Oligocene planktonic Foraminifera: Globigerina angustiumbilicata Bolli, Chiloguembelina cubensis (Palmer), Pseudohastigerina aff. p. micra Cole and Globorotaloides suteri Bolli. The Eocene of Cameroon is not well characterized,being found in isolated outcrops (Belmonte, 1966). In Kwa-Kwa the basal Eocene contains a fauna of Nummulites indicating a basal (Ypresian) Eocene age (Belmonte, 1966). In Bomono Eocene strata are found which provided a poor foraminifera1 microfauna such as (Belmonte, 1966): Cancris cf. cocoaensis Cushman, Planulina sp., Loxostomum s p . and Sagrina sp.. De KlasZ, Le Calvez and R&at (1964a) recorded Bolivina crassicostata from the Paleogene of Cameroon and Blondeau (1965) mentioned the large foraminifer Nummulites (Operculinoides) furoni Blondeau which is also found in Senegal and Ivory Coast. The middle
427
l a t e Eocene a n d O l i g o c e n e s t a g e s a r e unknown i n Cameroon (Belmonte, 1966). I n t h e Gabon B a s i n s e q u e n c e s o f Eocene a g e a r e m e n t i o n e d from t h e w e s t e r n p a r t of t h e b a s i n .
I n t h i s a r e a t h e Eocene
Upper O z o u r i , Animba a n d N ' G o l a f o r m a t i o n s were r e c o g n i z e d ( L e C a l v e z , K l a s z and B r u n , 1 9 6 4 ) . The m i c r o f a u n a o f t h e Eoc e n e Upper O z o u r i F o r m a t i o n i s n o t w e l l known, b u t i n c l u d e s
many s p e c i e s i n common w i t h t h e o v e r l y i n g Animba F o r m a t i o n ( d e
K l a s z and G a g e o n n e t , 1 9 6 3 ) .
The Eocene Animba F o r m a t i o n i s
s e p a r a t e d f r o m t h e O z o u r i F o r m a t i o n by a d i s c o n f o r m i t y and i t s u p p e r p a r t i s o v e r l a i n d i s c o n f o r m a b l y by m a r i n e Miocene s t r a t a . The Animba F o r m a t i o n i s m a i n l y composed of p h o s p h a t i c c l a y s t o n e s w i t h some d e t r i t i c i n t e r c a l a t i o n s .
The m i c r o f a u n a l con-
t e n t o f t h i s f o r m a t i o n w a s s t u d i e d by d e K l a s z and R & a t
(1961)
d e K l a s z and Gageonnet ( 1 9 6 3 ) , d e K l a s z , L e C a l v e z and R & a t (196433, c ) , Graham, d e K l a s z and R & a t
( 1 9 6 5 ) and Adegoke
(1966). The l a t e Eocene N ' G o l a F o r m a t i o n i s o n l y f o u n d i n s u b s u r f a c e samples i n b o r e h o l e s a t t h e s o u t h e a s t of P o r t G e n t i l . This formation yielded the following Foraminifera Gageonnet, 1963) :
( d e K l a s z and
H a n t k e n i n a p r i m i t i v a Cushman and J a r v i s ,
B o l i v i n a s t r i a t e l l a t a Bandy and M a n d j i n a e x c a v a t a d e K l a s z and R&at.
The N ' G o l a a s s o c i a t i o n , a l t h o u g h i t p o s s e s s e s i t s own
f e a t u r e s , shows some s i m i l a r i t i e s w i t h t h e w i t h t h e u n d e r l y i n g Animba F o r m a t i o n ( d e K l a s z a n d G a g e o n n e t , 1 9 6 3 ) . The Eocene of Angola i s r e p r e s e n t e d by t h e G r a t i d z o and Cunga f o r m a t i o n s (Brognon a n d V e r r i e r , 1 9 6 6 ) .
The G r a t i d z o
F o r m a t i o n i s made up o f w h i t i s h c h a l k , r i c h c h e r t o c c u r r e n c e s i n some p l a c e s , and t h e p r e s e n c e o f s i l i c i f i e d l i m e s t o n e s ;
it
i s i n t e r b e d d e d b l a c k b i t u m i n o u s m a r l a n d a r g i l l i t e (Brognon and V e r r i e r , 1 9 6 6 ) . A c c o r d i n g t o Happener ( 1 9 5 8 , i n A n t u n e z , 1 9 6 4 ) t h e r e e x i s t s a r i c h m i c r o f a u n a o f G l o b o r o t a l i a and G l o b i g e r i n a d a t e d a s e a r l y Eocene. The Cunga F o r m a t i o n i s composed of g r a y marls w i t h c a l -
it c o n t a i n s charact e r i s t i c d o l o m i t i c c o n c r e t i o n s w i t h H a s t i g e r i n e l l a and
careous i n t e r c a l a t i o n s , p a r t l y s i l i c i f i e d ;
428
Hantkenina. I n S o u t h A f r i c a (Chapman (1928) from a f o r a m i n i f e r a 1 l i m e s t o n e of l a t e Eocene a g e of t h e A l e x a n d r i a F o r m a t i o n , r e c o r d e d s e v e r a l benthonics associated with t h e l a r g e Foraminifera D i s c o c y c l i n a p r a t t i (Mich. ) and D i s c o c y c l i n a v a r i a n s
(Kaufmann)
.
A t l a n t i c Ocean I n t h e l a s t two d e c a d e s t h r o u g h o u t s e v e r a l c r u i s e s o v e r t h e o c e a n s and t h e i n t e n s i v e development of m a r i n e r e s e a r c h , s e v e r a l p r o j e c t s were c a r r i e d o u t . Oceanographic v e s s e l s s u c h a s Vema, C h a i n , Glomar C h a l l e n g e r , e t c . , o b t a i n e d d e e p - s e a c o r e s from s e v e r a l a r e a s of t h e S o u t h A t l a n t i c Ocean.
The A t -
l a n t i c Ocean i s of i m p o r t a n c e b e c a u s e t h e o b t a i n e d d a t a a l l o w e d i n f o r m a t i o n t o b e added a b o u t t h e a g e of t h e c o r e d s e d i m e n t s and a b o u t t e c t o n i c p l a t e s , s e a f l o o r s p r e a d i n g and c o n t i n e n t a l d r i f t g e o l o g i c a l h y p o t h e s i s and o t h e r e v e n t s which o c c u r r e d d u r i n g t h e l a s t 135 m i l l i o n y e a r s . Several authors contributed with micropaleontological stud i e s t o t h e b e s t knowledge of t h e s e d i m e n t a r y s e q u e n c e s found i n t h e S o u t h A t l a n t i c Ocean.
S a i t o , Ewing and B u r c k l e ( 1 9 6 6 )
from c o r e s o b t a i n e d by t h e v e s s e l Vema, r e c o r d e d Upper C r e t a c e o u s t o P l i o c e n e s e d i m e n t s from t h e S o u t h A t l a n t i c .
Paleogene
s e d i m e n t s w e r e o b t a i n e d from t h e Rio Grande R i s e l a t i t u d e 282
36's and l o n g i t u d e 2 9 2 01'W; a t t h i s s i t e . Eocene s e d i m e n t s w i t h Hantkenina a l a b a m e n s i s Cushman, Hantkenina p r i m i t i v a
-
Cushman and J a r v i s , G l o b i g e r a p s i s i n d e x F i n l a y l a n d G l o b o r o t a l i a c e n t r a l i s Cushman and Bermudez w e r e r e c o r d e d . T h i s , t h e o l d e s t s e d i m e n t c o r e d from Rio Grande R i s e i s from a f a u l t s c a r p on the north side.
C i f e l l i , Blow and N e l s o n ( 1 9 6 8 ) , from a d r e d g e
sample r e c o v e r e d by t h e Chain from a f r a c t u r e zone at. 01P 23.5'S,
2 9 Q 1 2 W r e c o r d e d P a l e o c e n e and Lower Eocene p l a n k t o n i c s .
The P a l e o c e n e ( T h a n e t i a n ) was m a i n l y r e p r e s e n t e d by b o r o t a l i a ( G l o b o r o t a l i a ) v e l a s c o e n s i s and G l o b o r o t a l i a
e-
(G.)
p s e u d o m e n a r d i i w h e r e a s t h e Eocene s e d i m e n t s c o n t a i n e d t y p i c a l Y p r e s i a n p l a n k t o n i c F o r a m i n i f e r a , m a i n l y G l o b o r o t a l i a formosa f o r m o s a , G l o b o r o t a l i a ( T u r b o r o t a l i a ) c o l l a c t e a and P s e u d o h a s t i g e r i n a e o c e n i c a , a l l t y p i c a l of t h e G l o b o r o t a l i a formosa formosa Zone. Maxwell e t a l .
( 1 9 7 0 ) recorded t h e r e s u l t s of t h e c r u i s e
of t h e Glomar C h a l l e n g e r from Leg 3 which c o v e r e d t h e S o u t h
429
A t l a n t i c b e t w e e n D a k a r , S e n e g a l and R i o d e J a n e i r o , B r a z i l . The r o c k s a m p l e s w e r e o b t a i n e d from s e v e n s i t e s l o c a t e d o n t h e m i d - A t l a n t i c R i d g e f l a n k s a n d t w o s i t e s on t h e R i o Grande R i s e : m o s t of t h e s e d i m e n t s c o r e d c o n t a i n e d p l a n k t o n i c F o r a m i n i f e r a r a n g i n g i n a g e from Campanian t o l a t e P l e i s t o c e n e . Maxwell
g. (1970
) d e s i g n a t e d n i n e submarine subsur-
f a c e f o r m a t i o n s r a n g i n g from Upper C r e t a c e o u s t o Yolocene i n t h e mid-Atlantic Province.
The a u t h o r s u s e d t h e names of h i s -
t o r i c exploratory v e s s e l s i n an a l p h a b e t i c a l o r d e r .
They a r e
i n o r d e r o f i n c r e a s i n g d e p t h and r a n g e o f a g e of e a c h formation: A l b a t r o s Ooze, P l i o c e n e - Q u a t e r n a r y B l e k e Ooze, L a t e P l i o c e n e - P l i o c e n e C h a l l e n g e r Ooze, L a t e Miocene-Early P l i o c e n e D i s c o v e r y C l a y , L a t e O l i g o c e n e - M i d d l e Miocene Endeavor Ooze, L a t e O l i g o c e n e - M i d d l e Miocene Fram Ooze, E a r l y O l i g o c e n e - E a r l y Miocene G a z e l l e Ooze, M i d d l e - L a t e Eocene Grampus Ooze, M i d d l e Eocene-Early
Miocene
H i r o n d e l e Ooze, L a t e C r e t a c e o u s - e a r l y M i d d l e Eocene Maxwell
g. (1970
)
found t h a t t h e Early t o Middle
C e n o z o i c p l a n k t o n i c f o r a m i n i f e r a 1 f a u n a s a r e d i v e r s e and b e a r a c l o s e r e s e m b l a n c e t o t h o s e r e p o r t e d from t r o p i c a l r e g i o n s o f t h e world.
Paleogene sediments w e r e recovered a t :
Site 1 4
w i t h t h e Zone o f C r i b o h a n t k e n i n a i n f l a t a , and a t S i t e 19 w i t h t h e Zone of H a n t k e n i n a a r a g o n e n s i s ,
A t
s i t e s 2 1 and 2 2 on t h e
R i o Grande R i s e Lower Miocene t o M a a s t r i c h t i a n a s s e m b l a g e s were
recognized. B l o w ( 1 9 7 0 ) made a n e x t e n s i v e s t u d y o f c o r e s r e c o v e r e d
d u r i n g Leg 3 o f t h e Glomar C h a l l e n g e r c r u i s e .
P a l e o g e n e se-
q u e n c e s w i t h t y p i c a l C a r i b b e a n p l a n k t o n i c F o r a m i n i f e r a were r e c o g n i z e d a t s i t e s 1 4 , 1 7 , 1 8 , 1 9 , 2 0 , 2 1 and 2 2 ; a t t h e s e h o l e s P a l e o g e n e p l a n k t o n i c zones were i d e n t i f i e d by Blow ( o p . c i t . ) which a r e summarized below.
Funnel1 (1971) i n t h e index of pre-Quaternary occurrences i n t h e o c e a n s , r e c o r d e d from t h e S o u t h A t l a n t i c s e v e r a l locat i o n s i n w h i c h P a l e o g e n e a n d p a r t i c u l a r l y Eocene s e d i m e n t s occur. T h i s i n f o r m a t i o n i s summarized below. B e r g g r e n and Amdurer ( 1 9 7 3 ) r e c o r d e d P a l e o g e n e s e d i m e n t s ,
430 a l s o r e c o v e r e d i n t h e S o u t h A t l a n t i c by t h e Deep S e a D r i l l i n g P r o j e c t d u r i n g t h e c r u i s e of Leg 3 .
The l o c a t i o n s o f t h e P a -
l e o g e n e s i t e s a n d t h e i r c o r r e s p o n u i n g a g e s a r e summarized b e l o w . Berggren
(1973) s y n t h e s i z e d t h e Paleogene p l a n k t o n i c f o r -
a m i n i f e r a 1 f a u n a s on l e g s I - I V
f r o m t h e Deep S e a D r i l l i n g P r o -
i s given.
j e c t i n which a d d i t i o n a l and d e t a i l e d i n f o r m a t i o n
The s e v e r a l c o r e s o b t a i n e d f r o m t h e A t l a n t i c Ocean i n w h i c h P a l e o g e n e s e d i m e n t s a r e r e c o r d e d a r e summarized b e l o w , i n w h i c h the location, a s w e l l a s t h e a g e a n d t h e p l a n k t o n i c z o n e s found i n t h e c o r e d s e d i m e n t s are i n d i c a t e d .
Text Fig.
2 shows
t h e l o c a t i o n as w e l l as t h e P a l e o g e n e s e q u e n c e s e n c o u n t e r e d d u r i n g t h e c r u i s e s of s e v e r a l v e s s e l s . 002 1 5 ' s
-
1 4 0 25'W
P a l e o c e n e a n d Lower E o c e n e , i n o r d e r of i n c r e a s i n g a g e Zones o f
-G .
g.
velascoensis
pseudomenardii
-G . 012 2 3 . 5 ' s
282 58.72s 2EQ 0 2 . 7 4 s 280 1 9 . 8 9 s 28Q 3 1 . 4 7 s
-
29Q 1 2 W
P a l e o c e n e - E a r l y Eocene ( T h a n . - Y p r e s i a n )
0 8 2 00.70W-Late
06Q36 W
aquiensis
-
Oligocene-Early Mioc.-Pleist.
Z.N3
L a t e O l i g o c e n e - E a r l y Miocene Z.P19-N3
L a t e E o c e n e - E a r l y Miocene Z.Pl7-19
20Q 54.46W
-
26p 50.75W
,
Oligocene
Z .P18 1 9
L a t e Eocene
Z .P16-18
M i d d l e Eocene
Z .P9-14
E a r l y Eocene
Z .P8
B a s a l Eocene o r l a t e Upper Paleocene Z.P6, 282 3 2 . 0 8 s
-
232 4 0 . 1 3 W
Z .P6
Oligocene
Z .P18-N2
L a t e Eocene
-
302 3 5 . 8 5 W
Z .P14-17?
M i d d l e Eocene
z . P 1 1 ,1 2 z .P10,11
E a r l y Eocene
Z.P
M i d d l e Eocene 28Q 3 5 . 1 0 s
7
Paleocene
8
E a r l y Eocene t o L a t e Upper P a l e o c . Z.P4,6 28Q 36 S
-
29Q 0 1 W
L a t e Eocene w i t h H a n t k e n i n a a l a b a m e n s i s Hantkenina p r i m i t i v a Globigerapsis index Globorotalia c e n t r a l i s
302 00.31s
-
352 15.00W
-
L a t e Oligocene
Middle hocene
Z.N
1-3
z .P11,12
431
42p 32's 4 7 s 29's 4 8 2 29's 65p 4 3 ' s
-
562 29'W
Upper Cretaceous-Lower T e r t i a r y
5gP 21 W 5gP 21 W 162 25 E
Eocene
Text Fig. 2 .
Eocene Eocene
L o c a t i o n s a t which P a l e o g e n e s e q u e n c e s i n t h e The f i g u r e i l l u s SouOh A t l a n t i c were f o u n d . trates a l s o t h e marine sedimentary b a s i n s i n which P a l e o g e n e t r a n s g r e s s i o n s t o o k p l a c e . Some of t h e P a l e o g e n e s e q u e n c e s were o n l y r e c o r d e d from t h e c o n t i n e n t a l m a r g i n . Summary
I n t h e S o u t h American c o n t i n e n t P a l e o c e n e s e q u e n c e s a r e more c o m p l e t e i n n o r t h e a s t e r n B r a z i l t h a n i n s o u t h e r n r e g i o n s , s u c h as i n A r g e n t i n a where o n l y t h e Danian S t a g e i s r e p r e s e n t e d P a l e o c e n e p l a n k t o n i c F o r a m i n i f e r a a r e common t o a l l S o u t h A t l a n t i c a r e a s and a r e s t r o n g l y r e l a t e d t o t h o s e r e c o r d e d by B o l l i (1966) from t h e C a r i b b e a n r e g i o n . Paleocene b e n t h o n i c assemblages are s p r e a d o v e r l a r g e a r e a s ; t h e y show many s p e c i e s common t o s e v e r a l S o u t h American basins.
A l t h o u g h c o s m o p o l i t a n and European s p e c i e s a r e f o u n d ,
t h e p r e v a i l i n g a s s e m b l a g e s a r e t h o s e r e c o r d e d from t h e U n i t e d
432
States
A t l a n t i c a n d Gulf C o a s t a r e a s , p a r t i c u l a r l y t h o s e o f
t h e Midwayan S t a g e . Eocene p l a n k t o n i c a s s e m b l a g e s e n c o u n t e r e d n o r t h o f L a t . 30's
a r e s i m i l a r t o t h o s e r e c o r d e d from t h e C a r i b b e a n a r e a ,
whereas A u s t r a l a s i a n i n f l u e n c e s a r e observed i n h i g h e r l a t i tudes, Eocene b e n t h o n i c F o r a m i n i f e r a show s i m i l a r f e a t u r e s .
In
a d d i t i o n , i n t h e southernmost areas, such a s t h e A u s t r a l B a s i n some P a c i f i c i n f l u e n c e s a r e r e g i s t e r e d . O l i g o c e n e p l a n k t o n i c s are o n l y r e p r e s e n t e d on t h e c o n t i n e n t a l margin of n o r t h e r n S o u t h America, whereas i n s o u t h e r n l a t i t u d e s , i . e . i n t h e C o l o r a d o and A u s t r a l B a s i n O l i g o c e n e transgression i s recorded. A l a r g e p a r t of
t h e P a l e o g e n e seems t o b e a b s e n t f r o m
southeastern Brazilian basins. No l a r g e F o r a m i n i f e r a a r e r e c o r d e d from t h e S o u t h American Paleogene s equences . I n t h e A f r i c a n c o n t i n e n t P a l e o c e n e b e n t h o n i c s show some common s p e c i e s t o t h e Midwayan f a u n a , a l t h o u g h some endemic s p e c i e s do o c c u r . D u r i n g Eocene times t h e endemism o f b e n t h o n i c f o r a m i n i f e r -
a 1 s p e c i e s becomes s t r o n g e r t h a n d u r i n g t h e P a l e o c e n e .
Al-
t h o u g h i n Gabon t h e m i c r o f a u n a l a s s e m b l a g e s shows t h e s t r o n g e s t endemic f e a t u r e s , s e v e r a l new b e n t h o n i c s p e c i e s a r e r e c o r d e d from Togo-Dahomey,
N i g e r i a a n d Cameroon.
L a r g e F o r a m i n i f e r a a r e r e c o r d e d o n l y from t h e Eocene o f t h e A f r i c a n c o n t i n e n t s u c h as R a n i k o t a l i a and D i s c o c y c l i n a i n S o u t h A f r i c a and Nummulites from Cameroon, T y p i c a l warm water p l a n k t o n i c s , s u c h a s H a n t k e n i n a a r e r e c o r d e d o n l y f r o m Gabon and A n g o l a , b e i n g a b s e n t i n t h e American S o u t h A t l a n t i c . O l i g o c e n e s t r a t a seems t o b e m i s s i n g i n A f r i c a a l t h o u g h some l o n g r a n g i n g s p e c i e s a r e r e c o r d e d f r o m N i g e r i a . References Adegoke, 0 . S . 1 9 6 6 . Eocene S t r a t i g r a p h y o f S o u t h e r n N i g e r i a . C o l l o q u e s u r 1 ' E o c e n e . P a r i s . v . 8 , p p . 23-49. A n d r e f s , R . R . , Mazzonf, M . M . a n d S p a l l y t t i , L . A . 1 9 7 3 . Geol o g i a y s e d i m e n t o l o g i a d e 1 , C e r r o Bororo ( P r o v i n c i a d e C h u b u t ) . A c t a s Q u i n t o C o n g r e s o GeoLoTico A r g e n t i n o . T . 1 1 1 , p . 21-55. A n t u z , M . T . , 1 9 6 4 . 0 N e o c r e t a c e o e o C e n o z o i c o do l i t o r a l d e J u n t a d e I n v e s t i g a c o e s d e U l t r a m a r , L i s b o a . p . 1-259. Angola.
433
B e l m o n t e , Y . C . , 1 9 6 6 . S t r a t i g r a p h i e du b a s s i n s e d i m e n t a i r e du Cameroun. P r o c . 2nd. W . A f r i c a n M i c r o p a l . C o l l . I b a d a n , 1 9 6 5 . B e r g g r e n , W . A . , 1 9 7 3 . The P l i o c e n e t i m e - s c a l e : c a l i b r a t i o n o f p l a n k t o n i c f o r a m i n i f e r a 1 and c a l c a r e o u s n a n n o f o s s i l z o n e s : N a t u r e , v . 243, n o . 5407, p . 391-397. , and Amdurer, M . , 1 9 7 3 . L a t e P a l e o g e n e ( O l i g o c e n e ) and Neogene p l a n k t o n i c f o r a m i n i f e r a 1 b i o s t r a t i g r a p h y o f t h e A t l a n t i c Ocean ( L a t . 30' N t o L a t . 30' S ) R i v . I t a l . P a l e o n t . v . 7 9 , no. 3 , p . 337-392. B e r t e l s , A . , 1964. M i c r o p a l e o n t o l o g i a d e l P a l e o c e n o d e G e n e r a l Roca ( P r o v i n c i a d e Rio N e g r o ) . Rev. Museo La P l a t a . Nueva S e r i e . P a l e o n t o l o g i a No. 2 3 , T . I V , p . 125-$84. , 1969. E s t r a t i g r a f i a d e l l i m i t e C r e t a c i c o - T e r c i a r i o en P a t a g o n i a S e p t e n t r i o n a l . Rev. Asoc. G e o l . Arg. T . X X I V , No. 1, p . 41-54. , , 1 9 7 0 . Los F o r a m i n i f e r o s p l a n c t 6 n i c o s d e l a c u e n c a C r e t a c i c o - T e r c i a r i a en P a t a g o n i a S e p t e n t 5 i o n a l ( A r q e n t i n a ) , con c o n s i d e r a c i o n e s s o b r e l a e s t r a t i g r a f i a d e F o r t i n G e n e r a l Roca ( P r o v i n c i a d e R i o N e g r o ) . A m e g h i n i a n a , T . V I I , No. 1, p . 1-47. , 1 9 7 3 . B i o e s t r a t i g r a f i a d e l C e r r o Boror;, P r o v i n c i a d e l C h u b u t , R e p 6 b l i c a A r g e n t i n a . A c t a s V . C o n g r e s o G e o l . Arg. T . 111, p . 71-90. , 1975. B i o e s t r a t i g r a f i a d e l Palehgeno , e n l a R e p h b l i c a A r g e n t i n a . Rev. E s p a E o l a d e M i c r o p a l e o n t o l o g i a , v o l . V I I , no. 3 , p . 429-450. , 1 9 7 6 a ( i n p r e s s ) . B i o e s t r a t i g r a f l a d e l P a l e o c e n o mar i n o en l a P r o v i n c i a d e l Chubut, Republica A r g e n t i n a . B l o n d e a u , A , , 1 9 6 6 . D 6 c o u v e r t e d e Nummulites a u Cameroun. P r o c . 2nd. W . A f r i c a n M i c r o p a l . C o l l . I b a d a n . p . 2 4 - 2 6 . Blow, W . H . , 1 9 7 0 . A Deep S e a D r i l l i n g P r o j e c t . Leg 3 . Foramin i f e r a from s e l e c t e d s a m p l e s . I n Maxwell, A . E . e t a l . , 1970. I n i t i a l R e p o r t s of t h e Deep S e a D r i l l i n g Project,v,III. W a s h i n g t o n ( U . S . Government P r i n t i n g O f f i c e ) p . 629-661 B o l l i , H . M . , 1966. Z o n a t i o n of C r e t a c e o u s t o P l i o c e n e marine s e d i m e n t s b a s e d o n p l a n k t o n i c F o r a m i n i f e r a . B o l . I n f . Asoc. B e n e z o l a n a d e G e o l . Min. y P e t r o l e o . , v . 9 , no. 1: p . 1-32. Brognon, G . a n d V e r n i e r , G . , 1 9 6 6 . T e c t o n i q u e e t s e d i m e n t a t i o n d a n s l e b a s s i n du Cuanza ( A n g o l a ) . I n : B a s s i n s s k d i m e n t a i r e s du l i t t o r a l a f r i c a i n . , I & r e p a r t i e : L i t t o r a l A t l a n t i q u e . A s s o c . d e s S e r v i c e s G e o l . A f r i c a i n s . p . 207-252. Chapman, F . , 1 9 2 8 . O n a F o r a m i n i f e r a 1 L i m e s t o n e o f Upper E o c e n e Age from t h e A l e x a n d r i a F o r m a t i o n , S o u t h A f r i c a . A n n a l s of t h e S o u t h A f r i c a n Museum, v o l . X X V I I I , P t . 2 , p . 291-296. C h a r r i e r , R . and L a h s e n , A . , 1 9 6 9 . S t r a t i g r a p h y of L a t e C r e t a c e o u s - E a r l y E o c e n e , S e n o S k y r i n g S t r a i t of M a g e l l a n A r e a , M a g a l l a n e s P r o v i n c e , C h i l e . American A s s o c i a t i o n of P e t r o l e u m G e o l o g i s t s B u l l . v . 5 3 , no. 3 , p . 568-590. C i f e l l i , R . , Blow, W . H . , and N e l s o n , W . G . , 1 9 6 8 . P a l e o g e n e S e d i m e n t f r a m a F r a c t u r e Zone o f t h e m i d - A t l a n t i c R i d g e . J o u r n a l of M a r i n e R e s e a r c h . v . 2 6 , p . 105-109. d e K l a s z , I . , and G a g e o n n e t , R . , 1 9 6 3 . B i o s t r a t i g r a p h i e du B a s s i n Gabonais. C o l l o q u e I n t e r n a t . d e M i c r o p a l . Dakar, p. 227-304.
434
de Klasz, I., Le Calvez, Y. and RLrat, D., 1964 a. Deux importantes espsces de Foraminifgres du Miocbye inferieur,de 1' Afrique occidentale. C.R. Sommaire des Seanc. SOC. Geol. France, Fasc. 5, p. 194-195. , 1964 b. Deux nouveaux Buliminidae (Foraminifgrys) du Pal&og+ne de 1'Afrique occidentale. CR. Sommaire des Seanc. S O C . Geol. France. Fasc. 5, p. 208-209. , 1964 c. Deux nouveaux genres de Foraminif$res de Gab?n (Afrique Equatoridle). C.R. Sommaire des S6anc. SOC. Geol. France. Fasc. 6, p. 236-237. de Klasz, I., Marie, P., and Meijer,.M., 1960., Gabonella nov. gen. un nouveau genre de Foraminifhres du Cretace Suphrieur et du Tertiaire basal de l'Afrique occidentale. Revue Micropal. v. 3 , no. 3, p. 167-182. Funnel, B. M., 1971. The occurrence of pre-Quaternary microfossils in the oceans. (In) Funnel, B. M. and Riedel, W. (eds.) "Micropaleontology of Oceans," Cambridge, Cambridge Univ. Press, 1967. p. 507-534. , Graham, J. J., de Klasz, I., and Rerat, D., 1965. Quelques importants Foraminif&res du Tertiaire du Gabon (Afrique &quatoriale). Rev . Micropal., v. 8, no. 2, p. 71-84. Herm, D., 1966. Micropaleontological aspects of the Magellanese Geosyncline, southernmost Child, South America. Proc. 2nd. W. African Micropal. Coll. Ibadan. p. 72-86. Kaasschieter, J. P. H., 1963. Geology of the Colorado Basin. Tulsa Geol. S O C . Digest, p. 177-187. Le Calvez, Y., de Klasz, I. and Brun, L., 1974. Nouvelle contribution & la connaissance des microfaunes du Gabon. Rev. Espagola de Micropal. v. VI, no. 3, p. 38;-400. Lima, E. C., 1973. Bioestratigrafia da Bacia de Barreirinhas. Anais do XXVI Congreso SOC. Brasilera de Geol. Belem., v. 3, p. 81-91. Mabesone, J. M., Tinoco, I. M., and Coutinho, P. N., 1968. The Mesozoic-Tertiary Boundary in northeastern Brazil. Palaeogeography, Paleoclimatology, Palaecology, v. 4, p. 161-185. Malumian, N., 1970b. Foraminiferos danianos de la Formacibn Pedro Luro, Provincia de Buenos Aires, Argentina. Ameghinianal T. VII, no. 4 , p. 355-367. Malumian, N., Masiuk, V . $ and Riggi, J. C., 1971. Micropaleontologla y sedimentologia de la perforacidn SC-1 Provincia Santa Cruz, RepAblica Argentina. Su importancia y correlaciones. Rev. ASSOC. Geol. Argentina. T. XXVI, no. 2, p.175-208. Masiuk, V., 1967. Estratigrafia del Rocanense del Puesto P. Alvarez. Curso inferior del Rio Chico, Prov. del Chubut. Rev. Museo La Plata (Nueva Serie) , T. V, p. 197-258. Maxwell, A. E . , et al,., 1970. Initial Reports of the Deep Sea Drilling Project, v. III., Washington, U.S. Government Printing Office, xx+ 806 p . Mendezl I. A., 1966. Foraminiferos, edad y correlaci&n estratigrafica del Salamanquense de Punta Peligro (45O 30's; 67O 1l'W) Provincia del Chubut. Rev. ASOC. Geol. Arg., T. XXI, no. 2, p. 127-159. Noguti, I. , 19j5. ZonaciAn Bioestratigrifica de 10s Foramidferos planctonicos del Terciario de Grasil. Rev. Espagola de Micropal., v. VII, no. 3 , p. 391-401. Noguti, I., and Fontaga dos Santos, J., 1972. Zoneamiento pre liminar por foraminiferos planctonicos do Aptiano ao Mioceno
435
na p l a t a f o r m a c o n t i n e n t a l d o B r a s i l . B o l . T e c . P e t r o b r a s . Rio d e J a n e i r o , v . 1 3 , n o . 3 , p . 265-283. Reyment, R . A . , 1 9 5 9 . N o t e s o n some G l o b i g e r i n i d a e , Globot r u n c a n i d a e and G l o b o r o t a l i i d a e f r o m t h e Upper C r e t a c e o u s and Lower T e r t i a r y o f W e s t e r n N i g e r i a . Rep. G e o l . S u r v e y N i g e r i a , p . 68-86. , 1 9 6 0 . S t u d i e s o n N i g e r i a n Upper C r e t a c e o u s and Lower T e r t i a r y O s t r a c o d a . P a r t 1: S e n o n i a n a n d M q e s t r i c h t i a n Ostrac o d a . S t o c k h o l m C o n t r i b . G e o l . , 7 : l - 2 3 8 , l a m . 23. , 1 9 6 5 . A s p e c t s of t h e Geology o f N i g e r i a . The s t r a t i g r a p y y of t h e C r e t a c e o u s and C e n o z o i c d e p o s i t s . I b a d a n Univ. P r e s s , p . 1-145. , 1966. B r i e f r e v i e w of t h e s t r a t i g r a p h i c s e q u e n c e s of West A f r i c a ( A n g o l a t o S e n e g a l ) . P r o c e e d . 2nd. West A f r i c a n M i c r o p a l e o n t o l o g i c a l C o l l . I b a d a n 1 9 6 5 , p . 162-176. T e r t i a r y SediS a i t o , T . , Ewing, M. and B u r c k l e , L . H . , 1 9 6 6 . ment from t h e m i d - A t l a n t i c R i d g e . S c i e n c e , v . 1 5 1 , no. 3 7 1 4 , p . 1075-1079. S c h a l l e r , H . , 1969, Revisao 7 st r a t i g r A f i c a da b a c i a d e S e r g i p e A l a g o a s . B o l . T e c n . P e t r o b r a s . R i o d e J a n e i r o , v . 1 2 , no. 1, p . 21-86. EsS c h a l l e r , H . , V a s c o n c e l o s , D. N . , and C a s t r o , J . C . , 1 9 7 1 . t r a t i g r a f i a p r e l i m i n a r d a Bacia s e d i m e n t a r d a Foz d o Rio Amaz o n a s . S O C . B r a s i l e i r a d e G e o l . A n a i s XXV C o n g r e s o , v . 3 , p . 189-202. S l a n s k y , M . , 1 9 6 3 . C o n t r i b u t i o n 'a l ' k t u d e g h o l o g i q u e du b a s s i n s & d i m e n t a i r e , c o t i e r du Dahomey e t du Togo. Mkmoires Bureau d e R e c h e r c h e s G e o l o g i q u e s e t M i n . , n o . 11, p . 1-275). S t o l k , J . , 1963. C o n t r i b u t i o n l ' k t u d e des correlations m i c r o f a u n i q u e s du T e r t i a i r e i n f e r i e u r d e la N i g e r i a M e r i d i o n a l e . C o l l . I n t e r n . d e M i c r o p a l . , D a k a r . 17. 247-275. T i n o c o , I . d e M . , 1967. M i c r o p a l e o n t o l o g l a d a f a i x a s e d i m e n t a r c o s t e r a R e c i f e - J o a o P e s s o a . Bol. SOC. B r a s i l e r a G e o l . , v . 1 6 , no. 1, p . 81-85. Todd, R . , and K n i k e r , H . T . , 1 9 5 2 . An Eocene F o r a m i n i f e r a 1 f a u n a f r o m t h e Agua F r e s c a S h a l e o f M a g a l l a n e s P r o v i n c e , S o u t h e r n m o s t C h i l e . Cushman Found. Foram. R e s . S p e c i a l Pub. no. 1, p . 1 - 2 8 . PLATE I Danian assemblaqe Fig.
1
-
Fig. 2
-
Fig. 3
-
Fig. 4
-
Fig. 5
-
Fig. 6
-
Fig. 7 Fig. 8
-
G l o b o c o n u s a d a u b j e r g e n s i s Brgnnimann, x 3 0 8 , San J o r g e B a s i n - S a l a m a n a u i a n S t a a e (. U t m e r D a n i a n ) G l o b o r o t a l i a ( T ; r b o r o t a l i a ) p s e u d o b u l l o i d e s (Plummer) , x 3 4 2 , Neuquen B a s i n - R o c a n i a n S t a g e (Lower D a n i a n ) G l o b o r o t a l i a ( T u r b o r o t a l i a ) c o m p r e s s a (Plummer, x 3 0 8 , San J o r g e B a s i n - S a l a m a n q u i a n S t a g e (Upper D a n i a n ) P u l s i p h o n i n a prima (Plummer) , x 2 0 5 , Neuqu6n B a s i n R o c a n i a n S t a a e (Lower D a n i a n ) S i p h o g e n e r i n o i d e s e l e g a n t u s ( P l u m m e r ) , x 1 7 1 , Neuqu6n B a s i n - R o c a n i a n S t a g e (Lower D a n i a n ) L o x o s t o m o i d e s a p p l i n a e (Plummer) , x 1 5 0 , San J o r g e B a s i n - S a l a m a n q u i a n o S t a g e (Upper D a n i a n ) Alabamina m i d w a y e n s i s B r o t z e n , x 1 7 1 , Neuquen B a s i n R o c a n i a n S t a g e (Lower D a n i a n ) L e n t i c u l i n a k l a g s h a m n e n s i s ( B r o t z e n ) , x 4 1 , San J o r g e B a s i n - Salamanquiano S t a g e (Upper Danian) L
L
436
Fig. 1
-
Fig. 2
-
Fig. 3
-
Fig. 4
-
Fig. 5
-
Fig. 6
-
Fig. 7
-
Fig. 1 Fig. 2
-
Fig. 3
-
Fig. 4
-
Fig. 5
-
Fig. 6
-
Fig. 7
-
PLATE I1 Eocene-Oligocene Assemblage Uvigerina tenuistriata Reuss, x217, Austral Basin Julian Stage (Eocene-Oligocene) Uvigerina abbreviata Terquem, x181, Austral Basin Julian Staqe (Upper Eocene-L. Oliqocene) Cribrorotaiia 1b;nensis Hornibrook , x144 , Austral Basin - Julian Stage (U. Eocene-L. Oligocene) Notorotalia sp., x209, Austral Basin - Julian Stage (U. Eocene-L. Oligocene) Cribrorotalia hornibrooki forma typica Malumian and Masiuk, x 1 1 5 , Austral Basin - Julian Stage (U. EoceneL. Oligocene) Globigerina gortanii Borsetti, x361, Austral Basin Leonian Stage (Lower Oligocene) Globigerina officinalis Subbotina, x361, Austral Basin - Julian and Leonian Stages (Oligocene) PLATE I11 Oligocene Assemblage Globigerina anguliofficinalis Blow, x351, Austral Basin - Leonian Stage (Lower Oligocene) Globigerina ciperoensis Bolli, x281, Austral Basin Leonian Stage (Upper Oligocene) Globorotalia opima Bolli, x351, Austral Basin Leonian Stage (Upper Oligocene) Cribrorotalia s p . , x141, Austral Basin - Leonian Stage (01igocene) Bolivina finlayi Hornibrook, x211, Austral Basin Leonian Stage (Oligocene) Nonionella magnalin ua Finlay, x281, Austral Basin Leonian Stage (Oligzcene) Uvigerina germanina (Cushman and Edwards), x225, Austral Basin - Leonian Stage (Oligocene)
437
438
PLATE I i
439
PLATE I 1 1
This Page Intentionally Left Blank
441
NEOGENE FORAMINIFERA - SOUTH ATLANTIC Alwine Bertels and Marly Madeira-Falcetta Facultad de Ciencias Exactas y Naturales Universidad de Buenos Aires, Rephblica Argentina Consejo Nacional de Investigaciones Cientificas y TLcnicas, RepAblica Argentina Escola de Geoloqia, Universidade Federal do Rio Grande do Sul Porto Alegre, Brazil Resumen En el presente trabajo se sintetizan 10s aportes cientificos efectuados por diversos autores, relacionados con las asociaciones foraminiferol6gicas que prevalecieron durante el Ne6geno en el Atlantic0 Sur. Se analizan diversas cuencas sedimentarias, tanto en el continente sudamericano como en el africano en donde tuvieron lugar depositaciones marinas durante el NeAgeno. Los conjuntos bent6nicos del Mioceno de ambos continentes , el sudamericano y africano, manifiestan escasas caracterlsticas I en comun. Las asociaciones de sudamLrica son aproximadamente comparables a las de la Provincia AtlAntica actual, en tanto que las africanas muestran un marcado endemismo. Los conjuntos p l a n c t d c o s , en cambio, son similares en ambos continentes. De acuerdo con 10s registros obtenidos hastala actualidad, I 10s depositos post Miocenos en el continente americano del sur contienen faunas similares a las actuales de iguales latitudes; las microfaunas africanas, por otra parte, no exhiben el marcado endemismo de Apocas anteriores. Abstract In the present work the scientific contributions related to the foraminifera1 associations which prevailed during the Neoqene in the South Atlantic Ocean made by several authors are synthesized. Several South American and African sedimentary basins in which Neogene deposits are registered are analyzed and the diagnostic taxa are listed. The Miocene South American and West African benthonic assemblages show little features in common; the South American benthonic Foraminifera are approximately comparable to those of the present Atlantic Province whereas the African assemblages
442
show a marked endemism.
Miocene p l a n k t o n i c F o r a m i n i f e r a a r e
comparable i n b o t h c o n t i n e n t s . According t o t h e r e c o r d s o b t a i n e d up t o t h e p r e s e n t , p o s t Miocene S o u t h American d e p o s i t s c o n t a i n m i c r o f a u n a s more s i m i l a r t o e a c h o t h e r t h a n t o t h o s e l i v i n g t o d a y a t t h e same l a t i t u d e s ; t h e West A f r i c a n m i c r o f a u n a s , on t h e o t h e r hand, d o n o t e x h i b i t t h e marked endemism o f o l d e r e p o c h s , Introduction The p u r p o s e o f t h e p r e s e n t work i s t o show t h e most d i a g n o s t i c f o s s i l f o r a m i n i f e r a 1 a s s e m b l a g e s which p r e v a i l e d i n t h e S o u t h A t l a n t i c Ocean d u r i n g Miocene t h r o u g h P l e i s t o c e n e .
Holo-
c e n e f a u n a s w i l l n o t be d i s c u s s e d . The s e d i m e n t a r y b a s i n s i n which Neogene d e p o s i t s o c c u r a r e analyzed i n both c o n t i n e n t s ; t h e f a u n a l assemblages a r e l i s t e d following a l a t i t u d i n a l order according t o t h e sedimentary b a s i n s from which t h e y were r e c o r d e d . The a v a i l a b l e d a t a o b t a i n e d from s e v e r a l o c e a n o g r a p h i c exp e d i t i o n s a r e s y n t h e s i z e d and t h e p r i n c i p a l p l a n k t o n i c zones a r e mentioned. Acknowledgments The a u t h o r s w i s h t o e x p r e s s g r a t i t u d e t o t h e A r g e n t i n a N a t i o n a l C o u n c i l f o r S c i e n t i f i c and T e c h n i c a l R e s e a r c h , f o r t h e e c o n o m i c a l a i d i n t h e accomplishment of t h e p r e s e n t work, and f o r g e n e r o u s l y p r o v i d i n g t h e u s e of t h e J e o l c o JSM
-
U-3 s c a n -
n i n g e l e c t r o n m i c r o s c o p e w i t h which t h e m i c r o g r a p h s w e r e t a k e n . The w r i t e r s a r e a l s o g r a t e f u l t o Buenos A i r e s U n i v e r s i t y , Department of Geology and t o t h e Rio Grande d o S u l U n i v e r s i t y , B r a z i l f o r f a c i l i t i e s p r o v i d e d i n a c h i e v i n g t h e c o m p l e t i o n of t h i s work. P a r t i c u l a r thanks a r e a l s o expressed t o D r . Carlos A. R i n a l d i and t o M r . P a t r i c i o G a n d u g l i a , who k i n d l y and g e n e r o u s l y h e l p e d i n t h e c o n s t r u c t i o n of t h e s t r a t i g r a p h i c c h a r t s and r a n g e c h a r t s which i l l u s t r a t e t h i s p a p e r . The i l l u s t r a t e d m a t e r i a l i s d e p o s i t e d i n t h e L a b o r a t o r y of M i c r o p a l e o n t o l o g y , E s c o l a d e G e o l o g i a , U n i v e r s i d a d e do Rio Grande do S u l , P o r t o A l e g r e , B r a z i l .
443
S o u t h America _ I _
The Amazonas B a s i n i s l o c a t e d i n n o r t h e r n B r a z i l .
Miocene
sediments w e r e deposited i n t o a graben with a southeast-north-
w e s t trend.
The known Miocene l i t h o s t r a t i g r a p h i c u n i t s a r e t h e
Maraj6 and P i r a b a s F o r m a t i o n s and t h e Par:
Group ( T e x t F i g . 1
i n Bertels, Paleogene, t h i s volume). F o r m a t i o n i s composed of c l a s t i c s e d i m e n t s
The Maraj;
which g r a d e i n t o t h e c o n t i n e n t a l m a r g i n t o a t h i c k s e q u e n c e of c a r b o n a t e s which c o n s t i t u t e t h e Amap;
_. a1
1
Formation ( S c h a l l e r
1971). r
The M a r a j o F o r m a t i o n e x t e n d s o v e r t h e whole a r e a of Maraj;
I s l a n d and a l o n g t h e l i t t o r a l a r e a s of w e s t e r n Amap;
e a s t e r n Par:
States.
and
I t o v e r l i e s c o n c o r d a n t l y t h e Lirnoeiro
F o r m a t i o n and i s d i s c o r d a n t w i t h t h e o v e r l y i n g P a r i Group.
The
L i m o e i r o F o r m a t i o n i s c o n s i d e r e k t o b e Upper C r e t a c e o u s t o Low-
er T e r t i a r y ( S c h a l l e r e t a l . , 1 9 7 1 ) , b a s e d o n p a l y n o l o g i c a l data, Petri
( 1 9 5 4 ) a s s i g n e d a Miocene a g e t o t h e s e q u e n c e of t h e
MarajA F o r m a t i o n and f o u n d reworked C r e t a c e o u s and Eocene Foraminifera.
H e considered t h a t Globigerina cretacea d'orbigny,
G l o b i g e r i n a c f . t r i a n g u l a r i s White, G l o b o r o t a l i a compressa (Plummer) , G l o b o r o t a l i a c f . c r a s s a t a (Cushman) and Guembelina g l o b u l o s a E h r e n b e r g w e r e reworked s p e c i e s . F o r t h e Miocene s e q u e n c e P e t r i ( 1 9 5 4 ) r e c o g n i z e d f o u r bent h o n i c f o r a m i n i f e r a 1 zones: 1) A m p h i s t e g i n a l e s s o n i Zone
2) B o l i v i n a p l i c a t e l l a Zone 3 ) E l p h i d i u m poeyanum e l o n g a t a Zone
4) Q u i n q u e l o c u l i n a l a m a r c k i a n a Zone The a g e s a s s i g n e d by S c h a l l e r e t a l .
( 1 9 7 1 ) , however, t o
t h e r e c o g n i z e d members of t h e Maraj6 F o r m a t i o n a r e :
Afui Mem
b e r : P a l e o c e n e ; C u r u r 6 Member; Eocene; Mexiana Member: Eocene t o Lower Miocene and A r a g u a r i Member: Lower Miocene.
Thus a
problem e x i s t s a s t o a g e i n t e r p r e t a t i o n of t h e f o r m a t i o n . The reworked s p e c i e s m e n t i o n e d by P e t r i ( 1 9 5 4 ) i n d i c a t e t h e e x i s t e n c e i n t h e a r e a of C r e t a c e o u s and P a l e o c e n e s t r a t a s u c h a s was r e v e a l e d by p a l y n o l o g i c a l s t u d i e s .
444 The p a r t l y e q u i v a l e n t AmapA F o r m a t i o n o c c u r s i n t h e c o n t i n e n t a l m a r g i n o f t h e Amap;
and P a r ;
States.
The s e d i m e n t a r y
sequences a r e a l g a l m i c r i t e s w i t h i n t e r c a l a t i o n s of g r e e n i s h g r a y c l a y s t o n e s and s i l t s t o n e s , f r e q u e n t l y g l a u c o n i t i c and pyritic. 1
The a g e o f t h e Amapa F o r m a t i o n r a n g e s f r o m t h e P a l e o c e n e t o t h e m i d d l e Miocene
(Schaller e t a l . , 1971):
a ) C a n d i r u S e q u e n c e , o f Eocene a g e I
b ) J a c u n d a S e q u e n c e , t o w h i c h a l a t e Eocene t o O l i g o c e n e a g e was a s s i g n e d . c ) Tambaqui S e q u e n c e : p l a n k t o n i c z o n e s o f G l o b o r o t a l i a m a y e r i and G l o b o r o t a l i a f o h s i p e r i p h e r o a c u t a w e r e i d e n t i f i e d ; t h e a g e of t h e s e c t i o n i s e a r l y t o m i d d l e Miocene. The P a r i Group was d e p o s i t e d i n t h e Amap;
and Par;
States
(Foz d e Amazonas B a s i n ) ; t h e i r s t r a t a h a v e b e e n a l s o b o r e d i n I
t h e c o n t i n e n t a l m a r g i n of t h e P a r a S t a t e . The Par;
Group h a s b e e n s u b d i v i d e d i n t o two f o r m a t i o n s by
Schaller et a l . ,
(1971).
These a r e t h e P i r a r u c u F o r m a t i o n , of
c l a y e y n a t u r e and t h e TucanarA F o r m a t i o n , e s s e n t i a l l y a r e n a ceous.
The g r o u p h a s a f l u v i a t i l e a n d p a r a l i c o r i g i n i n t h e
a r e a s o v e r t h e p r e s e n t c o n t i n e n t , whereas i t i s of n e r i t i c f a c i e s on t h e c o n t i n e n t a l margin.
Throughout t h e whole s e c t i o n
I
of t h e T u c a n a r e F o r m a t i o n Nummulites s p . h a v e b e e n f o u n d ,
The
Zones o f O r b u l i n a u n i v e r s a and G l o b o r o t a l i a m a y e r i h a v e b e e n recognized i n both formations.
Schaller e t al.(1971) attribu-
t e d Miocene t o H o l o c e n e a g e t o t h e s e d i m e n t a r y s e q u e n c e o f t h e Group. The u p p e r p a r t of t h e Amapi and M a r a j o f o r m a t i o n s g r a d e i n t h e s o u t h e r n b o r d e r of t h e Par;
s t a t e i n t o sediments described
p r e v i o u s l y by P e t r i ( 1 9 5 7 ) a s t h e P i r a b a s F o r m a t i o n . The o u t c r o p o f t h e P i r a b a s F o r m a t i o n i s composed o f l i m e s t o n e s and c o q u i n a s w i t h a r e n a c e o u s b e d s a t t h e t o p .
I t i s ex-
p o s e d a l o n g a l a r g e p a r t of t h e l i t t o r a l o f P a r & S t a t e .
Petri
( 1 9 5 7 ) a s s i g n e d a Lower Miocene a g e t o t h e f o r m a t i o n b a s e d o n t h e m i c r o f a u n a l a s s e m b l a g e , a l t h o u g h h e f o u n d many s i m i l a r i t i e s w i t h t h e m i d d l e Miocene a s s o c i a t i o n s from o t h e r r e g i o n s o f t h e w o r l d e s p e c i a l l y t h o s e of t r o p i c a l America.
445
Based o n t h e p r e s e n c e of A r c h a i a s a n g u l a t a ( F i c h t e l and M o l l ) Elphidium poeyanum ( d ' o r b i g n y ) , C a n c r i s s a g r a ( d ' o r b . ) , Eponides r e p a n d u s ( F i c h t e l and M o l l ) , Pyrgo s u b s p h a e r i c a ( d ' Orbigny) and o t h e r s , P e t r i ( 1 9 5 7 ) i n f e r e d s h a l l o w w a t e r d e p o s i t i o n of t h e P i r a b a s F o r m a t i o n . N o g u t i (1975) r e c o r d e d from t h e c o n t i n e n t a l m a r g i n of t h e Amazonas B a s i n t h e Miocene Zones of G l o b i g e r i n a t r i p a r t i t a , Praeorbulina glomerosa, Globorotalia f o h s i peripheroronda, G l o b o r o t a l i a f o h s i p e r i p h e r o c a u t a and G l o b o r o t a l i a m a y e r i . F e r r e i r a (1973) r e c o r d e d t h e o c c u r r e n c e of t h e F i r a b a s F o r m a t i o n ( e a r l y Miocene) i n t h e B a r r e i r i n h a s B a s i n , Maranhao State. I n t h i s Basin t h e P i r a b a s Formation i s developed over t h e whole b a s i n d i s c o r d a n t l y o v e r l y i n g C r e t a c e o u s s e d i m e n t s and c o v e r e d by non-marine and l i t t o r a l Q u a t e r n a r y d e p o s i t s . The P i r a b a s F o r m a t i o n i n t h i s b a s i n i s composed of c a l c a r e o u s m a r l s and m a r l y c h a l k s a t t h e b a s e p a s s i n g i n t o b l u e c l a y s t o n e s , f o s s i l i f e r o u s l i m e s t o n e s , g r a y c l a y s t o n e s and c u l minating i n t o beige limestones.
These l i m e s t o n e s c o n t a i n l a r g e
Foraminifera recorded a s "Orbitoid foraminifera." From t h e c o n t i n e n t a l margin of t h e S e r g i p e / A l a g o a s B a s i n Noguti (1975) r e c o r d e d t h e Zone of G l o b i q e r i n a t r i p a r t i t a . The c o n t i n e n t a l m a r g i n of t h e E s p i r i t o S a n t o B a s i n y i e l d e d a p l a n k t o n i c a s s o c i a t i o n i n which t h e P l a n k t o n i c Zones of
e-
b i g e r i n a t r i p a r t i t a , P r a e o r b u l i n a g l o m e r o s a and G l o b o r o t a l i a m a y e r i were r e c o g n i z e d by Noguti ( 1 9 7 5 ) . I n t h e c o n t i n e n t a l m a r g i n of Rio d e J a n e i r o ( s o u t h of t h e Campos B a s i n ) Noguti (1975) found t h e Miocene P l a n k t o n i c Zones of G l o b i g e r i n a t r i p a r t i t a and P r a e o r b u l i n a g l o m e r o s a . The n o r t h e a s t e r n B r a z i l i a n Reconcavo B a s i n c o n t a i n s Miocene s e d i m e n t s whose m i c r o f a u n a l c o n t e n t was s t u d i e d by P e t r i ( 1 9 7 2 ) . The a b s e n c e of r e p r e s e n t a t i v e s o f g e n e r a of t h e M i l i o l i d a e and t h e genus Elphidium i n d i c a t e a n o f f s h o r e e n v i r o n m e n t . S o u t h of B a h i a ( p o s s i b l y Reconcavo B a s i n ) Noguti (1975) r e c o r d e d t h e Zones of G l o b i g e r i n a t r i p a r t i t a and P r a e o r b u l i n a glomerosa of e a r l y Miocene a g e . I n t h e s o u t h e a s t e r n B r a z i l i a n P e l o t a s B a s i n (Closs, 1 9 6 7 , 1 9 7 0 ; F e r n a n d e s , 1975 M S ; T h i e s e n , 1975 M S ;
and F a l c e t t a , 1 9 7 6
446
MS) t h i c k s e q u e n c e s o f Miocene t o P l e i s t o c e n e a g e were d e p o s i t e d . The P e l o t a s B a s i n i s of t e c t o n i c n a t u r e ; t h e s e d i m e n t a r y s e q u e n c e s a r e found i n s u b s u r f a c e and o v e r l y t h e g r a n i t i c b a s e The s e d i m e n t s a r e m a i n l y composed of s a n d s t o n e s w i t h
ment.
i n t e r c a l a t e d c l a y s t o n e s ; f a c i e s changes are observed n e a r t h e b o r d e r s of t h e b a s i n i n which c o n g l o m e r a t i c and a r k o s i c l e v e l s occur. The p r e d o m i n a n t m i c r o f a u n a l a s s e m b l a g e s t u d i e d by C l o s s ( 1 9 6 7 , 1970) i s composed of G l o b i g e r i n o i d e s b i s p h a e r i c u s , l i n a s u t u r a l i s , Globorotalia mayeri, Globorotalia obesa,
ee-
b i g e r i n o i d e s g l o m e r o s u s , G l o b i g e r i n o i d e s t r a n s i t o r i u s , Globorot a l i a f o h s i and --
( o p . )&c
Globorotalia menardii archaeomenardii.
Closs
c o n s i d e r e d of i m p o r t a n c e t h e e v o l u t i o n a r y t r e n d of
Globigerina bisphericus
- 2.
s u t u r a l i s although he pointed out
the d i f f i c u l t i e s i n establishing a definite correlation with Miocene S t a g e s . F e r n a n d e z ( 1 9 7 5 , MS) s t u d i e d t h e u v i g e r i n i d s o f t h i s b a s i n and found s e v e r a l s p e c i e s of U v i g e r i n a .
Boll-
v i n i d s a r e a l s o r e c o r d e d from t h e p e l o t a s B a s i n m a r i n e s e q u e n c e by T h i e s e n ( 1 9 7 5 , MS). R e c e n t work of o n e of t h e w r i t e r s (M-M.F.)
b a s e d on t h e
p r e s e n c e of G l o b o r o t a l i a a c o s t a e n s i s , G l o b i g e r i n a n e p e n t h e s and o t h e r p l a n k t o n i c s p e c i e s , found t h a t t h e t r a n s g r e s s i o n i n t h e a r e a t o o k p l a c e i n l a t e Miocene t i m e s , i . e . , a p p r o x i m a t e l y a t Zone N 1 4 of Blow ( 1 9 6 9 ) . The O r b u l i n a s e r i e s mentioned by C l o s s ( 1 9 6 7 , 1 9 7 0 ) a s w e l l a s o t h e r Lower Miocene a s s e m b l a g e s a r e p r o b a b l y reworked from a d j a c e n t a r e a s . I n A r g e n t i n a t h e n o r t h e a s t e r n and c e n t r a l P a r a d B a s i n o r Chaco-Paranense B a s i n c o n t a i n s s h a l l o w w a t e r m a r i n e d e p o s i t s of t h e E n t r e R i o s and P a r a & f o r m a t i o n s ( E n t r e r r i a n o a n d / o r P a r a n i a n o s t a g e s ) of l a t e M i o c e n e - e a r l y
Pliocene? age. I
The s e d i m e n t s of t h e E n t r e R i o s and P a r a n a f o r m a t i o n s i n t h e P a r a n i b a s i n e x t e n d i n l a n d o v e r a l a r g e a r e a . S u r f a c e exp o s u r e s a r e found a t t h e e a s t and i n s u b s u r f a c e w e s t w a r d s .
The
s e d i m e n t a r y s e q u e n c e i s composed o f c l a y s t o n e s a t t h e b a s e and s a n d s t o n e s and c o q u i n a s t o t h e t o p . The f a u n a l a s s e m b l a g e a s w e l l a s t h e l i t h o l o g i c a l f e a t u r e s
447
indicate shallow waters and an almost closed marine environment. This condition may be due to the Precambrian shield located at the eastern border of the basin. During Miocene-Pliocene times the Parani Basin was probably connected with the Salado Basin which might have acted as a route for the large marine transgression. The microfaunal assemblage described by Pisetta (1968, MS) is composed of Buccella frigida (Cushman), Ammonia beccarii (Linn&) and Protelphidium tuberculatum (d'Oribigny). In the Salado Basin Malumian (1970a) studied subsurface sedimentary sequences. The Entrerrian Stage was found to be represented by Globorotalia pachyderma forma sinistrorsa, pI0telphidium tuberculatum, Cibicides berthelotti, Cibicides pseudoungerianus, and others. The age of the Entrreriano Stage was considered to be of late Miocene Age (Malumian 1970a). The microfaunal assemblage indicates warmer sea water conditions than that of today at the same latitude. Sedimentary subsurface sequences were also studied by Malumian (1972) in the Colorado Basin. The assemblage found by this author is listed in Text Fig. 1. The assigned age was Oligocene/Miocene based on the planktonics G&bigerina woodi woodi, Cassigerinella chipolensis etc. Malumian (1972) found that the planktonic assemblage is equivalent to the Globigerina ciperoensis ciperoensis and Globorotalia opima opima from tropical areas, and to Globigerina woodi woodi and Globigerina euapertura from temperate regions. The Austral Basin is the southernmost South American Basin. The sedimentary sequences range in age from the Valanginian to the Miocene. Miocene Foraminifera were recorded by Malumian (1968) who found an assemblage in subsurface samples. Subsequent studies carried out by Malumian, Masiuk and Riggi (1971) in the Austral Basin confirmed the marine Miocene section in this basin. The following Miocene Foraminifera were recorded: Buccella frigida Cushman, Nonion affine (Cushman), Pullenia subcarinata d'Orbigny, Nonionella atlantica Cushman, Lenticulina rotulata Lamark, Cibicides pseudoungerianus
448 (Cushman) and G l o b i g e r i n a e x . g r . p r a e b u l l o i d e s Blow. Africa I n t h e A f r i c a n c o n t i n e n t s e q u e n c e s r e f e r r e d t o t h e Miocene a r e p r e s e n t i n N i g e r i a , Cameroon, Gabon, Angola a n d S o u t h A f r i -
ca. I n Nigeria
(Reyment, 1 9 6 5 ) Miocene s t r a t a a r e e x p o s e d i n
t h e western region; t h e recognized l i t h o s t r a t i g r a p h i c u n i t s are t h e non-marine Ogwashi-Asaba F o r m a t i o n a n d t h e m a r i n e I j e b u Formation.
The l a s t i s l o c a t e d e a s t o f Lagos; a f t e r Reyment
( o p . c i t . ) i t i s p a r t l y m a r i n e and may b e p a r t l y e q u i v a l e n t t o t h e non-marine Ogwashi-Asaba F o r m a t i o n .
The b e d s a r e p o o r l y
e x p o s e d and c o n s i s t s m a i n l y o f c l a y s and s a n d s , i m p r e g n a t e d w i t h bitumen.
The m a r i n e f a c i e s c o n t a i n b e n t h o n i c F o r a m i n i f e r a .
Miocene s e q u e n c e s a r e a l s o f o u n d i n b o r e h o l e s w e s t of Lagos i n w e s t e r n N i g e r i a ; t h e s t r a t a c o n t a i n E p o n i d o p s i s e s h i r a I
d e K l a s z and R e r a t , Daucina e r m a n i a n a Bornemann and D a u c i n o i d e s c i r c u m t e g e n s d e K l a s z and R & a t
(Reyment, 1 9 6 5 ) .
Although t h e I j e b u marine beds l a c k p l a n k t o n i c Foraminif e r a , i n Gabon E p o n i d o p s i s e s h i r a d e K l a s z and R h a t
i s asso-
c i a t e d w i t h G l o b o r o t a l i a o f t h e f o h s i g r o u p which would i n d i c a t e a e a r l y - m i d d l e Miocene a g e . O t h e r t y p i c a l s p e c i e s which o c c u r i n Gabon a r e p r e s e n t i n s o u t h e a s t e r n N i g e r i a such as Megastomella a f r i c a n a compressa F a u l k n e r , d e K l a s z and R & a t , L e Calvez and R & a t
Nonion c e n t r o s u l c a t a d e K l a s z ,
and G a v e l i n e l l a b e n i n e n s i s K l a s z , L e C a l v e z
and R & r a t . I n Cameroon Miocene s e d i m e n t s r e s t d i s c o r d a n t l y on o l d e r stratigraphic units. I n t h e Duala B a s i n t h e Miocene i s r e p r e s e n t e d by s a n d s t o n e s a n d c l a y s t o n e s , w i t h m i o g y p s i n i d s , i n t e r c a l a t e d by ba-
s a l t s (Belmonte, 1 9 6 6 ) . I n t h e c e n t r a l p a r t of t h e B a s i n t h e Miocene r e a c h e s cons i d e r a b l e t h i c k n e s s and i s s u b d i v i d e d by Belmonte ( 1 9 6 6 ) i n t o : a ) b a s a l b e d s c o n s i s t i n g of c l a y s t o n e s w i t h i n t e r c a l a t e d sandstones. b ) a r e g r e s s i v e s e q u e n c e p r e d o m i n a n t l y d e t r i t i c which beg i n s w i t h n e a r l y marine s t r a t a and ends w i t h d e l t a i c deposits.
Belmonte ( o p . c i t . ) a s s i g n e d t o t h e s e M i o -
449
cene s t r a t a a 3urdigalian-Vindobonian age. I n W e s t e r n Cameroon a l i m e s t o n e i n t h e M i s s e l l e l e R i v e r (Reyment, 1 9 6 5 ) h a s y i e l d e d d i a g n o s t i c m i o g y p s i n i d s s u c h a s Miogypsina ( M i o l e p i d o c y c l i n a ) b u r d i g a l e n s i s G b b e l , M_. n e g r i i ( F e r r e r o ) , M_.
fi. (M.)
bantamensis
(M.)
(Miogypsinoides) complanata Schlumberger, (Tan S i n Hok) and M_.
(z.)n i g e r i a n a
Kupper.
Drooyer ( 1 9 6 6 ) r e c o r d e d from d e e p w e l l s i n Cameroon app r o x i m a t e l y t h e same a s s o c i a t i o n of l a r g e F o r a m i n i f e r a a s from outcrops i n t h e Missellele River.
Drooger ( o p . c i t . ) s u g g e s t s
a B u r d i g a l l a n age f o r s t r a t a c o n t a i n i n g Miogypsinoides bantam e n s i s Tam and
M.
b u r d i g a l e n s i s Gimbel.
A c c o r d i n g t o Reyment ( 1 9 6 5 ) t h e b e d s of t h e M i s s e l l e l e R i v e r may b e p a r t of t h e Mpundu F o r m a t i o n . F a u l k n e r , d e K l a s z and R & a t
(1963) have r e c o r d e d M e g a -
s t o m e l l a a f r i c a n a from a b o r e h o l e a t Uquo, 1 . 5 km s o u t h w e s t of C a l a b a r , w h i c h i n d i c a t e s t h e p r e s e n c e of Miocene and p e r m i t s c o r r e l a t i o n w i t h t h e Mandorov;
F o r m a t i o n of Gabon of e a r l y ?
Miocene a g e . I n t h e Chang r e g i o n a r e l i g n i t i c b e d s which i n some l e v e l s contain Foraminifera. Gaze1 e t a l . ( 1 9 5 6 , c i t e d i n Reyment, 1 9 6 5 ) s u g g e s t e d a
--
p o s s i b l e c o r r e l a t i o n o f t h e s e b e d s w i t h t h e non-marine OgwashiAsaba F o r m a t i o n of N i g e r i a . I n Gabon t h e m a r i n e Miocene s e q u e n c e s a r e e x p o s e d i n t h e v i c i n i t y of P o r t G e n t i l , a l o n g s i d e t h e p r e s e n t c o a s t l i n e , where Miocene s t r a t a unconformably o v e r l i e Eocene s e q u e n c e s . The s e d i m e n t a r y b a s i n a p p e a r s t o b e made of two s e p a r a t e d u n i t s , d i v i d e d by a n i n t e r m e d i a t e h i g h c a l l e d t h e Lambarene h o r s t ( B e l m o n t e , H i r t z and Wenger, i n Reyment, 1 9 6 5 ) . The Neogene s e q u e n c e of Gabon seems t o b e s i m i l a r t o t h a t of N i g e r i a and t h e r e a r e numerous F o r a m i n i f e r a i n common. I n W e s t e r n Gabon t h e r e c o g n i z e d c h r o n o s t r a t i g r a p h i c u n i t s a r e ( L e C a l v e z , d e K l a s z and B r u n , 1 9 7 4 ) : Mandorov&, M'Bega and N'Tchengu;! which c o n s t i t u t e t h e Alewana S y s t e m . The Akosso S t a g e c o m p r i s e s s t r a t a b e l o n g i n g t o t h e Miocene and y o u n g e r deposits. Many i m p o r t a n t c o n t r i b u t i o n s d e a l w i t h t h e m i c r o p a l e o n t o l o g y of t h e Gabon B a s i n ( c f . d e K l a s z and R h r a t , 1 9 6 2 ; d e K l a s z and G a g e o n n e t , 1 9 6 3 ; d e K l a s z , L e C a l v e z and R e h a t , 196433;
450
F a u l k n e r , d e K l a s z a n d R Q r a t , 1 9 6 3 ; d e K l a s z , L e C a l v e z and R & r a t , 1 9 6 4 a , c ; Graham d e K l a s z and R k r a t ,
1 9 6 5 ; d e K l a s z and
Micholet, 1970). The Mandorov;!
F o r m a t i o n i s d e t r i t a l and c l a y e y ; b a s e d on
t h e p l a n k t o n i c c o n t e n t , d e K l a s z and M i c h o l e t ( 1 9 7 0 ) a s s i g n e d a Lower B u r d i g a l i a n a g e t o t h i s f o r m a t i o n . c o r d e d f r o m t h e Upper Mandorov;!
T h e s e a u t h o r s re-
S e r i e s t h e p l a n k t o n i c s Globoro-
talia fohsi barisanensis, Globorotalia fohsi -t a l i a fohsi lobata. Taking i n t o a c c o u n t t h e ---
f o h s i and Globorop l a n k t o n i c zona-
t i o n p r o p o s e d by Blow ( 1 9 6 9 ) t h e c o r r e s p o n d i n g s t r a t a a r e o f e a r l y - m i d d l e Miocene a g e c o m p r i s i n g z o n e s N 1 2 t o N13. The M'Bega F o r m a t i o n , w h i c h o v e r l i e s t h e Mandorov;? Format i o n i s i n g e n e r a l clayey w i t h b a s a l d e t r i t a l beds. assemblages a r e found i n t h e d e t r i t a l b e d s .
Arenaceous
Most o f t h e s p e -
c i e s f o u n d i n t h e M'Bega F o r m a t i o n a r e common t o t h e u n d e r l y i n g MandorovE! F o r m a t i o n . The f o l l o w i n g s p e c i e s a p p e a r i n M'Bega: Nonion boueanum ( d ' o r b i g n y ) , C a s s i d u l i n a l a e v i g a t a c a r i n a t a Cushman, P u l l e n i a t i n a o b l i q u i l o c u l a t a P a r k e r a n d J o n e s a n d G l o b o r o t a l i a m e n a r d i i ( d ' o r b i g n y ) miocenica Palmer. D e K l a s z and G a g e o n e t
( 1 9 6 3 ) b a s e d o n t h e l a r g e number o f
common F o r a m i n i f e r a t o b o t h Mandorov;? a n d M'Bega F o r m a t i o n s ass i g n e d a n e a r l y Miocene a g e t o t h e M'Bega F o r m a t i o n . The l o w e r p a r t o f t h e N'Tchengu;? F o r m a t i o n i s a r g i l l a c e o u s , w h i l e t h e upper p a r t i s d e t r i t a l w i t h c l a y e y i n t e r c a l a tions. The f o r a m i n i f e r a 1 c o n t e n t i s m a i n l y l i k e t h a t o f t h e und e r l y i n g Mandorov;!
and M'Bega F o r m a t i o n s ; o n l y a few o t h e r s p e c i e s a p p e a r i n t h e N'Tchengu;? F o r m a t i o n s u c h a s : P t y c h o m i l i o l a s e p a r a n s ( B r a d y ) and E l p h i d i u m c f . c r i s p u m ( L i n n 6 )
.
The m o s t c h a r a c t e r i s t i c F o r a m i n i f e r a o f N'Tchengu;
are
s p e c i e s of A m p h i s t e g i n a w h i c h a r e f o u n d t o g e t h e r w i t h r a r e Operculinoides. D e K l a s z a n d G a g e o n n e t ( 1 9 6 3 ) s u g g e s t a l a t e Miocene a g e
f o r a t l e a s t p a r t o f t h e s t r a t a o f t h e N'TchenguE! F o r m a t i o n . I n Angola t h e known Miocene s e q u e n c e s a r e e x p o s e d i n t h e Cuanza B a s i n and i n t h e B e n g u e l a B a s i n , Brognon and V e r r i e r
451 (1966). sin.
No O l i g o c e n e s e d i m e n t s a r e p r e s e n t i n t h e Cuanza Ba-
The Miocene i s r e p r e s e n t e d o n l y by s e d i m e n t s o f A q u i t a n i -
a n and B u r d i g a l i a n a g e . A t t h e b a s e o f t h e Miocene s e c t i o n i s r e c o g n i z e d t h e Lower
Q u i f a n g o n d o F o r m a t i o n w h i c h i s b l a c k o r v a r i e g a t e d , h i g h l y gyps i f e r o u s and u n f o s s i l i f e r o u s .
I t r e s t s unconformably on e a r l y ,
m i d d l e o r l a t e E o c e n e , o r i n some p l a c e s , on Campanian s t r a t a (Brognon a n d V e r r i e r , 1 9 6 6 ) .
Above t h e u n f o s s i l i f e r o u s s t r a t a ,
b l a c k t o brown s h a l e s w i t h G l o b i g e r i n a d i s s i m i l i s a r e r e c o g n i zed. Edllowing t h e s t r a t a of t h e
5. d i s s i m i l i s
Zone, t h e Upper
Quifangondo w a s d e p o s i t e d , c o n s i s t i n g o f s i l t y s h a l e , v e r y r i c h i n F o r a m i n i f e r a , and w h i c h g r a d e s upward i n t o c o q u i n o i d l i m e s t o n e , w i t h sandy l i m e s t o n e i n t e r b e d d e d .
T h i s member c o r r e -
s p o n d s w i t h t h e G l o b i g e r i n a t e l l a i n s u e t a Zone (Brognon and Verrier, 1966).
The Luanda F o r m a t i o n h a s brown s h a l e s a t i t s b a s e , cont a i n i n g F o r a m i n i f e r a o f t h e G l o b o r o t a l i a f o h s i Zone; upwards t h i s f o r m a t i o n i s a b r u p t l y t r a n s i t i o n a l t o l i t t o r a l and d e l t a i c sand w i t h sands tone i n t e r b e d s t h a t c l o s e t h e c y c l e .
The m a r i n e
s e d i m e n t a r y c y c l e b e g a n d u r i n g e a r l y A q u i t a n i a n t i m e a n d ended d u r i n g B u r d i g a l i a n t i m e (Brognon a n d V e r r i e r 1 9 6 6 ) . The Miocene Cacuaco F o r m a t i o n w a s d e p o s i t e d i n s h a l l o w wat e r e n v i r o n m e n t w h e r e l i m e s t o n e w i t h a l g a e , b i v a l v e s and e c h i noids w a s deposited. I n t h e B e n g u e l a B a s i n , Antunez
( 1 9 6 4 ) r e c o r d e d Neogene se-
d i m e n t s c o n s i s t i n g o f s a n d s t o n e s , c l a y s t o n e s , c h a l k s and g y p s i f e r o u s m a r l s i n which s p e c i e s of G l o b i g e r i n o i d e s w e r e found. F o r a m i n i f e r a 1 a s s e m b l a g e s r e c o r d e d by Rocha and F e r r e i r a ( 1 9 5 7 ) a r e composed o f b e n t h o n i c s and t h e p l a n k t o n i c s G l o b i g e r -
-
-~
ina bulloides d'orbigny, Globigerinoides trilobus cf. irregul a r i s L e Roy, B i o r b u l i n a b i l o b a t a
( d ' o r b i g n y ) , Orbulina
e-
r a l i s Brcnnimann, 0. u n i v e r s a d ' 0 r b i g n y and G l o b o r o t a l i a tumida ( B r a d y ) ; t h i s a s s e m b l a g e r e p r e s e n t s a Miocene a g e (Rocha and
.
F e r r e i r a , op. c i t . ) Only a f e w s p e c i e s o f Angola s u c h a s E p o n i d o p s i s e s h i r a K l a s z and R & a t
a r e common t o N i g e r i a and Gabon.
The r e s t of
t h e r e c o r d e d s p e c i e s a r e d i f f e r e n t t o t h o s e of o t h e r a r e a s .
452 P 1i o c e n e. A s f a r a s information i s a v a i l a b l e , marine Pliocene i s not
r e p r e s e n t e d o n t h e p r e s e n t b o r d e r l a n d o f S o u t h America; i n Af-
r i c a , s e d i m e n t s y o u n g e r t h a n Miocene a r e r e f e r r e d t o a s " P o s t Miocene.
"
Pleistocene P l e i s t o c e n e m a r i n e s e d i m e n t s are exposed i n t h e R e c i f e / J o z o P e s s o a B a s i n and i n t h e R i o Grande C o a s t a l P l a i n , B r a z i l , i n Uruguay and i n A r g e n t i n a . From t h e B r a z i l i a n R e c i f e / J o g o P e s s o a B a s i n T i n o c o ( 1 9 5 8 ) d e s c r i b e d F o r a m i n i f e r a f o u n d i n s u b s u r f a c e s a m p l e s i n Pernambuco S t a t e , n o r t h e a s t e r n B r a z i l . The Q u a t e r n a r y s e d i m e n t s o v e r l i e t h e Upper C r e t a c e o u s I t a marac;
and G r a m a m e F o r m a t i o n s and P a l e o c e n e s e d i m e n t s o f t h e
Maria F a r i n h a F o r m a t i o n . The Q u a t e r n a r y a s s e m b l a g e i s m a i n l y composed of: l a r i a gramen --
d ' o r b i g n y , abundant m i l i o l i d s ,
Textu-
Robulus r o t u l a t u s
(Lamarck) , P e n e r o p l i s b r a d y i Cushman, P e n e r o p l i s p e r t u s u s
( F o r s k a l ) , Archaias angulatus
( F i c h t e l and M o l l )
,
Bolivina
E-
p a c t a Sidebottom, B o l i v i n a t o r t u o s a Brady, B o l i v i n a p s e u d o p l i -
c
s H e r o n - A l l e n and E a r l a n d , B o l i v i n a v a r i a b i l i s ( W i l l i a m s o n ) ,
B o l i v i n a s t r i a t u l a Cushman, D i s c o r b i s - m obtusa
( d ' o r b i g n y ) , Ammonia b e c c a r i i p a r k i n s o n i a n a
Ammonia b e c c a r i i skoy)
,
Cushman, D i s c o r b i s
(Linn;),
Buccella f r i g i d a
B u c c e l l a p e r u v i a n a campsi
(d'orbigny), (Boltov-
(Cushman) , R e u s e l l a s p i n u l o s a ( R e u s s ) ,
~-
P a t t e l l i n a c o r r u g a t a Williamson, Amphistegina l e s s o n i d ' o r b i g n y , ( B r a d y ) , Nonion a f f i n e ( R e u s s ) , N o n i o n e l l a
Loxostomum limbatum
a t l a n t i c a Cushman, E l p h i d i u m d i s c o i d a l e
( d ' o r b i g n y ) , Elphidium
advenum d e p r e s s u l u m Cushman, E l p h i d i u m i n c e r t u m ( W i l l i a m s o n ) , E l p h i d i u m e x c a v a t u m (Terquem) , E l p h i d i u m g a l v e s t o n e n s e Kornf e l d , E l p h i d i u m sagrum ( d ' o r b i g n y ) , E l p h i d i u m g u n t e r i C o l e , Cibicides bertheloti
(d'orbigny)
Cushman, C i b i c i d e s a c k n e r i a n u s
,
C i b i c i d e s pseudoungerianus
(d'Orbigny), Dyocibicides
s-
s e r i a l i s Cushman a n d V a l e n t i n e , C a s s i d a l i n a l a e v i g a t a d ' 0 r b i g ny, C a s s i d u l i n a sublobosa Brady, P l a n o r b u l i n a m e d i t e r r a n e a d ' o r b i g n y , Astrononion s t e l l i g e r u m ( d ' o r b i g n y ) Globigerina bulloides d'0rbigny
,
G l o b i g e r i n a pachyderma ( E h r e n b e r g ) and
Globigerinoides trilobus
(Reuss)
.
The p r e s e n c e o f a b u n d a n t M i l i o l i d s and t h e g e n u s E l p h i d i u m
453
i n d i c a t e s a s h a l l o w w a t e r e n v i r o n m e n t w h e r e a s t h e t o t a l assemb l a g e i n d i c a t e warm w a t e r s , e s p e c i a l l y by t h e p r e s e n c e of r e p r e s e n t a t i v e s o f P e n e r o p l i s , A r c h a i a s and Amphistegina ( T i n o c o , 1958). P l e i s t o c e n e and Holocene s e q u e n c e s i n s o u t h e r n B r a z i l cons i s t s of m a r i n e , e o l i a n and l a g o o n a l d e p o s i t s .
S u b s u r f a c e s a m p l e s from E l Chui w e l l were s t u d i e d by C l o s s and Madeira ( 1 9 6 8 ) .
The F o r a m i n i f e r a 1 a s s e m b l a g e i s composed
o f : T e x t u l a r i a gramen d ' o r b i g n y , Q u i n y u e l o c u l i n a seminulum (LinnL) , Ammonia b e c c a r i p a r k i n s o n i a n a ( d ' o r b i g n y ) , Elphidium d i s c o i d a l e ( d ' o r b i g n y ) , Elphidium excavatum (Terquem) , E l p h i d i um g a l v e s t o n e n s e K o r n f e l d , E l p h i d i u m g u n t e r i C o l e , Elphidium
s e l s e y e n s e (Heron-Allen -
and E a r l a n d ) , Amphistegina l e s s o n i
d ' o r b i g n y , C i b i c i d e s . b e r t h e l o t i ( d ' o r b i g n y ) , C i b i c i d e s pseudou n g e r i a n u s (Cushman) , C a s s i d u l i n a l a e v i g a t a d ' 0 r b i g n y and N o n i o n e l l a a t l a n t i c a Cushman. I n t h e P a l m a r e s do S u l l o c a l i t y , s o u t h e r n B r a z i l , a w a t e r w e l l y i e l d e d P l e i s t o c e n e o s t r a c o d e and f o r a m i n i f e r a 1 assemb l a g e s ( B e r t e l s , F a l c e t t a and K o t z i a n , 1 9 7 6 , i n p r e s s ) . The f o r a m i n i f e r a 1 a s s e m b l a g e i s mainly composed of t h e same s p e c i e s , e x c l u d i n g Amphistegina l e s s o n i , a s t h o s e d e s c r i b e d from t h e E l Chui w e l l by C l o s s and Madeira ( 1 9 6 8 ) . Throughout t h e v e r t i c a l s e c t i o n t h e a s s e m b l a g e shows mar i n e , m i x o h a l i n e and f r e s h w a t e r f a u n a : t h e s e f a u n a l changes p r o b a b l y c o u l d be r e l a t e d t o t h e P l e i s t o c e n e f l u c t u a t i o n s of sea l e v e l i n t h a t a r e a i n response t o possible g l a c i a l stages and i n t e r s t a g e s . The o u t c r o p s o f t h e m a r i n e P l e i s t o c e n e Chui F o r m a t i o n a r e disposed p a r a l l e l t o t h e p r e s e n t c o a s t l i n e ; l i t h o l o g i c a l l y it
i s composed of s e m i c o n s o l i d a t e d q u a r t z i t i c s a n d s w i t h p e l i t i c intercalations. The Holocene m a r i n e s e d i m e n t s s o f a r have n o t y i e l d e d Foraminifera i n southern Brazil. I n Uruguay t h e P l e i s t o c e n e F r a y B e n t o s , Camacho and E l Chui f o r m a t i o n s a r e r e c o g n i z e d . I n Western Uruguay t h e F r a y B e n t o s , of non-marine o r i g i n and t h e m a r i n e Camacho F o r m a t i o n a r e d i s t i n g u i s h e d . The m a r i n e Camacho F o r m a t i o n i s composed o f w h i t e , w e l l s o r t e d s a n d s , g r e e n i s h o r y e l l o w i s h c l a y and c o q u i n a .
The
454
f o l l o w i n g F o r a m i n i f e r a a r e r e c o r d e d by Delaney ( 1 9 6 7 ) : b e c c a r i i (Linn;),
Ammonia
Ammonia b e c c a r i i p a r k i n s o n i a ( d ' o r b i g q y ) ,
z-
l i v i n a s t r i a t u l a (Cushman) , B u c c e l l a f r i g i d a (Cushman) , Buccel-
l a peruviana
campsi ( B o l t o v s k o y ) , E l p h i d i u m d i s c o i d a l e d ' O r b i g n y , Elphidium excavatum (Terquem) and Q u i n q u e l o c u l i n a seminulum ( L i n n A ) . The m a r i n e Chui F o r m a t i o n h a s a l i m i t e d e x t e n t and i s conf i n e d t o n o r t h e a s t e r n Uruguay ( D e l a n e y , 1 9 6 7 ) . c o n s i t of w e l l - s o r t e d ,
Chui s e d i m e n t s
p r e d o m i n a n t l y q u a r t z s a n d o f medium
grain size. N o F o r a m i n i f e r a have been r e c o r d e d from t h i s f o r m a t i o n ,
b u t t a k i n g i n t o a c c o u n t t h a t i t i s t h e same s t r a t i g r a p h i c u n i t as t h a t o f s o u t h e r n B r a z i l , a s i m i l a r m i c r o f a u n a l a s s o c i a t i o n i s t o be e x p e c t e d .
I n Argentina t h e p r i n c i p a l m a n i f e s t a t i o n s of Quaternary i n g r e s s i o n s a r e r e p r e s e n t e d by t h e Pampeano and Post-Pampeano Series.
The Pampeano S e r i e s c o m p r i s e s t h e Ensenadense and
B o n a e r e n s e S t a g e s ; t h e Post-Pampeano S e r i e s c o m p r i s e s t h e B e l g r a n i a n o , Q u e r a n d i n i a n o and P l a t i a n o S t a g e s ( F i d a l g o , d e Franc e s c o and P a s c u a l , 1 9 7 5 ) . The Ensenadense a s s e m b l a g e from t h e Quequ6n l o c a l i t y , Buenos A i r e s P r o v i n c e , s t u d i e d by Boltovskoy ( 1 9 5 9 ) , i s comp o s e d of t h e same s p e c i e s a s t h o s e l i v i n g t o d a y a t t h e same l a titude. The f o r a m i n i f e r a 1 a s s e m b l a g e of t h e P l a t i a n S t a g e a t Mar C h i q u i t a , Buenos A i r e s P r o v i n c e ( S u a r e z S o r u c o , 1968 MS) i s composed o f :
Cyclogyra i n v o l v e n s ( R e u s s ) , Q u i n q u e l o c u l i n a
lam-
a r c k i a n a d ' o r b i g n y , Q u i n q u e l o c u l i n a p a t a g o n i c a d ' o r b i g n y , Q&g u e l o c u l i n a seminulum ( L i n n 6 ) , P y r g o n a s u t a Cushman, Pyrgo p e r uviana (d'orbigny)
, Pyrgo
r i n g e n s (Lamarck) , M i l i o l i n e l l a
*-
r o t u n d a (Montagu) , O o l i n a hexagona ( W i l l i a m s o n ) , O o l i n a melo
d ' o r b i g n y , B o l i v i n a s t r i a t u l a Cushman, B o l i v i n a v a r i a b i l i s ( W i l l i a m s o n ) , B u l i m i n a p a t a g o n i c a d ' o r b i g n y , B u l i m i n e l l a ele g a n t i s s i m a ( d ' o r b i g n y ) , B u c c e l l a f r i g i d a (Cushman) , Neoconorbina o r b i c u l a r i s --
(Terquem) , Ammonia b e c c a r i i p a r k i n s o n i a n a
-
( d ' o r b i g n y ) , Elphidium a r t i c u l a t u m ( d ' o r b i g n y )
, Elphidium
d i s c o i d a l e d ' 0 r b i g n y and C i b i c i d e s v a r i a b i l i s
(d'orbigny).
The a d d i t i o n a l s p e c i e s r e c o r d e d by B e r t e l s (1976b, i n p r e s s ) from t h e P l a t i a n S t a g e a r e :
Elphidium m a r g a r i t a c e o u m
455
Cushman, Bulimina m a r g i n a t a d ' o r b i g n y , B o l i v i n a p s e u d o p l i c a t a Heron-Allen and E a r l a n d , D i s c o r b i s f l o r i d a n u s Cushman, D i s c o r @ p e r u v i a n u s d ' o r b i g n y , D i s c o r b i n o p s i s a g u a y o i Bermudez and Cibicides dispars (d'orbigny)
.
P o s t Miocene of A f r i c a I n t h e A f r i c a n c o n t i n e n t Q u a t e r n a r y f o r m a t i o n s a r e found normally i n r i v e r v a l l e y s o r l i t t o r a l a r e a s . I n Dahomex, P l i o c e n e t o Recent s e q u e n c e s have been ment i o n e d by S l a n s k y ( 1 9 6 3 ) .
From s u b s u r f a c e samples t h e f o l l o w -
i n g a s s e m b l a g e s have been found:
Globigerinoides r u b e r d'Or-
b i g n y , Ammonia b e c c a r i i ( d ' o r b i g n y ) , G l o b o r o t a l i a i n f l a t a d ' o r b i g n y , Nonion yranosum d ' o r b i g n y , Nonion commune d ' o r b i g n y , Eponides p r o c e r a B r a d y , and D i s c o p u l v i n u l i n a b e r t h e l o t i d ' O r bigny
.
A f t e r S l a n s k y , (1963) t h i s s e q u e n c e may b e a t t r i b u t e d t o t h e P l i o c e n e u n t i l t h e Recent. I n N i g e r i a (Reyment, 1965) t h e P l i o - P l e i s t o c e n e Benin Form a t i o n i s composed of c l a y and s h a l e s , d e l t a i c t o f u l l y m a r i n e , i n p a r t e s t u a r i n e , lagoonal o r f l u v i o l a c u s t r i n e i n o r i g i n . T h i s f o r m a t i o n y i e l d e d F o r a m i n i f e r a such a s :
&-
Candorbulina
v e r s a J e d l i t s c h k a , G l o b o q u a d r i n a d e h i s c e n s (Chapman, P a r r and C o l l i n s ) , G l o b i g e r i n o i d e s t r i l o b u s ( R e u s s ) , b o l i v i n i d s and Cibicides spp. P o s t Miocene s t r a t a a r e r e p r e s e n t e d a l s o i n Gabon by t h e Akkoso S e r i e s which o v e r l i e c o n c o r d a n t l y l a t e Miocene s e d i m e n t s i n t h e w e s t e r n r e g i o n whereas i n t h e e a s t e r n a r e a they d i s c o r d a n t l y o v e r l i e s t r a t a of P a l e o g e n e a g e . The f a u n a l a s s e m b l a g e i s composed o f Gageonnet, 1963) :
( d e Klasz and
P t y c h o m i l i o l a s e p a r a n s (Brady) , Ammonia
e-
c a r i i ( d ' o r b i g n y ) , Nonion scaphum, G l o b i g e r i n a b u l l o i d e s d ' O r bigny, G l o b i g e r i n o i d e s r u b e r d ' o r b i g n y , G l o b i g e r i n o i d e s saccul i f e r (Brady) , G l o b i g e r i n o i d e s t r i l o b u s (Reuss) , e t c . According t o Bond (1965) t h e P l e i s t o c e n e o f C e n t r a l and South A f r i c a i s r e l a t e d t o p l u v i a l - i n t e r p l u v i a l e v e n t s ; f o s s i l s a r e generally lacking.
The Q u a t e r n a r y s e d i m e n t s a r e f o s s i l
dwnbos, c o l l u v i a l d e p o s i t s and s a n d s o f e o l i a n o r i g i n . A t l a n t i c Ocean During s e v e r a l c r u i s e s Neogene S o u t h A t l a n t i c c o r e s were o b t a i n e d ; from t h e r e c o v e r e d s e c t i o n s v a l u a b l e d a t a c o n c e r n e d
456
t o t h e s e d i m e n t a r y s e q u e n c e s were p u b l i s h e d .
s i s was g i v e n t o t h e P l i o - P l e i s t o c e n e
P a r t i c u l a r empha-
boundary and t o t h e
P l e i s t o c e n e c l i m a t i c c h a n g e s i n f e r r e d by means o f p l a n k t o n i c Foraminifera. Saito,Ewing and Burckle (1966) recorded Pliocene sediments Ridge a t t h e f o l l o w i n g c o o r d i n a t e s :
from m i d - A t l a n t i c 15'06'W;
-
22'59's
06'46'W;
-
26'15'5
03'01'W;
17'39'5-
-
22'06'5
00"
19'W.
From t h e R i o G r a n d e R i s e a n d W a l v i s R i d g e t h e s e a u t h o r s found:
-
25'27's
p e r Miocene; Miocene;
06'28'E
29'02's
21'28'5
-
-
40'05'W
-
Upper M i o c e n e ; 2 4 ' 0 7 ' s
09'13'W
Upper Miocene;
05'45'E
29'52's
-
Up-
36'48'W
Pliocene.
S a i t o , Ewing a n d B u r c k l e
(2. G.) found
t h a t t h e Miocene
i s c h a r a c t e r i z e d by G l o b o q u a d r i n a d e h i s c e n s , a n d t h e P l i o c e n e by G l o b i g e r i n o i d e s
fistulosus.
These a u t h o r s also suggested
t h a t t h e Rio Grande Rise and W a l v i s Ridge a r e r e c e n t f e a t u r e s r e s u l t i n g f r o m u p l i f t i n l a t e M i o c e n e o r Lower P l i o c e n e t i m e s . V a l u a b l e i n f o r m a t i o n was p r o v i d e d b y Blow
(1970a, b ) i n
t h e r e p o r t o f t h e r e s u l t s o f L e g s 3 a n d 4 o f t h e Glomar C h a l lenger Cruise.
Leg 3 c o v e r e d t h e S o u t h A t l a n t i c b e t w e e n D a k a r ,
S e n e g a l a n d R i o d e J a n e i r o , a n d some s i t e s o f Leg 4 l i e i n t h e South A t l a n t i c . The s i t e s o n Leg 3 i n w h i c h N e o g e n e s e d i m e n t s were obt a i n e d a r e s i t e s 1 4 - 1 8 a n d 21-23.
The l o c a t i o n s o f t h e s e s i t e s
and t h e p l a n k t o n i c Neogene Zones i d e n t i f i e d a r e t h e f o l l o w i n g : Hole 1 4 Hole 1 5
-
28" 1 9 . 8 9 s
-
20" 56.46W
E a r l y Miocene
30'
-
1 7 " 58.89W
Pleistocene
53.38s
L a t e Miocene
Middle Miocene E a r l y Miocene
-
30'
30.155
-
1 5 " 42.79W
Pleist.
and Holoc.
Pleistocene
-
28'
02.745
-
6'
36.15W
N19 N18 ,1 9 N15 N6,7 N23 N22
Pliocene
N19-21
L a t e Miocene
N16-18
Middle Miocene Hole 17
N22 N20 ,2 1
Pliocene Early Pliocene
Hole 1 6
N4-N6
Pleist.
-
Holocene
Late Miocene
N15 N22 ,23 N17,18
457
Hole 17 A 28' Hole 18
-
02.74s 58,718
27'
-
6' 36.15W 08' 00.7OW
E a r l y Miocene
N4-7 N22 N20 ,21
Pleistocene Pliocene Early Pliocene
N19 N18 ,19
L a t e Miocene
M i d d l e Miocene
N15 N6,7 N23 N22
E a r l y Miocene Hole 16
-
30' 30.15s
-
15" 42.79W
Pleist.
and Holoc.
Pleistocene Pliocene
N19,21 N16-18
L a t e Miocene
Middle Miocene Hole 1 7
-
28'
02.748
Hole 17A- 28'
02.74s 58.718
-
6'
36.15W
N15 N22-23
P1eist.-Holocene
N17 ,18 N4-7
L a t e Miocene
Hole 18
-
27'
-
6' 36.15W 08'00.70W
-
E a r l y Miocene Pleistocene
N22 N4-6 N19 ,20 N22 N4
E a r l y Miocene Hole 21
-
Hole 22
-
28' 35.10s - 30' 35.85W 30' 00.31 S 35.15.OOW
-
Pliocene Pleistocene Lower Miocene
On Leg 4 t h e s i t e s l y i n g i n t h e S o u t h A t l a n t i c and i n which Miocene t o Holocene s e d i m e n t s were r e c o v e r e d , a r e 23-25. The l o c a t i o n o f t h e s i t e s a n d t h e p l a n k t o n i c Zones i d e n t i f i e d are: Hole 23 - 06'
-
08.75 S
Hole 24
-
06'
16.30s
Hole 25
-
00'
31.00s
-
31" 02.60W P l e i s t o c e n e E a r l y Miocene 30' 53.53W 39' 14.40W
N22 N4 w i t h
Miogypsina E a r l y Miocene
N4 N22
Pleistocene Pliocene L a t e Miocene-Ehrly
Maxwell e t a l .
N19 P l i o . N18
L a t e Miocene N16-18 (1970) r e c o r d e d , from t h e R i o Grande R i s e , s t r a -
t i g r a p h i c s e c t i o n s a t s i t e s 21 and 22 of Leg 3 of t h e Glomar Challenger.
S e q u e n c e s r a n g e i n a g e from t h e Campanian o r o l d e r
t o t h e P l e i s t o c e n e , t h e O l i g o c e n e a n d Miocene b e i n g m i s s i n g a t S i t e 21, and P l i o c e n e d i r e c t l y o v e r l i e s e a r l y Miocene a t S i t e
22. S e v e r a l Neogene p l a n k t o n i c Zones were r e c o g n i z e d by
458
Maxwell, e t a l .
( o p . c i t . ) a t t h e f o l l o w i n g s i t e s of Leg 3
S i t e 15
Zone of G l o b i g e r i n i t a d i s s i m i l i s
Site 16
Zone of G l o b o r o t a l i a a c o s t a e n s i s and
Site 18
Zone of G l o b o r o t a l i a k u g l e r i
G l o b o r o t a l i a merotumida F u n n e l e t a l . ( 1 9 7 1 ) r e c o r d e d s e v e r a l Neogene s e c t i o n s found i n t h e S o u t h A t l a n t i c ; A l b e r i c i , B a r b i e r i , I a c c a r i n o and R o s s i (1973) mentioned Neogene l o c a l i t i e s a t t h e w e s t e r n f l a n k of t h e m i d - A t l a n t i c Ridge a t S i t e s 1 5 and 1 6 , Leg 3 , of t h e Glomar C h a l l e n g e r . Berggren and Amdurer (1973) from s t u d i e s of Leg 3 a t s i t e s 1 4 , 1 7 , and 1 8 , which p e n e t r a t e d Neogene s e d i m e n t s , found c l o s e
r e l a t i o n s h i p s w i t h N e w Zealand and A t l a n t i c F o r a m i n i f e r a d u r i n g t h e O l i g o c e n e and Miocene and t h e c o r r e l a t i o n s t h a t c a n b e made between N e w Zealand and S o u t h A t l a n t i c F o r a m i n i f e r a 1 Zones. B e r g g r e n and Amdurer ( o p . c i t . ) a l s o remarked t h a t b e g i n n i n g w i t h t h e Middle Miocene t h e S o u t h A t l a n t i c f a u n a s a p p e a r t o e x h i b i t a g r e a t e r s i m i l a r i t y w i t h mid-low l a t i t u d e A t l a n t i c f a u n a s t h a n w i t h New Zealand. R e l a t i v e t o t h e A t l a n t i c Ocean t h e B r a z i l i a n c o n t i n e n t a l m a r g i n was i n v e s t i g a t e d i n s e v e r a l a r e a s ; t h e f o r a m i n i f e r a 1 res u l t s w e r e o b t a i n e d by N o g u t i and F o n t a n a d o s S a n t o s ( 1 9 7 2 ) and Noguti ( 1 9 7 5 ) . Miocene s e q u e n c e s were found i n t h e p r o l o n g a t i o n i n t o t h e c o n t i n e n t a l m a r g i n of t h e Amap;, sins.
Par;,
S e r g i p e , and Alagoas ba-
The f o l l o w i n g p l a n k t o n i c zones w e r e r e c o g n i z e d :
e-
b i g e r i n a t r i p a r t i t a , P r a e o r b u l i n a glomerosa, G l o b o r o t a l i a f o h s i peripheroronda, Globorotalia f o h s i peripheroacuta, Globorotalia m a y e r i , and G l o b o r o t a l i a f o h s i r o b u s t a , a l l of e a r l y Miocene age. T e x t F i g . 1 i l l u s t r a t e s t h e l o c a t i o n and t h e a g e of t h e o b t a i n e d c o r e s i n t h e South A t l a n t i c a s w e l l as around t h e s e d i m e n t a r y b a s i n s i n which m a r i n e Miocene and younger s e d i ments were r e c o r d e d . G r e a t emphasis h a s been p l a c e d on t h e P l i o - P l e i s t o c e n e boundary and t h e c l i m a t i c c h a n g e s t h a t o c c u r r e d d u r i n g t h e P l e i s t o c e n e ; r e l a t e d t o t h i s t o p i c s e v e r a l s t u d i e s were undert a k e n i n t h e S o u t h A t l a n t i c , a s w e l l a s i n o t h e r Oceans i n s e a r c h of d a t a r e l a t e d t o t h e t e m p e r a t u r e c h a n g e s d u r i n g t h e
459
Pleistocene.
The d a t a i n d i c a t e t h a t d u r i n g t h e P l e i s t o c e n e
t h e r e were p e r i o d i c o s c i l l a t i o n s of t e m p e r a t u r e w i t h a n a m p l i t u d e o f a b o u t 6 O C w i t h some d i f f e r e n c e s between t h e d i f f e r e n t oceans. E r i c s o n and W o l l i n ( 1 9 5 6 ) s t u d i e d c o r e s i n t h e e q u a t o r i a l m i d - A t l a n t i c Ridge.
Based on p l a n k t o n i c F o r a m i n i f e r a , t h e s e
a u t h o r s found t h a t d u r i n g t h e P l e i s t o c e n e t h e a r e a w a s i n f l u enced by g l a c i a l c l i m a t e , a l t h o u g h t h e c l i m a t e c h a n g e was somewhat l e s s d r a s t i c t h a n i n h i g h e r l a t i t u d e s .
Globorotalia
m e n a r d i i m e n a r d i i , G l o b o r o t a l i a m e n a r d i i t u m i d a and G l o b o r o t a l i a flexuosa -
were c o n s i d e r e d warm w a t e r
s p e c i e s whereas
G&-
b o r o t a l i a p u n c t u l a t a , G l o b o r o t a l i a s c i t u l a and G l o b o r o t a l i a f l a t a a r e c o o l water s p e c i e s and r e l a t e d t o g l a c i a l s t a g e s . Rudiman ( 1 9 7 1 ) , from t h e p i s t o n c o r e s of t h e e q u a t o r i a l
A t l a n t i c , found t h a t i n d i c a t i v e w a r m w a t e r s p e c i e s a r e G l o b i g e r i n i t a a e q u i l a t e r a l i s , Sphaeroidinella dehiscens, Globigerin o i d e s conglobat=,
gona and
=-
G l o b i g e r i n o i d e s t e n e l l u s , Candeina n i t i d a ,
Globigerina rubescens, Globigerina d i g i t a t a , Globigerina Hastigerina pelagica; cold species are:
Globorotalia
h i r s u t a , G l o b o r o t a l i a p u n c t u l a t a , G l o b o r o t a l i a s c i t u l a and bigerina quinqueloba.
g-
Taking i n t o a c c o u n t t h e s e a s s e m b l a g e s
Rudiman c o n c l u d e d t h a t i n t h e e a s t e r n e q u a t o r i a l A t l a n t i c f r a c t u r e z o n e , t h e r e a r e documented c l i m a t i c v a r i a t i o n s o v e r t h e l a s t 1 . 8 m.y. w h i c h c a n b e u s e d t o c o r r e l a t e s e v e r a l c o r e s . Boltovskoy
(1973) s t u d i e d s e v e r a l South A t l a n t i c submarine
c o r e s between L a t 2 2 and 4 7 S and Long. 2 2 and 6 7 W . a r e a Boltovskoy r e c o r d e d t h e P l i o c e n e / P l e i s t o c e n e
In this
boundary w i t h
t h e e v o l u t i o n a r y z o n a l m a r k e r s G l o b o r o t a l i a t o s a e n s i s and
G-
borotalia truncatulinoides. On t h e o t h e r h a n d , B o l t o v s k o y
( o p . c i t . ) b a s e d on:
1) t h e
t o t a l r e l a t i o n s h i p s of w a r m and c o o l s p e c i e s ; 2 ) t h e r e l a t i o n G l o b o r o t a l i a m e n a r d i i / G l o b o r o t a l i a i n f l a t a and 3 ) t h e c o i l i n g d i r e c t i o n of G l o b i g e r i n a pachyderma and G l o b i g e r i n a b u l l o i d e s found i n t h e P l e i s t o c e n e two c o o l and two w a r m p e r i o d s which were r e l a t e d t o t h e E a r l y Wurm-Principal Wurm and Riss-Wurm/ i n t e r s t a d i a l Wurm r e s p e c t i v e l y . Summarv S o u t h American A t l a n t i c Miocene p l a n k t o n i c a s s e m b l a g e s a r e w e l l r e p r e s e n t e d i n t h e P e l o t a s B a s i n ( s o u t h e r n B r a z i l ) and i n
460
Text F i g .
1.
L o c a t i o n o f t h e c o r e s r e c o v e r e d from t h e S o u t h A t l a n t i c and c o r r e s p o n d i n g a g e s , and t h e s e d i m e n t a r y b a s i n s i n which Miocene and P l e i s t o c e n e F o r a m i n i f e r a were r e c o r d e d .
t h e c o n t i n e n t a l m a r g i n of t h e Amazonas and S e r g i p e / A l a g o a s basins. The y o u n g e s t Miocene m a r i n e s e q u e n c e s a r e r e c o r d e d i n t h e P e l o t a s B a s i n i n which m i d d l e - l a t e Miocene p l a n k t o n i c Foraminif e r a a r e r e c o r d e d by F a l c e t t a ( 1 9 7 6 , M S )
.
P l a n k t o n i c a s s e m b l a g e s from low l a t i t u d e s a r e comparable t o t h o s e of t h e C a r i b b e a n a r e a , w h e r e a s t h o s e of t h e P e l o t a s B a s i n and o t h e r b a s i n s from h i g h e r l a t i t u d e s show more a f f i n i t i e s w i t h t e m p e r a t e assemblages. A t 30'
bigerinids.
S g l o b o r o t a l i i d s a r e s c a r c e , b e i n g dominant g l o -
The z o n a l scheme p r e s e n t e d by Blow ( 1 9 6 9 ) i s a
u s e f u l one f o r l a t i t u d e s n e a r 30's
o r h i g h e r , whereas a t lower
l a t i t u d e s t h a t of B o l l 1 ( 1 9 6 6 ) seems t o b e more r e l i a b l e . Most of t h e u v i g e r i n i d s and b o l i v i n i t i d s p r e s e n t i n t h e P e l o t a s B a s i n a r e r e s t r i c t e d t o t h e Miocene; a l t h o u g h some a r e of A u s t r a l o a s i a t i c o r i g i n , most of t h e s p e c i e s a r e r e l a t e d o r
common t o European o n e s .
461
Other benthonic Miocene assemblages show close relationships to those faunas living today in the western Atlantic Ocean, and many cosmopolitan species are common components of the associations. African Miocene benthonic foraminifera1 associations, a1 though they contain some cosmopolitan and Caribbean species, show strong endemic features: the planktonics are those known from the Caribbean and equatorial areas. No Australoasiatic influences are perceptible. Large Foraminifera, although scarce are recorded from the South American continent only in northern Brazil, that is, the equatorial region. Below this latitude no large Foraminifera were recorded. In the African continent, large Foraminifera were recorded from Gabon and Cameroon. South American and African benthonic foraminifera1 assemblages show clear latitudinal zonation. In South America this is documented by the abundance of warm water species down to latitudes of 20-25OS whereas south of this latitude the faunas assemblage lack typical warm water species or they become scarce. Pleistocene South American assemblages are similar to those of the present Atlantic Ocean. In African basins, where post-Miocene marine sediments were recorded, the strong endemic features, typical of older faunal assemblages seem to have been replaced by more cosmopolitan faunal elements. References Alberici, A., Barbieri, F., Iaccarino, S. and Rossi., U. 1973. Considerazioni biostratigrafiche e paleoecologiche sul Neogene del Fianco Occidentale della dorsale Medio-Atlantica Meridionale (Foraminifer1 dei "Sites" 15 e 16 del "Leg" I11 D.S.D.P. "L'Ateneo Parmense" - Acta Naturalia, Vol. IX Fasc. 2, p. 137-151. Antunez, M. T. , 1964. 0 Neocretdceo e o Cenozoic0 do litoral de Angola. Junta de Investigacoes de Ultramar, Lisboa. Belmonte, Y. C., 1966. Stratigraphie du bassin sgdimentaire du Cameroun. Proc. 2nd W. African Micropal. Coll. Ibadan, 1965. Berggren, W. A. and Amdurer, M., 1973. Late Paleogene (Oligocene) and Neogene planktonic foraminiferal biostratigraphy of the Atlantic Ocean (Lat. 30°N to Lat. 30's) Riv. Ital. Paleont. v. 79, no. 3,,p. 337-392. Bertels, A,, 1976b (in press). Micropaleontology and Paleogeography of the Upper Cretaceous and Cenozoic of Argentina.
462
Bertels, A . , 1976. Paleogene Foraminifera - South A t l a n t i c , t h i s volume. B e r t e l s , A . , F a l c e t t a , M. M . , and K o t z i a n , S . B . , 1 9 7 6 . (in p r e s s ) . Micropaleontologia (Foraminiferos y OstrAcodos) d e l Cuaternario d e Palmares do S u l (Formaci6n C h u i ) , B r a s i l . Blow, W . H . , 1 9 6 9 . L a t e Middle Eocene t o R e c e n t p l a n k t o n i c f o r a m i n i f e r a 1 b i o s t r a t i g r a p h y . I n t e r n , C o n f e r e n c e on P l a n k tonic Microfossils. p. 199-422. , 1 9 7 0 . A Deep S e a D r i l l i n g P r o j e c t . Leg 3 . F o r a m i n i f e r a from s e l e c t e d s a m p l e s . I n Maxwell, A . E . , e t a l . , 1 9 7 0 . I n i t i a l k e p o r t s of t h e Deep S e a D r i l l i n g P r o j e c c 111. W a s h i n g t o n (U.S. Government P r i n t i n g O f f i c e ) p . 629-661. , 1 9 7 0 b . Deep S e a D r i l l i n g P r o j e c t , Leg 4 , F o r a m i n i f e r a from s e l e c t e d s a m p l e s . I n Maxwell, A . E . , e t a l . , 1970. I n i t i a l R e p o r t s o f t h e Deep S e a D r i l l i n g P r s e z , v . I V . W a s h i n g t o n (U.S. Government P r i n t i n g O f f i c e ) p . 383-400. B o l t o v s k o y , E . , 1 9 5 9 . Los f o r a m i n i f e r o s d e 10s s e d i m e n t o s c u a t e r n a r i o s e n 10s a l r e d e d o r e s d e P u e r t o Q u e q u e ( P r o v i n c i a d e Buenos A i r e s ) , Rev. Asoc. G e o l . A r g e n t i n a . T . X I V , n o s . 3 - 4 , p . 251-277. , 1973. E s t u d i o d e t e s t i g o s submarinos d e l A t l A n t i c o s u d o c c i d e n t a l . Rev. Museo Arg. C i e n c i a s N a t . " B e r n a r d i n o R i v a d a v i a " G e o l . T . V I I , n o . 4 , p . 213-340. Bond, G . , 1 9 6 5 . Q u a n t i t a t i b e a p p r o a c h e s t o r a i n f a l l a n d t e m p e r a t u r e changes i n t h e Quaternary of s o u t h e r n A f r i c a . Int e r n a t i o n a l S t u d i e s on t h e Q u a t e r n a r y . V I I I n t e r n . Congress f o r Q u a t e r n a r y R e s . Ed. W r i g h t , H . E . J r . , a n d F r e y , D . G . , p . 323-336. Tectonique e t sedimentation B r o g n o n , G . and V e r r i e r , G . , 1 9 6 6 . d a n s le b a s s i n du Cuanza ( A n g o l a ) . I n : B a s s i n s s 6 d i m e n t a i r e s du l i t t o r a l a f r i c a i n . I h e p a r t i e : L i t t o r a l A t l a n t i q u e . A s s o c , d e s S e r v i c e s G g o l . A f r i c a i n s . , p . 207-252. C l o s s , D . , 1 9 6 7 . Miocene p l a n k t o n i c F o r a m i n i f e r a f r o m s o u t h e r n B r a z i l . M i c r o p a l e o n t o l o g y l v . 1 3 , n o . 3 , p . 337-344. , 1 9 7 0 . E s t r a t i g r a f i a d a B a c i a d e P e l o t a s , R i o Grande d o S u l . I h e r i n g i a , G e o l . , n o . 3 , p . 3-76. Cenozoic F o r a m i n i f e r a from C l o s s , D . , and Madeira M . L . , 1968. t h e Chuy D r i l l Hole, n o r t h e r n Uruguay. A m e g h i n i a n a , T . V . , n o . 7 , p . 229-236. B i o s t r a t i g r a p h i e du d e K l a s z , I . , and Gageonnet, R . , 1963. B a s s i n Gabonais. C o l l o q u e I n t e r n a t . d e M i c r o p a l . Dakar. p . 277-304. D. 1964a. Deux impord e Klasz, I . , Le Calvez,Y.,and R & a t , t a n t e s e s p ' e c e s d e F o r a m i n i f h r e s du Miockne i n f e ' r i e u r d e 1 ' A f r i q u e o c c i d e n t a l e . C . R . Sommaire d e s S Q a n c . S O C . G h o l . F r a n c e , F a s c . 5 , p . 194-195. , 1 9 6 4 b . Deux nouveaux B u l i m i n i d a e ( F o r a m i n i f L r e s ) , du P a l 4 o g e n e d e 1 ' A f r i q u e o c c i d e n t a l e . C . R . Sommaire d e s S e a n c . S O C . G e o l . F r a n c e F a s c . 5 , p . 208-209. , 1 9 6 4 c . Deux nouveaux g e n r e s d e F o r a m i n i f L r e s d ? Gabon ( A f r i q u e E q u a t o r i a l e ) . C . R . Sommaire d e s S & a n c . S O C . G e o l . F r a n c e . F a s c . 6 , p . 236-237. d e Klasz, I . , and Micholet, J . , 1 9 7 0 . E l g m e n t s nouveaux c o n c e r n a n t l a B i o s t r a t i g r a p h i e du B a s s i n G a b o n a i s . I V C o l l o q u e A f r i c a n M i c r o p a l . l p . 109-141. d e K l a s z , I . , and R e r a t . , 1 9 6 2 . Une n o u v e l l a esp'ece d ' E p o n i d o p s i s ( F o r a m i n i f e r a ) d e 1 ' A f r i q u e o c c i d e n t a l e . C . R . Sommaire S e a n c . SOC. G e ' o l . F r a n c e . F a s c . 4 , p . 112-113.
,
463
D e l a n e y , P . J . V . , 1967. Geomorphology a n d Q u a t e r n a r y c o a s t a l g e o l o g y of Uruguay, p . 1-39. D r o o g e r , C . W . , 1966. N o t e s o n Mio s i n a of Cameroon. P r o c e e d . of t h e Second W e s t A f r i c a n M i c b . , Ibadan, p. E r i c s o n , D . B . , and W o l l i n , G . , 1956. C o r r e l a t i o n of s i x c o r e s from t h e e q u a t o r i a l A t l a n t i c and t h e C a r i b b e a n , Deep-sea R e s e a r c h , v . 3 , no. 2 , p . 104-125. F a l c e t t a , M. M . , 1976 (MS). F o r a m i n i f e r o s P l a n c t 6 n i c o s e Est r a t i g r a f l a d o Cenozoico d a Bacia d e P e l o t a s , B r a s i l . F a u l k n e r , L . S . , d e K l a s z , I . , and R b r a t , D . , 1963. Megastom e l l a nov. g e n . nouveau F o r a m i n i f e r e d e l ' A f r i q u e o c c i d e n t a l e Rev. M i c r o p a l . , v . 6 , no. 1, p . 19-22. F e r n a n d e s , J . M. G . , 1975 (MS) 0 Ghnero U v i g e r i n a ( F o r a m i n i f e r i d a ) d o C e n o z o i c o S u p e r i o r na Bacia d e P e l o t a s , Rio Grande do $ 1 1 1 ,. A r a s i l . D i s s e r t a c a o d e Mestrado. F e r r e i r a , C . S . , G o n z a l e z , B . B . , and R o d r i g u e z , F . B . H . , 1973. O c c o r r e n c i a d a Formacao P i r a b a s (Mioceno i n f . ) na Bacia d e B a r r e i r i n h a s , Maranhao. Rev. B r a s i l e i r a d e Geoc., v . 3 . F i d a l g o , F . , d e F r a n c e s c o , F. O . , and P a s c u a l , R . , 1975. Geol o g i a S u p e r f i c i a l d e l a L l a n u r a B o n a e r e n s e . V I Congreso Geol. A r s . R e l a t o r l o G e o l . P r o v . Buenos A i r e s , p . 103-138. F u n n e l l , B . M . , 1 9 7 1 . The o c c u r r e n c e o f p r e - Q u a t e r n a r y microf o s s i l s i n t h e o c e a n s . ( I n ) F u n n e l , B . M. and R i e d e l W . , ( e d s . ) " M i c r o p a l e o n t o l o g y of O c e a n s , " Cambridge, Cambridge Univ. P r e s s , 1967. p . 507-534. Graham, J . J . , d e K l a s ? , I . , and R h r a t , D . , 1965. Q u e l q u e s i m p o r t a n t s F o r a m i n i f e r e s du T e r t i a i r e du Gabon ( A f r i q u e equat o r i a l e ) . Rev. micro pal.,^. 8 , no. 2 , p. 71-84. L e C a l v e z , Y,., d e K l a s z , I . and Brun, L . , 1974. N o u v e l l e cont r i b u t i o n a l a c o n n a i s s a n c e d e s m i c r o f a u n e s du Gabon. Rev. E s p a g o l a d e M i c r o p a l . , v . V I , no. 3 , p. 381-400. Malumian, N . , 1968. F o r a m i n i f e r o s d e l C r e t a c i c o S u p e r i o r y T e r c i a r i o d e l subsuelo de l a Provincia Santa Cruz, Argentina. Ameghiniana, T . V . , no. 6 , p.,191-227. , 1970a. B i o s t r a t i g r a f i a d e l T e r c i a r i o m a r i n o d e l subs u e l o d e l a P r o v i n c i a d e Buenos A i r e s ( A r g e n t i n a ) Ameghinian a , T . V I I , no. 2 , p.,173-203. , 1972. F o r a m i n i f e r o s d e l O l i g o c e n o y Mioceno d e l subs u e l o d e l a P r o v i n c i a d e Buenos A i r e s . Ameghiniana, T . I X , no. 2 , p. 97-137. Malumian, N . , Masiuk, V . , and R i g g i , J . C . , +971. M i c r o p a l e o n t o l o g i a y sedimentologia d e l a p e r f o r a c i o n SC-1 Provincia S a n t a C r u z , RepAblica A r g e n t i n a . Su i m p o r t a n c i a y c o r r e l a c i o n e s . Rev, Asoc. G e o l . A r g e n t i n a . T . X X V I , no. 2 , p . 175208. Maxwell, A, E . e t a l . , 1 9 7 0 . I n i t i a l R e p o r t s of t h e Deep Sea D r i l l i n g P r o j e c t , Volume 111. Washington (U.S. Government P r i n t i n g O f f i c e ) , xx + 806 PP. N o g u t i , I . , 1975. Zonaci6n B i o e s t r a t i g r i f i c a d e 10s F o r a m i n i f e r o s p l a n c t o n i c o s d e l T e r c i a r i o d e G r a s i l . Rev. Espa"n1a d e M i c r o p a l . v . V I I , no. 3 , p . 391-401. N o g u t i , I . , a n d F o n t a n a d o s S a n t o s , J . , 1972. Zoneamiento p r e l i m i n a r p o r f o r a m i n f f e r o s p l a n c t o n i c o s d o A p t i a n o ao Mioceno na
464
p l a t a f o r m a c o n t i n e n t a l do B r a s i l . Bol. T e c . P e t r o b r a s . Rio d e J a n e i r o . v . 1 3 , n o . 3 , p . 265-283. P e t r i , S . , 1 9 5 4 . F o r a m i n i f e r o s f o s s e i s d a Bacia d o M a r a j 6 . Univ. Sao P a u l o . F a c u l t a d e d e F i l o s o f i a , C i e n c i a s e L e t r a s . B o l . n o . 1 7 6 , G e o l . n?. 11, p . 17173. 1 9 5 7 . F o r a m i n i f e r o s m i o c e n i c o s d a Formacao P i r a b a s . Univ. Sao P a u l o , F a c u l t a d e d e F i l o s o f i a , C i e n c i a s e L e t r a s . B o l . n o . 216, G e o l . ny. 1 6 , p . 1 - 7 9 . , 1972. F o r a m i n i f e r o s e o a m b i e n t e d e d e p o s i c a o d o s s e d i m e n t o s d o Mioceno d o Reconcavo B a i a n o . Rev. B r a s . d e Geoc i h n c i a s . v . 2 , n o . 1, p . 51-67. P i s e t t a , J . L . , 1968 (MS). D e s c r i p c i 6 n d e una f a h u l a d e F o r aminiferos de l a Provincia de Entre R i o s . Trabajo Final de L i c e n c i a t u r a , F a c u l t a d d e C i e n c i a s E x a c t a s y N a t u r a l e s , Univ. Buenos A i r e s ( u n p u b l i s h e d ) . Rocha, A . T . , and F e r r e i r a , J . Y., 1 9 5 7 . C o n t r i b u c a o p a r a 0 e s t u d o d o s F o r a m i n i f e r o s d o T e r c i a r i o d e Luanda. Garcia d e O r t a , v . 5 , n o . 2 , p . 297-310. Reyment, R . A . , 1 9 6 5 . A s p e c t s o f t h e Geology o f N i g e r i a . The s t r a t i g r a p h y o f t h e C r e t a c e o u s and C e n o z o i c d e p o s i t s . Ibadan Univ. P r e s s . p . 1-145. Rudiman, W . F . , 1 9 7 1 . P l e i s t o c e n e S e d i m e n t a t i o n i n t h e e q u a t o r i a l A t l a n t i c : S t r a t i g r a p h y and F a u n a l P a l e o c l i m a t o l o g y . G e o l . S O C . Am. B u l l . v . 8 2 , n o . 2 , p . 283-301. S a i t o , T . , Ewing, M . , and B u r c k l e , L . H . , 1 9 6 6 . T e r t i a r y Sediment from t h e m i d - A t l a n t i c Ridge. S c i e n c e , v . 1 5 1 , no. 3714; p . 1075-1079. S c h a l l e r , H . , V a s c o n c e l o s , D . N . , and C a s t r o , J . C . , 1 9 7 1 . E s t r a t i g r a f i a p r e l i m i n a r d a B a c i a s e d i m e n t a r d a Foz d o R i o Amazonas. SOC. B r a s i l e i r a d e G e o l . A n a i s XXV C o n g r e s o , v . 3 , p . 189-202. S l a ? s k y , M . , 1 9 6 3 . C o n t r i b u t i o n 'a 1 ' 6 t u d e g & o b o g i q u e d u b a s s i n s e d i m e n t a i r e & t i e r du Dahomey e t du Togo. Memoires B u r e a u d e R e c h e r c h e s G 6 o l o g i q u e s e t Min. no. 11, p . 1 - 2 7 0 . S u a r e z S o r u c o , J . R . , 1968 (MS) Estudio micropaleontol6gico d e l c o r d 6 n l i t o r a l d e l a l o c a l i d a d d e Mar C h i q u i t a , P r o v i n c i a d e Buenos A i r e s . T r a b a j o F i n a l d e L i c e n c i a t u r a , F a c u l t a d d e C i e n c i a s E x a c t a s y N a t u r a l e s , U n i v e r s i d a d d e Buenos A i r e s (unpublished). T h i e s e n , z . V . , 1 9 7 5 (MS). B o l i v i n i t i d a e e C a u c a s i n i d a e ( F o r a m i n i f e r i d a ) do C e n o z o i c o S u p e r i o r d a Bacia d e P e l o t a s , R i o Grande d o s u l , B r a s i l . D i s s e r t a c a o d e Mestrado. U n i v e r s i d a d e F e d e r a l d o R i o Grande d o S u l , B r a s i l . ( u n p u b l i s h e d ) T i n o c o , I . d e M . , 1958. F o r a m i n i f e r o s Q u a t e r n a r i o s d e O l i n d a , E s t a d o d e Pernambuco. Dep. Nac. P r o d . Min. G e o l . e Min. Monog r a f i a X I V , pp. 1 - 6 1 .
,
.
465 Plate I
Fig.
2
Fig.
3
-
Fig. 4
-
Fig.
5
-
Fig.
6
Fig.
7
-
Fig.
8
-
Fig. 1
G l o b o r o t a l i a conomiozea J e n k i n s , x 2 7 8 , P e l o t a s B a s i n (Middle-Upper M i o c e n e ) G l o b o r o t a l i a c r a s s u l a v i o l a Blow, x 1 7 3 , P e l o t a s B a s i n (Middle-Upper Miocene) G l o b i g e r i n a n e h e n t e s Todd, x 1 4 9 , P e l o t a s B a s i h - U p p e r Miocene) G l o b o q u a d r i n a d e h i s c e n s (Chapman,Parr a n d C o l l i n s j l x 1 8 6 , P e l o t a s B a s i n (Middle-Upper M i o c e n e ) G l o b o r o t a l i a humerosa ( T a k a y a n a g i and S a i t o ) , ~ 1 8 6 P e l o t a s B a s i n (Middle-Upper Miocene) Globorotalia 3mpha Jenkins, xi86 P e l o t a s B a s i n (Middle-Upper Miocene) G l o b i g e r i n a d e c o r a e r t a T a k a y a n a g i and S a i t o , x186 Pelotas B a s i f p e r Miocene) G l o b i g e r i n a d r u r y i A k e r s , x186 P e l o t a s B a s i n (Middle-Upper Miocene)
466
PLATE I
46 7
CENOZOIC OSTRACODA M.C.Keen,
- NORTH ATLANTIC
University o f Glasgow, Scotland
A B S TRACT The ostrocod fauna o f each Cenozoic epoch i s described where possible from western Europe (Denmark, Belgium, England, France, N.Spain), the A t l a n t i c seaboard o f N .America, Greenland, Rockall, and from the N . A t l a n t i c Basin. No species are known i n common to the two sides o f the N . A t l a n t i c for the Palaeogene, although genera and species groups i n d i c a t e some communication. Most Palaeogene ostracods c a n be related to Cretaceous forebears. The Neogene saw a renewol o f the ostracod fauna, w i t h w i d e and r a p i d dispersal o f genera, Fourteen ostracod w i t h a few species present o n both sides o f the A t l a n t i c . zones are proposed for the Palaeogene o f N.W.Europe, but these cannot be recognised a t present i n A q u i t a i n e due to p r o v i n c i a l differences.
R ~ S U M ~ O n a d g c r i t la faune d'ostracodes de tous les Qpoques cknozoiques, s i possible, de Danemark, Belgique, Angleterre, France, e t Espagne; de lo rnarge orientale de I'Amgrique d u Nord; de Rockall, e t du bassin de I ' A t l a n t i q u e N o r d . A u cours d u Paliog'ene o n ne trouve aucun des gsp'eces en tous les deux c6tes de I ' A t l a n tique Nord, b i e n que l a pr&ence du mgme genres suggcerent une liaison foible. Le Ngog'ene v o i t I ' a p p a r i t i o n de plusieurs formes nouvelles avec u n net renouvellement de la faune d'ostracodes. O n trouve un c e r t a i n nombre d'gsp'eces comO n a reconnu wesuccession de 14 munes aux deux cstes de I ' A t l a n t i q u e . biozones dons le PaIQog'ene de I'Europe nord-ouest.
Text fig.1.
Palaeogene areas discussed
Ip
m m
0 s TR AC 0D ZONES FOR
STAGE
N. W. EUROPE
PARIS
BRACKISH FRESHWATER OSTRACOD ZONES
ENGLAND
BAS,N
Noocyprideis Virgatocypris / / / /
/
-AT TORFIAN
THANETIAN
Fig.2.
Zonation of the Palaeogene, partly bosed on the work of Berggren and Curry.
BELGIUM
469
FAUNAL SUMMARY
A vast amount of work has been carried out on the Palaeogene ostracods of western Europe and North America, but as it varies from area to area it is impossible to treat all regions in similar detail, In %rope most workers have been concerned with the faunas of Belgium, the Paris Basin, southern England, Germany, and Aquitaine.
The
present state of knowledge makes it possible to attempt a zonation of the Palaeogene of these areas. In North America most work has been concentrated on the faunas of the Gulf states with relatively few studies along the Atlantic seaboard, so a zonal scheme cannot be justified at present.
On the other hand, Neogene faunas are
well known in the area from Florida to Maryland so that ostracods can be used for very accurate correlation.
The only Neogene faunas
described from the Atlantic coasts of Furope are from the classic localities of Aquitaine. ?A LJAw C'ENE
Marlibre (1958) and Deroo (1965) recognised three faunizones in the Upper Danian and Montian of Belgium and Holland, refcrred to as the "Couches
a Cytherelloidea" (upper P1) , "Couches k Cytheretta"
an2 "Couches h Triginglymus" ( ? > = Montian).
Mme Damotte (1964)
was able to recognise the lower two in the Faris Basin, so they probably have a regional significance. zones in the zonal scheme given below.
They form the three lowest The 'eelandian fauna of
;>enmarkis not very well known, although ostracods have been listed by leroo and some described by Triebel.
Particular importance is
placed on the presence of ?aracgtheretta reticosa Triebel, which is also present in the Thanet Beds of Kent and the Palaeocene of noland (5ecezechura, 1965).
The ostracods of the i:zbles de
Brachcux of the Paris Basin 3re the only Upper Palseocene fauna to it is difficult to know how many of its species are truly indigenous. Of 26 doscribed be described in detail from western %rope,
so
species (!.postolescu, 1956) only three are present in the Thmet Beds. in particular, the Hszelina of the Thanct Beds (undescribed species) is not the same as that of the Crbles de Brachcux, "?uriana" sculpta.ADO:,T3LXSCU, while the absence of P.reticosa from the lables de Br?.chcux is considered important, although bearing in mind that .this m?.y be due to geographical factors, i.e., it may be
a
northern "cold watert'species.
However, the evidence
470
sugg5st.; that thi Thanetian of the Taris Basin is rath-r younger thsn the typa ;hanet Beds. shoxz in ?ext Pig. 3.
7h.i distribution of lmportant specles are
Moraeleberir conaliculata (APOST.) Opimocythere incis0 (APOST.) Clithrocytheridea mogna APOST. Cytheroptemn thilienrir APOST. Cytheretto new0 montenrir MARLIERE m o l l f a n . Cytheretta newa montenrir MARLIERE large f o n . Cytheretto news newa APOST. Parocytherette reticora TRIEBEL
.
Limburgino longiparoceo (DEROO) Liniburgincl arnobide(la firwrata DEROO bmantino stelloto DEROO Paleomonmirabilia geulenrir DEROO Limburgi no bi lame Ilom bi lame Ilosa (MAR LI ERE) Krithe montensir DEROO Boirdio montensir MARLIERE Cytherellcl montenrir MARLIERE Cythere I lo logenalir MAR LlERE
Fig.3.
Distribution o f ostracods in the Palaeocene of Europe.
The marine Thanetian is succeeded by continental deposits with a Vetustocytheridea lignitarum DOLLE1US (=V.guitrancourtensis AP0‘:T. ) , found throughout northern France, southern England, and possibly in northern hquitaine. Other important species are Neocgprideis durocortoriensis APOST. and Cgtheromorpha aillyensis BIGNOT. Palaeocene faunas have been described from the north and south of the Pyrenees (Tambareau 1972, Ducasse 1972, Tambareau et Villatte 1974). The “Lower Thanetian” has a poorly preserved marine
471 o s t r a c o d fauna.
The "Upper Thanetian" h a s a b r a c k i s h water fauna
i n i t s lowest beds, with Vetustocytheridea sp.,
Cytheromorpha a f f .
a i l l y e n s i s , and Neoc.yprideis m a n d i n a t u s TAIBARIUU.
The l a t t e r i s
o f great importance b e c a u s e i t i s i n t e r m e d i a r y between t h e M a a s t r i c h t i a n N.murciensis DAXOTT", F F FQURCADE and N . d u r o c o r t o r i e n s i s . T h i s is t a k e n t o r e p r e s e n t zone 1 o f I h e Ducasse ( 1 9 7 2 ) d e s c r i b e d from Campo i n n o r t h e r n Spain (see Text F i g . 2 ) .
The upper p a r t o f
t h e T h a n e t i a n of t h e P y r e n e e s c o n t a i n s a n u n d e s c r i b e d marine f a u n a , w i t h two s p e c i e s r e l a t e d t o T o l i s h P a l a e o c e n e o s t r a c o d s : cf
Kingmaina
opima. 5ZCEZECHUXA and T r a c h y l e b e r i d e a a f f . semiplana SCZZECHURA.
; h e Yucasse ( 1 9 7 2 ) h a s a l s o d e s c r i b e d I l e r d i a n f a u n a s , d i s c u s s e d below under t h e Eocsne.
!m :e
Ezcezechura (1971) h a s d e s c r i b e d a l a r g e Lower Palaeocene
fauna from V e s t e r n Greenland. many unassigned.
Thirty nine species a r e i l l u s t r a t e d ,
The fauna cannot be r e l a t e d t o any i n North
America o r Thrope. O s t r a c o d s from t h e Canian B r i g h t s e a t Formation o f IJaryland have been d e s c r i b e d by Hazel (1?68), where r e f e r e n c e t o o t h e r works may be seen.
The f a u n a , c o n s i s t i n g of t h i r t y f i v e s p e c i e s , i s v e r y
d i s t i n c t i v e , h a v i n g l i t t l e in common w i t h t h e o v e r l y i n g A p i a Formation, b u t w i t h t h i r t e e n s p e c i e s p r e s e n t i n v a r i o u s Lower '3alaeocene Gulf Coast l o c a l i t i e s .
The o s t r a c o d s i n d i c a t e
c o r r e l a t i o n wi-th t h e upper p a r t s o f t h e K i n c a i d Formation o f Texas and ?.he Clayton Formation o f 8lr,bama.
The f a u n a i s dominated
V e s t u s t o c y t h e r i d e a and Opimocythere, i n d i c a t i n g s h a l l o w marine i n n e r e u b l i t t o r a i conditions.
The p r e s e n c e o f V e t u s t o c y t h e r i d e a on
both s i d e s o f th.c A t l a n t i c i s i n t e r e s t i n g ; thc, i a r l i e s t s p e c i e s , V . f o r n i c a t a
Hazel p o i n t s o u t t h a t
f o r n i c a t a (.4L
similar t o H a p l o c y t h e r i d e a , and s u g g e s t s i t evolved from t h e l a t t e r . H a p l o c y t h e r i d c a i s a v e r y common g m u s i n t h e C r e t a c e o u s and T e r t i a r y o f ii.hmerica, t h a t Vetustocgthsridea,
b u t is r a r e i n m o p e . a'
So i t i s p r o b a b l e
s h a l l o w marine and b r a c k i s h w a t e r o s t r a c o d
somshow m i g r a t e d a c r o s s t h ? 4 t l a n t i c t o LGrops, p o s s i b l y v i a GrePnland where t h e ocean was narrow p r i o r t o ocean f l o o r s p r m d i n g ? The :?quia Formation i s p r o b a b l y o f idontisn a g e i n p a r t , as v h l l as i n c l u d i n g younger members, and c o n t a i n s s l a r g e and 3 , i s t i n c t i v e fauna.
3ne s p e c i e s o f i n t e r e s t i s t h a t r e f e r r e d t o
by Hazel as : < o c y t h e r o p t ? r o n a f f . t h i l i e n s i s A?3ET.,
originally
d e s c r i b e d from t h e S a b l e s d e Bracheux o f t h e ;Jaris Basin.
Pooser
412
(1965) has described the fauna of the Black ;dingo Formation of South Carolina which suggest that the formation straddles the Palaeocene-Xocene boundary, most of it being Eocene. He recognised tuo assemblage zones, Brachycythere interrasilis A L X X A N D B and Cytherelloidea nanafaliensis HOI:.'F, which could represent differences in age, environment, o r both. The distribution of some common Palaaocene and Eocene species from the Atlantic coast are shown in Text Fig.4.
I
Y
i U. EOCENE
I 1
M. EOCENE
L!
EOCENE
ACUIA PALAEOCENE BRIGHTSEAT
7I T I +
Fig.4. Distribution o f ostracods in the Palaeogene of eastern U.S.A.
Eocene faunas have been described by Apostolescu (1955, 1956), Haskins (1968-72) and Keij (1957). The base of the Ypresian stage is taken to define the base of the Xocene, which lies within zone P6. This stage is represented by the London Clay and Argiles d'Ypres. Three subeones can be recognised (Text Fig.2) in England and Belgium, although it is not clear to what extent these are facies controlled. The fauna is very characteristic, with a deeper water element represented by Brachycythere triangularis REUSS, Tchinocythereis reticulatissum
473 E 4 G A R l Krithc londinensis JONE2, and Trachyleberidea prestwichiana (JONES AiU’3 BHZRBORN)
(JONEE)
,
, shallower water
by Cytheretta scrobiculoplicata Hazelina aranca (JONES A m CRERBORN) , Pterygocytheris
laminosa HASKIN’?, Zchuleridea perforata insignis ( J O N E ) and Trachyleberis spiniferimma ( cJOIJE‘J Aim SHFBBORN) , and coastal waters by Zytheridea primitia H.44CKIBS and Cytheridea newburyensis GOKCEX. The faunas have been described by Xagar ( 1 9 6 5 ) , Gokcen ( 1 3 7 1 ) , Haskins (1968-72), and Hillems (1975 ). Tho Cuisian substage of the Paris Basin contains a rich fauna (.4postolsscu, 1956) including Novoc.ypris whitecliffensis (HAZKIWS). This species first appears in the Zables de Mons-en-Frevele of Belgium with a fauna more characteristic of the London Clay, but lacking Cytheretta scrobiculoplicata.
Novocypris is a genus more
typical of the Aquitaine Basin. In Spain and Lquitaine the Ilerdian spans the PalaeoceneEocene boundary. Xme 3ucasse recognised three ostracod biozones from Camp0 in northern ?pain for the Ilerdian, numbered 2, 3, and 4 on Text Fig.2. The lowest zone is characterised by a littoral marine fauna with genera such as Bairdoppilata, Pokornyella, and Guadracythere. The middle zone is characterised by a deep shelf or continental slope fauna, with Cytherella consueta DELTEL, Echinocythereis, Krithe, and Pontocyprella aturica DELTXL. The two species of Idlle Peltel are found in many Tertiary deep water The upper zone marks a return to shallower deposits of Iquitaine. water.
Throughout the ?yrenean area the base of the Ilerdian is
marked by the appearance of Pokornyella citrea TAMBIBAREAU often accompanied by Rchinocytheris isabenana DFXTLI (Tambareau, 1972). Biozone 5 of Ducasse (1972) is equated with the Cuisian and the first appearance of Novocypris eocaenica l’lUCAS5Ein northern Spain, although this species occurred earlier in northern Aquitaine. The Xiddle Eocene commences with the Lutetian stage. The oldest ostracod fauna recognised so far is found in Fisher Jed 1 of Bracklesham ( = lowest Fisher Bed VI of Khitecliff Bay; the numerical subdivisions on Text Pig.2 are the Fisher Beds of Khitecliff Bay), but the fauna is undescribed. -4bove this, the ostracod zone Ob contains the typical Lutetian fauna seen in the This subzone sees the first Paris Basin (Apostolescu, 1355). appearance of Cytherelloidea damariacensis APOSTOLESCU,
474
'tergpocythcrzis 'i'i?Z:;BEL,
cornuta ( R O E X E R ) , Schixocythere appendiculata
Schuleridea perforata perforata ( R O ? X 8 ) and other species
indicated on Text 7ig.5. A major evolutionary radiation CM be recognised amongst the ostracods at this time (Keen 197Zc). The upper part of the Lutetian, correlated with the Biarritzian, is marked by the appearance of Cythoridea ripida rigida HASKINS (Text
This species, together with Cgtheretta forticosta K D J T and Fig. 5). Xchinocgthereis scabropapulosa JONES, differentiate ostracod Zone 9 from 8b. All three are present in Zone 10, but accompanied by descendants of Cgtheretta eocaenica KFIJ: Cytheretta cellulosa K T X and Cytheretta carita K,GN; and a descendant of Cytheretta costellata costellata BOSCUGT, C.costel1ata grandipora K W J .
-
w I I
I
Fig.5.
I
I
Distribution o f ostracods i n the Bracklesham Beds of England.
475
The Upper Zocene is equated with the Bartonian, where t w o ostracod zones can be distinguished.
The lower is characterised by
Cytheretta laticosta RELJSS, Cgtheridea intermedia RCLJSS, and Pterygocythereis bartonensis K T I J , all restricted to this zone. The fauna can be recognised in the Barton Clay and in the Xarnes
a
Pholadomya ludensis in the Paris Basin, but appears to be absent in Belgium.
The upper zone contains freshwater, brackish, and marine
horizons, and is characterised by an endemic fauna in southern ngland
.
In northern Aquitaine Mme 3ucasse (1969) has recognised eight biozones for the tocene (Text Fig.2).
These are really assemblage
zones, and to a certain extent reflect environmental conditions. HoTxever, they have proved useful for correlation over a limited area, Many of the species are present in the Anglo-Paris-Belgium area, but have more extensive ranges, thus thro:.:ing some doubt on their iientification.
On the whole, these zones are not easy to correlate
with northdrn Europe.
Faunas from the Western Approaches and the
Xnglish Channel will probably prove helpful in correlating ?pitainn and more northerly regions.
-3.
fauna recfntly describad from thz?
Loire valley near Nantes (Blondeau, 1971) probably belongs to the Cytheretta cellulona zone, and may provide a link with Aquitaine. Typical Atlantic coast "iocene ostracods are listed in 'i'ext Fig.4.
The main Lor;er Yoceni-:fauna is that describt;d by 'ooser
(1965) from the Black ldingo Formation of Eouth Carolinri and from boreholes in North Carolina by "wain ( 1951). been discussed under the Palaeocene.
The former has alreiidy
?coser also describ9d a rich
I.Iid51e 9occn? fauna from the Warley Hill ipormstion snd ?antes Limectone.
From the former he recorded Trzchyleberis spinosissima
(Jonas an? :herborn)
which he regards as a s m i o r synonym of
T.s?iniferrima ( J 3 N T >.NE cX?ZB~R€J); it was suggested that the locality yielding this species was probably older than most k.'arlr:y Hill localities.
If the identification is correct, it suggests a
l o w r Xocens age as this cpscies is typical of the London Clay, belonging to ostracod zone 6b-c.
Xost of the Iv~iddlc.Eocene ostracods
arc? also found in the Upper Eocene, and ind-ed are typical of the Jacksonion stage of the Gulf Coast.
The Upper Yoczne is difficult
to recognise in the ostracod faunas of the Atlantic coast, but is probably represented by part of th? Cooper Marl
476
OLIGOCFSJE There has been a great deal of debate in recent years over the position of the Eocene-Oligocene boundary.
In the Paris Basin there
is a marked faunal break between the Ludian Marnes ‘a P.ludensis and Marnes h Lucina inornata on the one hand and the Sannoidan on the other: of 27 Sannoisian and some 30 Ludian species none are found in common (Keen, 1972a). In England a similar break is seen between the marine Middle Head.on Beds and the Upper Hamstead Beds, the latter containing 13 species all of which are present in the Sannoisian or Rupelian of the Paris Basin. This Sannoisian fauna is also present in Belgium.
The difficulty is in correlating these
beds with the Lattorfian, which defines the base of the Oligocene. The presence of the typical Sannoisian ostracod Hemicyprideis montosa JONES PiNn SHmBORN just below the freshwater Bembridge Limestone of %gland and the Lattorfian Quadracythere diversinodosa (LIENENKLAUS) just above the limestone suggests that the Bembridge Beds might be correlated with the Lattorfian (Text Fig.2). Certainly thr Middle Headon Beds have a strong affinity with the Barton Beds (Keen, 1968). The Rupelian has a very distinctive ostracod fauna (the Sannoisian is regarded as a facies of the Lower Rupelian), with little in common with older faunas or with the Lattorfian.
Several of the species are present in Aquitaine,
allowing accurate correlation. The Upper Oligocene is only really seen in Aquitaine if the study is restricted to the Atlantic seaboard. Here, the Couches h Phare St. Martin at Biarritz contain several species found in the Chattian of Germany and Switzerland, including Flexus gutzwilleri (OTRTLI)
.
477
HAMSTEAD
1
13b
HEADON
I Fig.6.
Distribution of ostracods i n the Upper Eocene and Oligocene of England.
Along t h e e a s t e r n seaboard of N.Amsrica no Oligocene strata a r e d e f i n i t e l y present.
Swain has suggested t h a t C y t h e r e t t a howei
?MAIN might be t y p i c a l f o r t h e Oligocene, but t h e age of t h e rocks c o n t a i n i n g i t a r e not c l e a r .
Pooser placed t h e Cooper Marl i n t h e
Oligocene, but t h e fauna he r e c o r d s seems t o i n d i c a t e two a g e s , Upper Zocene and Miocene. Cytheretta alexander,
HOP!?
HOl,:K
The o l d e r group of samples c o n t a i n s & CHUIBYRC, Haplocytheridert montgomerxensis
& CH4iE3l.RS and Trachyleberis f l o r i e n s i s (HO!XE & CHAKEGRT,) which
s t r o n g l y suggest a Jacksonian age, i.e.
Upper Eoccne.
Pooser
suggests t h i s might c o n s t i t u t e a s e p a r a t e b i o s t r a t i g r a p h i c a l u n i t . The same fauna i s a l s o recorded from a l o c a l i t y r e f e r r e d t o t h e Miocene Duplin Marl by ?ooser, and t h i s u n i t must a l s o be Upper n
:'>ocene.
The younger assemblage, which i s more t y p i c a l of t h e
Cooper Marl, xas r e f e r r e d t o t h e Henrghowella evax Assemblage Zone, and t h e s p e c i e s l i s t e d suggest a Miocene age.
In f a c t , t h e main
reason f o r an Oligocme age was t h e apparent overlap of Eocene and
418 Tdiocene s p e c i e s , which i s o b v i o u s l y removed i f two d i f f e r e n t f a u n a s from two d i f f e r e n t h o r i z o n s a r e p r e s e n t . NS0G”NE
O s t r a c o d f a u n a s o f t h i s a g e have been d e s c r i b e d from s o u t h e r n She c o n t r o v e r s i a l l o c a l i t y o f Escornbgou i n s o u t h e r n
France.
A q u i t a i n e h a s y i e l d e d a r i c h f a u n a w i t h some s p e c i e s i n common w i t h t h e C h a t t i a n o f B i a r r i t z , Germany, and S w i t z e r l a n d b u t a l s o w i t h many s p e c i e s u s u a l l y r e g a r d e d as t y p i c a l l y Miocene: c a n a l i c u l a t a :REIJ
, Cnestocgthere
truncata
Callistocgthsre
(RE%?),
t r i - o s t a t a (RT.Jc~c), and Iismicythere d e f o r m i s (RXJUF,S).
:>!any o f t h e
apeci?s are p r c s m t i n t h e c l a s s i c l o c a l i t i e s of n o r t h e r n Aquitaine, b u t ~ i o m e t y p i c a l Xiocene g e n e r a s u c h as A u r i l a and ? u r i a n a a r e ‘?he Fscornb4ou h o r i z o n i s u n l o u b t e d l y o l d e r t h a n t h e t y p e
abypnt,
. i t a i n i a n , F O i f t h a l a t t e r d e f i n e s t h e base of t h e Miocene, ,,scornbeoa mu--t b s o f Oligocenr: age. ? h > ? > q u i t a i n i a nand younger o s t r a c o d f a u n a s o f t h e t y p e a r e a have bqen t h o r o u g h l y d e s c r i b e d by :Coyc?s (1965’1.
He found t h a t t h e
d i s t r i b u t i o n o f many s p e c i e s was s t r o n g l y i n f l u e n c e d by t h e changes . c i t h i n a shal‘io;. c o a s t ? ? environment, so i t was n o t e a s y t o
ii f f e r e n t i a t f t h ? c l a s s i c a l stagrs.
The Lower [diocene ( A q u i t a n i a n
- Burdigslian’t form-d on? u n i t w h i l e th; Upper Miocene could be divide! i n t o t
30.
1969)
Carborx1.l (
The P l i o c a n e i s m o s t l y u n f o s s i l i f e r o u s . Ctudi-d t h e o s t r a c o d s o f t h e Miocene o f th-? ?hone
Vali-:y and *):as a h a - t o r e c o g n i s e f o u r b i o z o n e s which a r e d i s c u s s e d i n t h . n-xt s e c t i o n .
Thc :dioc.snc: s t r a t a o f t h e e a s t e r n U.5.B.
out.crop from F l o r i d a
t o :krylan:! and have y i s l 4 e d a v e r y r i c h f a u n a , d e s c r i b e d by s u c h a u t h o r s as LTirich & Basnler, HOWF:, i!IcLnan, and ?uri.
Xany o f t h e
Ypncier,: h a v s a l o n g s t r a t i g r a p h i c a l r a n g e w i t h s s v e r a l s t i l l l i v i n g . !iox;;”ver, by u s i n g a s e m b l a g n s i: i s p o s s i b l e t o u s e t h e o s t r a c o d s i n
?mi ( l 9 5 , ? ) h a s i n d i c a t e d t h a t s p e c i e s o f C y t h c r e t t a
corr-istion.
hav-? v e r y l i m i t e i s t r a t i g r a p h i c a l r a n g e s and a r e i d e a l as z o n a i fo=iil?.
T h e i r ta:ionorny d o e s need s o r t i n g o u t though.
r i c a n Zyth5rL:tta
<,ever31 o f
s p e c i e s r e s e m b l e ’lhropean o n a s and i t i s
i n t w e - t i n g t o F p s c i i i a t ? ?h-.thsr t h e s e Were d e r i v e d from ‘%ropean iamigrmts.
Tho Xiocw:
such 4 s ? u r i i a ,
sa?.: t h s r a p i d d i s p e r s a l o f EJ?ogen$ genpra
, and
h r i a n q w i t h some s p e c i e s such as
) r e c o r d e d from b o t h s i d e s o f t h e
479 Atlantic.
Man-made taxonomy probably hides many others.
C'?XOZOIC OSTRACOIX FROT4 WITHIN THT ATLANTIC BA'IN The psychrospheric ostracods (i.e., those specially adapted to abyssal environments) have been studied by Benson (1972a, 197%). In the North Atlantic the three genera Agrenocythere, Bradleya, and F)oseidonamicus have been recorded.
Agrenocythere was probably
derived from Oertliella during the Palaeocene.
The latter is
typical of shelf Cretaceous and ?alaeogene deposits, 0.aculeata (BOSQUIET) being very common during the Eocone.
0.ducassae B?N?ON
was described from the Bartonian of Biarritz and is also found in the
Oligocene o f the 2ockall Plateau; Benson suggests it lived on the outw shelf and upper bathyal zone of western nurope and the N.T. Atlantic.
Oertliella has prominent. eye tubercles, which are lost
as the animal adapts to the psychrosphere where blind forms dominate. Three species of Agrenocgthere are recorded from the ?$.Atlantic: P..antiwata D7NIsOId (1,-X Eocene of ,Europe, America, Carribean; Oligocene of Rockall) regarded as upper bathyal; its descendant, BOL3 (Oligocene A.hazelae VAN !!3
-
Recent, W
&: T;
Atlantic,
Mediterranean, eastern Pacific, Carribean) truly psychrospheric; and P..gosnoldi BEXSON from the I.!I.Foceneof submarine canyons along the N.":.
shelf of U.S.A.,
upper bathyal environment.
The latter
is part of a fauna mentioned by Gibson et.al. (1968) consisting mostly of new species, but with the late Palaeocene Loxoconcha prava ALEMNDER and the M.Socene Trachyleberidea goochi YrlAIN.
Benson
listed a rich fauna from the Rockall Plateau; a Palaeocene fauna with such genera as Agrenocythere, Bairdia, Bythoc.Dris, Cytherella, Aazelina (cf.aranea), Hermanites, and Trachyleberidea (cf. prestwichiana), probably indicating depths of 200-600 rn; and an Oligocene fauna with truly pschrospheric blind species, Agrenocythere, Henryhowella, and Poseidonamicus with Bairdia, Bythoc.Dris, Cytherella and Krithe, indicating depths of lOOO-15OO m as today.
The Rockall story has been continued by Peypouquet
(1975) with drill samples from the Rockall trough.
He found a rich
Lower Miocene fauna with a generic composition similar to Bensonls Oligocene fauna, including the pschrospheric forms. The fauna became impoverished during the Miocene, although pschrospheric species were always present.
The Pleistocene saw a great renewal
of the fauna, but without the pschrospheric forms.
Peypouquet
480
explains their absence then, as today, by a deficiency in ?, N,
&
’1.
caused by a change in oceanic currenfs. B l OSTRATI GRAPH I C ZONES An attempt i s made at a preliminary zonation for the Palaeogene
of tho 4nglo-?aris-3slgim area, with Germany, T’enmzrk, and perhaps the whole :iorth @ea Basin.
Text Fig.; indicates their relrltionship
to the stratigraphy of the area and to other z o n a l schemes. Ostracod biozones. I.
%,imburqinaornatoidella fissurata.
9303
Base defined by first appearance of L.orn.fiss.,
top by first
appearance of Krithe monlensis. I .
Krithe montensis.
>?ROO
Base defined by first appearance of K.montensis, top by first appearance of Alatacgthere cirrusa.
3.
>-latacytherecirrusa.
3LTOO
Base defined by first appearance of P..cirrusa, top by first appearance of C.nerva nerva.
4.
E?IEBX Taracgtheretta reticosa. P.reticosa? Base defined by first appearance of C.nerva nerva, top by
disappearance of P.reticosa.
Probably equivalent to range of
?.reticosa.
5.
Cgtheretta nerva nerva.
P.?OF:T
Baee defined by disappearance of P.reticosa, top by disappearance of C. nerva nerva. 6a.
Cgtheretta sp.nov. cf. laticosta. Base defined by disappearance of C.nerva nerva, top by appearance of C.scrobiculoplicata.
6b.
Cgtheretta scrobiculoplicata (JONES) Base defined by appearance of C.scrobiculoplicata, top by appearance of E.reticulatissum.
6c.
Zchinocgthereis ret ioulatissum (JONES) Base defined by appearance of E.reticulatissum, top by disappearance of C.scrobiculoplicata.
I.
Novoc.mris whitecliffensis (HASKINS) 3ase defined by disappearance of C.scrobiculoplicata, top by disappearance of N.whitec1iffensi.s.
481
8a. Cytheretta
s p . nov
aff.eocesnic2.
aase defined by disappearznce of B.whitecliffensis, top defined by zppearance of Cythererta e o c a e u . 8b.
Cytheretta eocaenic%.
K'IJ
Bast dpfined by appearance of C.eocaenics, t o p by appearance of C.rigida rigida. Cythcridoa rigida rigida.
9.
K4'K dJC
Ease defined by appearance of C.rlg.rigida, 10
.
top by appearance of
C. celluioss. Cythpretta ccllulosa.
K J3:
Bar- defined by appsarance of C.ccllulosa, top by appearance of C.iatlcosta. 11.
Cytherctta laticosta (RLU-.) Basc drfined by appearance of C.laticozta, top by disappearance of Pai,lcnborohella -ocaenica.
1;.
iange zone of C.lasicosta.
Haplocytheridea dsbilis (JOMPS) Base dr,fined by disappearance of Paizenborchella eocaenica TfiIEE *$L, top by appearance of r.diverslnodosa.
138. %adracythere diversinodosa ( L I T 3 ? K L 4 U c ) Base defined by appearance of p.diversinodosa, top by appearance of H.hebertiana. 13b. Kammotocythere hebertiana (BO-CUTP) Base defined by appearance of H,hebertiana, top by appearance of Flexus wtzwilleri. 14.
"lems mtzwilleri (OTRITLI) Base defincd by appearance of F.gutzwilleri, top by appearance of H. deformis.
15.
Hemicythere deformis ( R - U S S ) Bass d3fined by appearance of H.deformis, top by appearance of Aurila.
','hiszone may belong to the FIiocene.
Carbonnel ( 1 9 6 9 ) has produced a zonal scheme for the Kiocene of the Rhone 3asin; this may n o t be applicable over a wide area, however. Biozone A.
Loxoconcha linearis linearis CAXBQ!OJEL
?his coincides with the Aquitanainian * Burdigalian. Biozone B. Neomonoceratina helvetica OXXTLI Lower Zelvetian. Biozone C.
Rhodanicites tripartita CARBONNJCL
Upper Helvetian.
482 Biozone
n.
Z l o f s o n e l l a amberii C!.4RBONX!?L
Squivalent t o t h e Tortonian. Thc Pliocene was not zoned, although c h a r a c t e r i s t i c s p e c i e s a r e listed. itany l i n e a g e s of o s t r a c o d s can be .traced through t h e Palaeogene Genera such as
and t h e s e o f f e r g r e a t scope f o r c o r r e l a t i o n .
C y t h e r e t t a , Cgtheridsa, and L e , m i n o c y t h e r e i s o f f e r g r e a t scope, with Neocgprideis i n t h s b r a c k i s h water sediments.
No zonal schemes a r e a v a i l a b l e f o r .the A t l a n t i c Seaboard of North America, although i t ought t o be p o s s i b l e t o c r e a t e one. Certain genera appear t o be i d e a l .
3pimocythere, as i n d i c a t e d by
Hazel (1968) i s an important genus i n t h e Palaeogene with w i d d y d i s p e r s e d s p e c i e s groups.
several
Vetustocytheridea a l s o h a s
p o t e n t i a l , as does Haplocytheridea, which seems t o occupy t h e same n i c h e as Cytheridea i n &rope. sncm t o have s h o r t ranges.
Xany s p e c i e s of C y t h e r i l l o i d e a
In t h e Ueogene a zonation based on
C y t h e r e t t a seems p o s s i b l e . Brackish water and freshwater o s t r a c o d zones a r e a l s o i n d i c a t e d on Text Fig.2.
FAC I ES R E LATlO NS H I PS T h s e have mostly been d e a l t with i n d i s c u s s i o n of t h e faunas. iFreshwater, b r a c k i s h , and marine environments a r e r e a d i l y recognised. I.!ithin t h e marine environment shallow c o a s t a l , s h e l f , o u t e r s h e l f , b a t h y a l , and a b y s s a l have been mentioned.
The zonal scheme given
above i s mostly f o r t h e shallow and s h e l f s e a s ;
not enough
information i s a v a i l a b l e f o r deeper w a t e r s , but a s t h e r e i s q u i t e a
l o t of mixing of t h e s e faunas t h e r e should be no d i f f i c u l t y i n c o r r e l a t i n g shallow and deep water faunas.
483 REFERENCES
Apostolescu, V., 1955. De'scription de quelques Ostracodes du Lut6tien du Bassin de Paris: Cah. gGol., no. 28/29, p. 241-279. 1956. Contribution l'gtude des ostracodes de -6'. l'Eocbne inferieur (s.1.) du Bassin de Paris: Rev.Inst.frang., Pgtrole, V.ll, p.1327-1352. Benson, R.H., 197.2a. Preliminary Report on the Ostracodes of 9SDP Leg 12, sites 117 and 117.4: Initial Reports of the Deep Sea Drilling Project, V.XII, p.427-432.
.
,1972b. The Bradleya Problem: Smithsonian contr. Paleobiol., no. 12., 138p. Blondeau, X.A., 1971. Contribution 5 lle/tudedes Ostracodes e'ocknes des Bassins de Campbon et de Saffre (Loire-Atlantique): Thesis, Univ. Xantes , I 50p. Carbonnel, G., 1969. Les ostracodes du Ilioc\ne Rhodanien: Docum. Lab.gebl.?ac,tci.Lyon, no.32, 469p. 3amotte, R., 1964. Contribution k l'dtude des "calcaircs montiens" du Bassin de Paris: la fame d'0stracodes: Bull,soc.g6ol.Prance, (7) VsVi, ~$349-356. 3)groo,G., 1965. Cytheracea (Ostrscodes) du idsastrichticn de !6aastricht (Pays-Bas): ;~eded.gcol.Stichf., ser.C, no.?, 197p. hcasse, O., 1969. Biozonation de 1'':ockne nord-aquitain: Bull. 'oc.geol.i7rance, (7) V. XI, p.491-501.
.,
197;. ies ostracodes de la coupe de Campo: Revista , p. 573-289. Kspan. Yicropaleont., V. %gar, r;.I-I., 1965. London Basin: i. p. 15-32.
Cistracoda of the London Clay (Ypresian) in the Reading District: Revue Xicropalebnt, V . 8 ,
Gibson, ri,G., Hazel, J.X., and IbIello, J.F.., 1968. Fossiliferous rocks from submarine canyons off the northeastern United States: prof .p&p.lj. ':.Gaol. 'urv. , 600-0. Gokpn, N., 1971. Las Ostracodes de l'Ypr4sien de l'ouesl du Dassin de tondres: Bull.min.expl.Inst.'i'urkey, no. 75, p.69-86.
1968 - 1971. Tertiary ostracoda from the Isle of blight and Barton, Hampshire, %gland: Revue !4icropaleont, V.10, p.250-260; V.11, p,3-I?, p.161-175; \!.I?, p.149-170; V 1 3 , D.13-Zyr p.207-iil; V.14, p.147-156.
Y a s k i n s , C.W.,
Iiazel, J.C., 1968. Ostracodes from the Brightseat, Formation (Danian) of IIaryland: J.?aleont., 8.42, p. 100-142.
484
Keen, M.C., 1968. Ostracodes de 1lEochne supkrieur et llOligoc8ne infe
.,
1972a. The Sannoisian and some other Upper Palaeogene Ostracoda from north-west Europe: Palaeontology, V.15, p.267-325.
., 1972b. Evolutionary patterns of Tertiary ostracods: Froc.Int.Geol.Congr. , 24(7), p.190-197. Keij, A . J . , 1957 Eocene and Oligocene Ostracoda of Belgium: Mem. Inst.Roy.Sci.Nat .Belgique, V. 136. Marliere, R., 1958. Ostracodes du Piontien de Mons et resultats de leur e'tude: Mkm.Soc.Belg.gdol., V.5, p. 1-53. Moyes, J., 1965. Les Ostracodes du Miockne Aquitain: Univ.Bordeaux, 338 p.
Thesis,
Peypouquet, J.P., 1975. La renouvellement de la f a m e d'0stracodes du bassin de Rockall entre le Miockne et le Pl&stockne sup6rieur.: Bull. SOC. Giol. France, (7) V. XVII, p . 886-895. Pooser, W.K., 1965. Biostratigraphy of Cenozoic Ostracoda from South Carolina: Paleont.Contr.Univ.Kansas, Arthropoda, Art.8., 80p. Ostracode genera Cytheretta and Paracytheretta Puri H.S., 1952. in America: J. Paleont., V.26, p.199-212. Swain, I".I?., 1951. Ostracoda from wells in North Carolina, Part 1, Cenozoic Ostracoda: Prof.pap.U.S.Geol.Surv., 234-A, 58p. Szcezechura, J., 1965. Cytheracea (Ostracoda) from the uppermost Cretaceous and lowermost Tertiary of Poland: Acta paleont.po1. V. 10, p.451-564.
,, 1971. Paleocene Ostracoda from Nbgssuaq, West Greenland: Bull.Gronlands geol.under., no.94, 42p. Tambarenu, Y., 1972. Thanetien supgrieur et Ilerdien infgrieur des Petites Pyrehees, du ?lantaurel et des chainons audois: Thesis, Univ.Toulouse, 37713. Tambareau, Y., and Villatte, J., 1974. Le Passage T'hanetienIlerdien dans la region de Campo: Bull.Soc.hist.nat.Toulouse, V. 110, p.340-361 Willems, !"I., 1975. Ostracoda from the Ieper Formation of the Kallo well (Belgium): Bull.Soc.g
485
PLATE 1. Figs. 1 and 3, C.ytheridea p r i m i t i a HASKINS; F i g . l , L = 0.55, Fig.2, L = 0.55. Fig. 2, C y t h e r e t t a eocaenica KEIJ;
London Clay, Alum Bay;
L u t e t i a n , Damery;
L = 0.80.
Pigs. 4 and 6 , Cytheridea r i g i d a r i g i d a HASKINS; Bracklesham W h i t e c l i f f Bay; Fig.1, L = 0.70, Fig.2, L = 0.69. Fig. 5, C.ytheretta c e l l u l o s a Km; L = 0.78.
Beds,
Sables de Beauchamp, Moiselles;
Figs. 7 and 9, Cytheridea intermedia (REUSS); Barton Clay, Barton; Fig.7, L = 0.B8, Fig.8, L = 0.86. Fig. 8, C y t h e r e t t a haimeana (SOSQUET);
L u t e t i a n , Damery;
L = 0.60.
Figs. 10 and 1 2 , Cytheridea pernota OERTLI & KEIJ; Couches de Sannois, Cormeilles; Fig.10, L = 0.76, Fig.12, L = 0.15. Fig. 11, C y t h e r e t t a s c r o b i c u l o R l i c a t a (JONES); Kingsclere; L * 0.92. Fig. 1 3, C y t h e r e t t a l a t i c o s t a (REUSS); L = 0.77, Fig. 14, C y t h e r e t t a f o r t i c o s t n KuW; Bay; L = 0.79. Fig. 15, C y t h e r e t t a porosacosta Bay; L = 0.75.
London Clay,
Barton Clay, Barton; Bracklesham Beds, W h i t e c l i f f
K m ; Middle Headon Beds, Colwell
Fig. 16, O e r t l i e l l a a c u l e a t a (BOSOUET); Verzy; L = 0.73.
Marnes a P.ludensis,
Fig. 17, C,ytheretta c o s t e l l a t a c r a t i s KEEN; Verzy; L = 0.73.
Marnes a P.ludensis,
,
Fig. 18 Trachyleberidea prestwichiana (JONXS AND SmBORN) ; London Clay, V h i t e c l i f f Bay; L = 0.65. Fig. 19, P a r a c y t h e r e t t a r e t i c o s a TRIXBXL; L = 0.78,
Thanet Beds, IIerne Bay;
Fig. 20, C y t h e r e t t a c o s t e l l a t a c o s t e l l a t a ( R O W B ) ; Bracklesham Beds, Selsey; L = 0.64.
486
PLATE 1
487
PLATE 2. Figs. 1 and 4, Haplocgtheridea montgomeryensis (HOWE and CHAPqfBEE3); Jackson, Louisiana; Fig.1, L = 0.83, Fig.4, L = 0.83. Fig, 2 , t c h i n o c g t h e r e i s Aacksonensis (HOWE and PYEATT) ; Jackson Creole B l u f f , Louisiana; L = 0.98. Fig. 3 , Zchinocythereis r e t i c u l a t i s s i m a EACAR; Sheppey; L = 0.93.
London Clay,
Fig. 5, T r a c h y l e b e r i s f l o r i e n s i s (Holm and CHAbBiIswS) j Creole B l u f f , Lauisiana; L = 0.82. Fig. 6 , !?uadracythere diversinodosa (LIETGXELAUS); “ h i t e c l i f f Bay; L = 0.66. Fig. 7 , C y t h e r e l l o i d e a montPorneryensis HOKE; Louisiana; L = 0.66,
Bembridge Beds,
Jackson Creole B l u f f ,
Fig. 8 , T a i j e n b o r c h e l l a eocaenica “RIFBEL; Verzy; L = 0.60.
Marries a P.ludensis,
Fig. 9 , Dimocythere r u s s e l l l (HOVE and L I X ) ; Louisiana; L = 1.20. Fig.10, C j t h e r e l l o i d e a darnerlacensis AFOST. I ’ h i t e c l i f f Day; L = 0.72. Fig. 11, Haplocytheridea d e b i l l s (JONES); Headon H i l l ; L = 0.62.
Jackson,
Jackson Creole B l u f f , Bracklesham Beds,
Pliddle Headon Beds,
Fig.11, C g t h e r e t t a alexander; HOKE and CHAIBERS; B l u f f , Luuisiana; i = 0.90. Fig. 1 3 , ” t e r y g o c y t h e r e i s p u s t u l o s a IiATKINS; Verzy; L = 0.79.
Jackson Creole
M a n e s a P.ludensis,
Fig. 14, Legumlnocythereis s t r i a t o p u n c t a t a ( R O 9 F E ) ; Bracklesham Beds, I : h i t e c l i f f Bay; L = 0.96. Fig. 15, Zocgtheropteron w e t h e r e l l i (JONE? and ?HERBORN), IJiddle L = 0.70. Headcn Beds, Headon H11l; Fig. 16, Hammatocythere o e r t l i i (DUCASS3); L = 0.85. Fig.
17, L e w i n o c y t h e r e i s o e r t l i i KGIJ;
Upper Socene, Blaye;
Sables de Lede, Bambrugge;
L = 0.84. Fig. 18, ?arac.ypris c o n t r a c t a (JON::) Verzy; = 0.96.
; Idarnes a P.ludensis,
Fig. 1 9 , L e g m i n o c g t h e r e i s d e l i r a t a lliddle Headon Beds, Headon H i l l ; 1, = 0.82.
488
PLATE 2
489 PLATS 3.
Fig. 1 , Cladarocgthere apostolescui (IfiRCWITE); Bembridge Limestone, Rouldnor Cliff; L = 0.73. Fig. 2, Fchinocythereis scabra (VOii i.IUSNST4;R) ; Gscornbeou; L = 0.85. Fig. 3, Iluellerina latirnarginata (':?Em);Chattion, Bunde; L = 0.69. Fig. 4, Cgtheretta geoursensis KEEN; Fig.
Escornbeou; L = 107.
5, Crestocsthere truncata (REUSS); Escornbeou; L
=
0.62.
Fig. 6, Hammatocythere hebertiana (BOSXET) ; Rupelian, Auvers-'tGeorge; L = 0.86. Fig.
7, Hemlcythere deforms minor
:JOy-iS;
Fscornbeou; L = 0.64.
Fig. 8, Cgtheretta tenuipunctata absoluta KEEN; Cormeilles; 1, = 0.84.
Marnes a Huitres,
Fig. 9, Callistocythere canaliculata (RWSS); Escornbeou; L = 0.55. Fig. 10, Aurila conradi conradi (HOVE and McGUIRT); Xarl, Florida, L = 0.60.
Choctawhatchee
Fig. 11, ProtocytherettaKarlana (HOkJE and P m T T ) ; Florida; L = 1.04.
Chipola Xarl,
Fig.12, Cstheretta orthozensis IIOYXS;
Helvetian, Orthez; L = 0.86.
Fig. 13, Henryhowella evax (ULRICH and BASSLGR); Chipola Liar1 , Florida; L = 0.82. Figs.14 and 18, Cgtheretta burnsi (ULRICH and BASSLIE'); 7ig.14 Calvert Formation, Calvert, L = 1.05; Fig.18 Area ?,one,Red Bay, Florida, L = 1.10. Fig.15, Cytheretta sahni PURI;
Duplin Marl, Florida; L = 0.98.
Fig. 16, Cgtheretta inaeauivalvis (ULRTCH and BASSLZR); Facies, Alum Bluff, Florida; L = 0.90.
Choal River
Fig. 17, Actinocythereis exanthernata (ULRICH and BASCLW); IIarl, Florida; L = 0.86.
Chipols
490
PLATE 3
191
PL,lTE 4. Pig. 1 , ‘ i i r p a t o c y p r i s g r i s y e n s i s ( LUGIEIT} ; L = 0.86. Fig. 2 , Virgatocypris edwardsi K Y X p!.S.; Colwell Say; L = 1.13.
Ludian, Verzy;
Upper IIeadon Beds,
Fig. 3 , i’irmtoc,ypris t e n u i s t r i a t a , (DOLLITIS); Cormeilles; L = 1.01.
‘jannoisian,
Figs. /i and 5, Hemicyprideis montosa ( J O N T F and ‘HYIBQFI;); Hamstead Beds, Bouldnor C l i f f ; l i g . 4 , L = 0.78; F1.g.5, L = 0.80.
Fig. 6 , Cytheromorpha b u l l a HASKITS; 1) = 0.58.
Headon Beds, Headon H i l l :
Fig. 7 , Iiemic,mrideis h e l v e t i c a (LI%V@JKL&US) ; Blaue T o r , Delemont, Switzerland; L = 0.70. Pig. 8 , Neocyprideis c o l w e l l e n s i s ( J O U T ) ; L = 9.63.
9, Neoc.yprideis
Fig.
IIezdon Beds, Headon I I i l l ;
d u r o c o r t o r i e n s i s APOS’T. ;
~ p p a m a c i s n , Ypernay;
L = 9.81. Fig. 10, Neocgarideis williamsoniana (BOSQUTT) ; Sannoisian, Cormeilles; L = 3.83. Fig.11, Neoc.yprideis a p o s t o l e s c u i ( K T J ) ; X o i s e l l e s ; L = 0.79.
::ables de Beauchamp,
Fig. 1 2, Agrenocythere a n t i q u a t a BZNS0N; Upper Zocene, Trinidad; L = 1.17. Photograph k i n d l y donated by D r . R.H. Benso n. Fig.13, Am enocythere gosnoldia BZNSON; BIiddle Zocene, A t l a n t i c Slope o f f Cape Cod; L = 1.14. Photograph k i n d l y donated by D r . R.H. Benson. Fig. 14, A n e n o c y t h e r e hazelae (VAN D E 3 BOLD) ; Xiocene, Cipero Formation, Trinidad; L = 1.10. Photograph k i n d l y donated by D r . R.H. Benson. Fig.15,
L
=
Candona d a l e y i KEmT b1.S.;
0.87.
Lower Headon Beds, Headon H i l l ;
49 2
PLATE 4
493
Discussion F . M . Swain: I n Pooser's paper t h e species l i s t s f o r t h e s o - c a l l e d Cooper Marl, O l i g o c e n e , and f o r t h e l a t e Miocene D u p l i n Marl w e r e r e v e r s e d i n p r i n t i n g - h e l a t e r s e n t a c o r r e c t i o n s h e e t . T h i s m i g h t a c c o u n t f o r your comment t h a t t h e O l i g o c e n e forms seem t o be of Miocene t y p e . Keen: T h a t m i g h t e x p l a i n i t . Swain: The o t h e r p o i n t i s w i t h r e f e r e n c e t o your remark a b o u t t h e l a t e Eocene a s p e c t of t h e d e s c r i b e d forms from t h e A t l a n t i c c o a s t a l p l a i n . Most of t h e c o l l e c t i o n s from t h e Eocene t h a t have been d e s c r i b e d from t h a t a r e a a r e e a r l y and m i d d l e Eocene, p r o b a b l y m o s t l y m i d d l e Eocene. T h e r e i s a n e x c e l l e n t l a t e Eocene o s t r a c o d e f a u n a i n S o u t h C a r o l i n a s i m i l a r t o t h a t d e s c r i b e d by Huff from Mississip p i . The Eocene forms from N o r t h C a r o l i n a a r e a l l , o r n e a r l y a l l e a r l y and m i d d l e Eocene t y p e s , n o t l a t e Eocene. Keen: I m a i n l y g o t t h e s e from P o o s e r a s t h e main r e f e r e n c e . I c e r t a i n l y t h o u g h t , l o o k i n g a t most of t h e s p e c i e s r e c o r d e d f o r t h e m i d d l e Eocene i n p a r t i c u l a r were o n e s t h a t a r e found i n t h e l a t e Eocene o f t h e Gulf C o a s t . I a l s o t h o u g h t t h a t t h e s o - c a l l e d O l i g o c e n e f o r m a t i o n , t h e Cooper M a r l , seemed t o m e t o b e d e f i n i t e l y l a t e Eocene on t h e s p e c i e s he r e c o r d e d . Swain: The r e a s o n f o r t h a t i s t h e l i k l i h o o d t h a t some of t h e forms r e c o r d e d f o r t h e O l i g o c e n e came from t h e u n d e r l y i n g l a t e Eocene. Dr.
This Page Intentionally Left Blank
495 C E N O Z O I C 1,lARINE OSTRACODA OF THE SOUTH ATLANTIC W.
A . van d e n Bold
L o u i s i a n a S t a t e U n i v e r s i t y , Baton Rouge
Abstract T e r t i a r y O s t r a c o d a o f t h e A t l a n t i c Ocean s o u t h o f t h e T r o p i c o f Cancer a r e known from w i d e l y s c a t t e r e d c l u s t e r s o f l o c a l i t i e s on t h e c o n t i n e n t and from a number o f d e e p - s e a c o r e s . The l a t t e r h a v e o n l y b e e n s t u d i e d i n d e t a i l f o r p a r t o f t h e i r f a u n a . P a l e o c e n e o s t r a c o d e s have b e e n s t u d i e d i n f o u r a r e a s : C e n t r a l America (Guatemala and B r i t i s h H o n d u r a s ) , n o r t h e r n S o u t h America ( V e n e z u e l a , T r i n i d a d , Guiana ) , A r g e n t i n a and West A f r i c a . The d i s t r i b u t i o n o f some g e n e r a ( e . g . B u n t o n i a , S o u d a n e l l a , T o g o i n a ) i n d i c a t e s s i m i l a r i t i e s between f a u n a s from w i d e l y d i f f e r e n t a r e a s and s u g g e s t s t h a t c u r i n g o r s h o r t l y p r i o r t o t h e e a r l y T e r t i a r y some s h a l l o w - w a t e r o s t r a c o d e s made t h e i r way a c r o s s t h e A t l a n t i c e i t h e r t h r o u g h s h a l l o w p a s s a g e s o r a t t a c h e d t o seaweed d r i f t i n g w i t h p r e v a i l i n g c u r r e n t s . Eocene o s t r a c o d e s a r e known from West A f r i c a and T r i n i d a d , b u t from v e r y d i f f e r e n t env i r o n m e n t s , which makes comparison h a z a r d o u s . O l i g o c e n e o s t r a c o d e s have o n l y b e e n r e p o r t e d from t h e C a r i b b e a n and from some d e e p - s e a c o r e s . Mioc e n e o n e s h a v e b e e n d e s c r i b e d from West A f r i c a ( G a b o n ) , t h e C a r i b b e a n , n o r t h e r n S o u t h America and A r g e n t i n a ( i n c l u d i n g s o u t h e r n B r a z i l ) . Here t h e p r e s e n c e o f m o r p h o l o g i c a l l y s i m i l a r g r o u p s ( S o u d a n e l l a ) i n West A f r i c a and s o u t h e r n S o u t h America and t h e r e p o r t o f t h e M u n s e y e l l a ? p u n c t a t a g r o u p from s o u t h e r n B r a z i l and n o r t h e r n S o u t h America s u g g e s t c o n n e c t i o n s o r par a l l e l f a u n a l d e v e l o p m e n t s which w i l l have t o be i n v e s t i g a t e d i n more det a i l . The West A f r i c a n f a u n a b e a r s a s t r o n g s i m i l a r i t y t o t h e M e d i t e r r a n ean o n e . F o r t h e P l i o c e n e - H o l o c e n e , o s t r a c o d e s a r e known from t h e same a r e a s a s i n . t h e Miocene and t h e d i s t r i b u t i o n o f t h e a u s t r a l form Patagonocythere as w e l l a s t h e p o s s i b l e p r o v i n c i a l i t y o f s u c h forms a s C y p r i d e i s , P e r i s s o c y t h e r i d e a , S u l c o s t o c y t h e r e and t h e d i s t r i b u t i o n o f C a l l i s t o c y t h e r e and N e o m o n o c e r a t i n a a p p e a r t o b e o f g r e a t p o t e n t i a l i n t e r e s t .
Zusammenfassung T e r t i z r - O s t r a c o d e n d e s A t l a n t i s c h e n Ozeans s i d l i c h d e r Krebstwende s i n d von r a u m l i c h w e i t v e r b r e i t e t e n L o k a l i t z t e n und e i n e r Anzahl T i e f s e e - B o h r k e r n e b e k a n n t . Nur e i n k l e i n e s T e i l d e r Fauna d i e s e r l e t z t e n i s t e i n e r grundl i c h e n D e t a i l - F o r s c h u n g a u s g e s e t z t worden. Palaozsne Ostracoden s i n d i n v i e r Gebiete s t u d i e r t worden, n h l i c h Z e n t r a l Amerika, n o r d l i c h e s SudameriF a u n i s t i s c h e c b e r e i n s t i m m i n g zwischen k a , A r g e n t i n i e n und West A f r i k a . einzelnen Gebiete i s t angedeutet durch k a r a k t e r i s t i s c h e Verbreitung e i n i g e r G e s c h l e c h t e und l h s t f r z h t e r t i & e o d e r e i n wenig a l t e r e s e i c h t e t r a n s a t l a n t i s c h e M e e r e s v e r b i n d u n g e n , o d e r a b e r B e f o r d e r u n g von F l a c h w a s s e r a t r a c o d e n m i t t e l s von Meeres-Stromungen v e r f r a c h t e t e s P f l a n z e n m a t e r i a l s , vermuten. Eozzne A r t e n s i n d a u s West A f r i k a und T r i n i d a d b e s c h r i e b e n , a b e r Nur a u s den f a z i e l l e t i n t e r s c h i e d e v e r s c h w e r e n e i n V e r g l e i c h d e r Faunen. K a r i b e n und e i n i g e n Tiefsee-Bohrungen s i n d O l i g o z 2 n e O s t r a c o d e n b e k a n n t . Miozane A r t e n s i n d von West A f r i k a ( G a b o n ) , dem K a r a i b i s c h e n G e b i e t , Venezu e l a , K o l o n b i e n , A r g e n t i n i e n und sGdlichem B r a z i l g e m e l d e t und d a s Vorkommen m o r p h o l o g i s c h a h n l i c h e r Formen ( w i e S o u d a n e l l a i n A f r i k a und SGdamerika) und d i e M u n s e y e l l a ? p u n c t a t a Gruppe i n n o r d l i c h e m und s g d l i c h e m sifdam e r i k a ) z e i g e n Verbindungen o d e r p a r a l l e l e E n t w i c k l u n g s g e s c h i c h t e n an. D i e
496 West A f r i k a n i s c h e Fauna h a t v i e l e m e d i t e r r a n e E l e m e n t e . I m Jung-Caenozoicum s i n d d i e Faunen u n t e r s u c h t i n f a s ; d e n s e l b e n G e b i e t e n und w e i t e r e S t u d i e n d e r V e r b r e i t u n g und P r o v i n c i a l i t a t d e r G a t t u n g e n P a t a q o n o c y t h e r e , Cyp r i d e i s , Perissocytheridea,Callistocythere,S u l c o s t o c y t h e r e , Neomonocerat i n a U.S.W. k 8 n n t e w i c h t i g e n w e i t e r e n Daten h e r v o r b r i n g e n . PREVIOUS WORK D i s t r i b u t i o n o f Recent O s t r a c o d a was summarized by P u r i ( 1 9 6 7 ) , who r e c o g n i z e d a N o r t h American B o r e a l P r o v i n c e ( N . o f t h e C a r o l i n a s ) , a C a r i b b e a n P r o v i n c e , s u b d i v i d e d i n t o Bahaman, s o u t h F l o r i d a n and Venezuelan s u b p r o v i n c e s and a Gulf o f Mexico P r o v i n c e .
The S o u t h A t l a n t i c was n o t d i v i d e d i n t o
p r o v i n c e s b e c a u s e o f i n s u f f i c i e n t d a t a and t h e A n t a r c t i c was n o t d i s c u s s e d . Hartmann ( 1 9 7 5 ) summarized t h e d i s t r i b u t i o n o f R e c e n t O s t r a c o d e s from West A f r i c a and made c o m p a r i s o n s t o t h e P a c i f i c C o a s t o f S o u t h America.
But
a p a r t from H a r t m a n n ' s ( 1 9 5 5 , 1956) s t u d i e s o f t h e f a u n a of s x t h e r n B r a z i l and some i n f o r m a t i o n on t h e A r g e n t i n e c o a s t by Whatley and Moguilevs k y ( 1 9 7 5 ) n o t much i s known a b o u t t h e Recent o s t r a c o d e s o f t h e w e s t e r n s i d e of t h e A t l a n t i c .
The summary o f t h e A r g e n t i n e f o s s i l o s t r a c o d e - f a u n a s
by B e r t e l s ( 1 9 7 5 ) g r e a t l y f a c i l i t a t e s t h e t a s k o f making c o m p a r i s o n s w i t h other areas.
However, n o t h i n g i s known o f t h e f o s s i l . o s t r a c o d e s on t h i s
s i d e of t h e A t l a n t i c from. a b o u t 5'N
t o 30°S.
Data on t h e l o c a t i o n o f Neo-
g e n e C a r i b b e a n s u b p r o v i n c e s c a n be found i n van d e n Bold ( 1 9 7 0 a , 1 9 7 1 ~ ~ 1?74a, 1 9 7 5 ) . C o m p a r a t i v e l y l i t t l e i s known a b o u t T e r t i a r y f a u n a s from t h e w e s t c o a s t of A f r i c a (Paleogene: gene:
A p o s t o l e s c u , 1 9 6 1 and Reyment, 1963-1966; Neo-
van d e n B o l d , 1 9 6 6 h ) , and no i n f o r m a t i o n a t a l l e x i s t s s o u t h of t h e
5 O S p a r a l l e l e x c e p t a r e c e n t s t u d y b y . D i n g l e on t h e P a l e o c e n e o f t h e e a s t
coast. LIMITATIONS OF THE DATA Although i n t h e A t l a n t i c Ocean i t s e l f w e h a v e some v e r y d e t a i l e d i n f o r m a t i o n on some g r o u p s o f deep-water o s t r a c o d e s ( A q r e n o c y t h e r e , B r a d l e y a , P o s e i d o n a m i c u s , A b y s s o c y t h e r e ) t h a n k s t o Benson ( 1 9 7 1 , 1 9 7 2 ) and t h e Deeps e a D r i l l i n g Program, l i t t l e i s known o f some o f t h e o t h e r f o s s i l g r o u p s , and even t h e d i s t r i b u t i o n o f R e c e n t forms i s s t i l l v e r y i n s u f f i c i e n t l y known.
Also i n t h e b o r d e r l a n d s it i s d i f f i c u l t t o o b t a i n a n o v e r a l l p i c -
t u r e o f t h e d i s t r i b u t i o n b e c a u s e t h e i n f o r m a t i o n i s g e n e r a l l y from w i d e l y s e p a r a t e d l o c a l i t i e s and moreover s e v e r e l y r e s t r i c t e d s t r a t i g r a p h i c a l l y . From West A f r i c a t h e i n f o r m a t i o n i s l i m i t e d t o t h e Danian-Lower Eocene ( A p o s t o l e s c u , 1 9 6 1 , Reyment, 1963-1966) arid t h e Lower Miocene t o P l i o P l e i s t o c e n e o f o n e s m a l l a r e a o n l y (var. d e n B o l d , 1 9 6 6 h ) .
I n Argentina
497 T e r t i a r y f a u n a s a r e r e p o r t e d from t h e Danian, Miocene ( n o t always c e r t a i n from what l e v e l s ) and P l e i s t o c e n e ( B e r t e l s , R o s s i d e G a r c i a ) . o n l y P a l e o c e n e f a u n a s a r e known.
I n Guiana
I n Venezuela and T r i n i d a d t h e r e i s f a i r l y
c o n t i n u o u s c o v e r a g e from P a l e o c e n e t o P l e i s t o c e n e , b u t o n l y deep-water f a u n a s a r e known from Middle and Lower Eocene and o n l y s h a l l o w - w a t e r f a u n a s from t h e Upper Eocene.
I n t h e Caribbean t h e r e i s o n l y s c a t t e r e d (and
l a r g e l y u n p u b l i s h e d ) i n f o r m a t i o n on a n y t h i n g o l d e r t h a n O l i g o c e n e , a l t h o u g h t h e Neogene i s f a i r l y w e l l known.
I n C e n t r a l America P a l e o c e n e and some
Lower E o c e n e a r e known from Guatemala and B r i t i s h Honduras, no Middle and Upper Eocene f a u n a s h a v e been r e p o r t e d and t h e Neogene i s o n l y known r e l a t i v e l y w e l l from Panama, C o s t a Rica and Guatemala. T h e r e f o r e I have summarized t h e more i m p o r t a n t d a t a f o r each r e g i o n i n t a b u l a r form, u s i n g o n l y t h e s p e c i e s t h a t were f a i r l y common o r had a relat i v e l y wide d i s t r i b u t i o n ; i n p l a c e s where no d e t a i l e d s t r a t i g r a p h i c i n f o r mation was a v a i l a b l e t h e s p e c i e s a r e m e r e l y l i s t e d . Myodocopida and f r e s h w a t e r o s t r a c o d e s have been o m i t t e d from t h e d i s c u s s i o n b e c a u s e o f t h e s c a r c i t y o f r e f e r e n c e s t o them i n t h e l i t e r a t u r e on Tertiary ostracodes.
The l a s t were summarized by McKenzie (1973), w h i l e
K o r n i c k e r d e v o t e d many i m p o r t a n t p a p e r s t o t h e d i s t r i b u t i o n of Recent Myodocopida.
A number o f o t h e r
forms h a v e been l e f t o u t o f t h e d i s c u s s i o n :
The s p e c i e s d e t e r m i n e d by A p o s t o l e s c u (1961) a s Orionina and Ambocythere a r e c e r t a i n l y d i f f e r e n t from t h e s e g e n e r a .
Several species referred t o
Anticythereis have been t r a n s f e r r e d t o o t h e r g e n e r a , b u t I have r e t a i n e d t h o s e t h a t r e s e m b l e t h e t y p e s p e c i e s a n d / o r s p e c i e s r e f e r r e d t o t h a t genus by B e r t e l s , i n o r d e r t o a r r i v e a t some c o n s i s t e n c y .
Some o f t h e s p e c i e s
d e s c r i b e d as Phacorhabdotus ( s e e a l s o Omatsola, 1 9 7 2 ) d o n o t b e l o n g t o t h a t genus.
I n t h e s e and some o t h e r c a s e s i t becomes v i r t u a l l y i m p o s s i b l e t o
make c o r r e c t g e n e r i c a s s i g n m e n t s w i t h o u t s t u d y o f t h e o r i g i n a l m a t e r i a l and t h e r e f o r e t h e s p e c i e s c a n n o t b e u s e d f o r comparison w i t h p o s s i b l y similar groups i n o t h e r a r e a s . A l s o many o f t h e unornamented g r o u p s have been l e f t o u t a s i t is v e r y
d i f f i c u l t t o draw c o n c l u s i o n s from t h e d i s p e r s a l o f s u c h g e n e r a a s Cyther-
ella, Bythocypris, Bairdia, Propontocypris and o t h e r s , and it would be o u t s i d e t h e s c o p e o f t h i s p a p e r t o i n c l u d e d e t a i l e d i n f o r m a t i o n on t h e s e .
The
same i s t r u e of t h e g e n e r a Haplocytheridea, Cyamocytheridea, Ovocytheridea,
Cyprideis, Cushmanidea and o t h e r s w i t h o u t s t u d y o f t h e o r i g i n a l m a t e r i a l . One f u r t h e r problem i s t h e i d e n t i t y o f t h e g e n u s Rocaleberis B e r t e l s ,
1969.
The d i s t i n c t i o n from Henryhowella l i e s , a c c o r d i n g t o B e r t e l s , i n t h e
p r e s e n c e o f a v e s t i b u l e and o c c a s i o n a l b r a n c h i n g o f t h e m a r g i n a l p o r e c a n a l s
However, i n Henryhowella ex g r . a s p e r r i m a , t h e r e i s a v e s t i b u l e and b r a n c h i n g p o r e c a n a l s b o t h i n t h e European material ( e . g . K e i j , 1957, R u g g i e r i ,
1 9 6 2 ) and i n t h e C a r i b b e a n ( v a n den B o l d , 1 9 6 0 ) .
Therefore, a t t h e present
t i m e , I c a n n o t s e p a r a t e t h e two f o r m s , and I have k e p t t h e name R o c a l e b e r i s f o r t h e Danian s p e c i e s and i n d i c a t e d t h e d i s t r i b u t i o n o f Henryhowella i n o t h e r areas u n d e r t h e same h e a d i n g w i t h q u e s t i o n m a r k s .
I t should be noted
t h a t R o s s i d e G a r c i a m e n t i o n s a Henryhowella c f . evax from t h e E n t e r r i e n s e i n n o r t h e r n A r g e n t i n a , and S a n g u i n e t t i o n e from t h e Miocene o f t h e P e l o t a s basin i n southern Brazil. The genus M a s i u k c y t h e r e R o s s i d e Garcia (1968) is d e s c r i b e d from t h e Danian o f A r g e n t i n a .
B e r t e l s h a s i g n o r e d t h i s genus i n h e r summary o f
A r g e n t i n i a n o s t r a c o d e s (1975) and d o e s n o t r e f e r t o it i n o t h e r p u b l i c a tions.
I have t h e r e f o r e i g n o r e d i t i n t h i s p a p e r .
A number o f o t h e r
u n c e r t a i n t i e s w i l l b e mentioned i n c o n n e c t i o n w i t h t h e t a b l e s , t h e i n f o r mation on which i s s e l e c t i v e and t h e r e f o r e o b v i o u s l y b i a s e d . ACKNOWLEDGEMENTS Noordermeer and Wagner c o n t r i b u t e d a s l i d e o f t h e more p r o l i f i c s p e c i e s o f t h e P a l e o c e n e o f G u i a n a , which o f f e r e d a n o p p o r t u n i t y f o r comparison w i t h s p e c i e s from t h e P a l e o c e n e of Venezuela and T r i n i d a d .
B e r t e l s , by s e n d i n g
r e p r e s e n t a t i v e specimens o f a number o f A r g e n t i n e f o r m s , h e l p e d m e t o devel-
op
a b e t t e r u n d e r s t a n d i n g o f t h e f a u n a of t h a t r e g i o n . OSTRACODB DISTRIBUTION
There i s a number of g e n e r a , which h a v e been r e p o r t e d from s i n g l e a r e a s o n l y and t h e s e a r e c o n s i d e r e d h e r e t o b e more o r l e s s p r o v i n c i a l .
Some
o t h e r s a r e d i s t r i b u t e d o v e r much l a r g e r r e g i o n s and t h e r e f o r e may b e c o n s i d e r e d t o i n d i c a t e p o s s i b i l i t i e s of communication between f a u n a s o f t h o s e regions.
Genera which a r e o f i n t e r e s t i n t h e s e two ways have b e e n summar-
i z e d i n T a b l e 7 and s u g g e s t i o n s f o r d i r e c t i o n s of d i s p e r s a l a r e found i n Text-fig.
6.
I t must b e p o i n t e d o u t , t h a t t h i s m i g r a t i o n - p a t t e r n
e n t i r e l y on what h a s been o b s e r v e d up t o t h e p r e s e n t .
i s based
We have t o a c c e p t
t h e o l d e s t r e p o r t e d o c c u r r e n c e as t h e f i r s t r e a l p r e s e n c e o f a g e n u s , unl e s s t h e r e i s good r e a s o n t o d o u b t t h i s .
I n some c a s e s e x t e n s i v e s t u d i e s
have been p u b l i s h e d on the f a u n a o f t h e u n d e r l y i n g s t r a t a .
If a particular
genus i s n o t r e p o r t e d from t h e s e , i t i s assumed t h a t t h e genus d i d n o t ex-
i s t p r i o r t o i t s f i r s t reported occurrence.
As b o t h i n A r g e n t i n a and West
A f r i c a t h e Cretaceous beds a r e reasonably w e l l explored; I t h i n k w e can a c c e p t t h a t a f i r s t o c c u r r e n c e r e p o r t e d i n t h e Danian o f t h e s e r e g i o n s i s i n d e e d t h e f i r s t p r e s e n c e of t h e g e n u s .
I n n o r t h e r n S o u t h America t h e
Upper C r e t a c e o u s i s much l e s s known and h a r d l y a n y t h i n g i s known a t p r e s e n t
499 on t h e o s t r a c o d e s of t h e C r e t a c e o u s of t h e C a r i b b e a n and C e n t r a l America. O n t h e o t h e r hand t h e C r e t a c e o u s o f North America i s much more e x t e n s i v e l y
s t u d i e d and may b e u s e d i n c o n s i d e r a t i o n s o f m i g r a t i o n of o s t r a c o d e - g e n e r a . To t h e c o n t r a r y i n S o u t h A f r i c a t h e r e i s a f a i r amount o f i n f o r m a t i o n of
C r e t a c e o u s s p e c i e s , b u t n o t h i n g a t a l l i s known from t h e T e r t i a r y .
In the
f o l l o w i n g t a b l e s and l i s t s ( i n t h e a p p e n d i x ) o u r knowledge of t h e Cenozoic o s t r a c o d e s o f t h e S o u t h A t l a n t i c i s summarized by r e g i o n and s t r a t i g r a p h i c level.
Text-fig.
1:
D i s t r i b u t i o n of f o s s i l d e p o s i t s from which o s t r a c o d e s
have b e e n d e s c r i b e d i n and around t h e S o u t h A t l a n t i c , w i t h t h e d i s t r i b u t i o n of some deep-water
species.
500
T e x t - f i g s . 2-3: L o c a t i o n map of t h e w e s t e r n South A t l a n t i c . L e t t e r symbols i n d i c a t e a u t h o r o r sample: B : B e r t e l s ; D : Drooger; H : Hartmann; HB: Hulings & Baker; J : J o n e s ; K : K e i j ; N : Noordermeer & Wagner; R o : Rcssi de Garcia; S: S a n g u i n e t t i ; T . Teeter; V: van den Bold; W : Whatley & Moguilevsky.
501
HOMEY-TOGO
-
fV12-54
/
.45'
15'
30'
45
0'
u'
I
Text-figs. 4-5: L o c a t i o n map e a s t e r n A t l a n t i c Ocean. A: Apostolescu; F: Brady (Fonds d e l a Mer); G : Grekoff; M: Maddocks o r Benson & Maddocks; 0: Omatsola; R: R e p e n t ; 11-11: DSDP ( B e n s o n ) ; V : Vema; A: A t l a n t i c ; Am: A l b a t r o s ; RC: Chain ( a l l Benson) ; CH : C h a l l e n g e r ,
3
Y
e
rt 7
c(
J
m
0
J
m
0
m
P
TJ
zr
rt
&'
. I n
w m
00.
P O
e n
i3 m w
P O
rto e m
m
c.. n o
art
g$
re
rtrt
0 t-. Crn
m u
w
m
P
E
I
,
Ortra
Xerioleberir muuryoe van den Bold Uroleberir sp off. U romkormno (Lothorn)
Cylherople,on sp, off. C. d e v u Aportolercu Eucyiheruro decorota Wemgetsi
Cylherella guomreniirran den Bold Cylherella kellettae (Munsey) Cyiherella s p Cylherelloideo ~p Propontoryprll 'p Parorypr~rcornmuntr van den Beld Boirdia rnagna Alexander Bode dolirha van den Bald Boirdio soldodensir v m den Bold Bowdm aH hondurorenrtr yon den Bold Bawdia sp Krilhe sp off K peronoro Alexander Clithroryther#dea'p. off C. lombqbeenrtr (Stephenlon) Clilhrorylheridea sp Munreyollo hyolokyrljr (Munley) logoma7 kuglcr, ("on den Bald) logomo? kuglcri soldadoenru (van den Bold) Bvnronio olabarnenrrr (Hawe L Pyeatt) Bvntonia sp Lrgurnmxylherear sp off 1 logoghwoboenrtr A p o d e s c u Acanlho
Rvggierio tottvmi Reyment Nigeroloroconcho oniseguni Reyment Loxoconcho logorensis Reyment Xertoleberir kelere Reyment Schizocythere edominnini Reyment
Soudonerlo lociniora Apostolescu Soudonem trionguloto Apostolercu Soudonello nebuloro Aportolescu Hoielino rongolkomenris (Apostolercu)
Protobuntonio keiji (Reyment) Phocorhobdotur songolkomensir Aportolercu
Cythereir deltoensir Reyment Cythereir teiskotenrir (Apostolercu) Trachyleberis teirkotensis (Apostolercu) Corto dohomeyi (Aportolescu) Borrleriter obolidiegwuenrir Reyment Veenio worrienris Reyment Veenio owticortoto Reyment Bvntonio fortunano Aportolercu Buntonio ottitogoenris Aportolercu Bvntonio pukinoto Apostolercu Bvntonio bopoensis Aportolescu Bvntonio livid0 Apostolescu Protobuntonio pundoto (Reyment)
Ovocyfherideo pulchro Reyment Meherello biofrenwr Reyment Cytheropteron obo Reyment Poijenborchellino ijinenrir Reyment Togoino ottitogoenrir Aportolescu Togorno obero Aportolercu loruboello ologuni Reyment Brochycythere ogboni Reyment Dohomeyello oloto Aportolercu Leguminocythereir bopoenris (Apostolercu) Leguminocythereir logoghiroboenrir Apostolescu Leguminocythereissenegolensis Aportolescu Quodrocythere logoghiroboenrir (Apostolercu)
N
u1 0
503
Table 3 :
Range of s e l e c t e d o s t r a c o d e s p e c i e s :n t h e P a l e o g e n e of Argentina. DISCUSSION
The f o l l o w i n g g e n e r a show a p a t t e r n i n t h e i r d i s t r i b u t i o n , which makes them of s p e c i a l i n t e r e s t . 1:
B u n t o n i a i s r e p o r t e d from t h e C e n t r a l A f r i c a n C r e t a c e o u s (Aposto-
l e s c u , Keyment), and shows i n t h i s r e g i o n a c o n t i n u o u s development througho u t t h e known T e r t i a r y .
I n B r a z i l Krommelbein (1975) r e p o r t s t h i s genus i n
t h e uppermost C r e t a c e o u s .
I n Venezuela i t s e a r l i e s t known p r e s e n c e i s i n
t h e M i t o J u a n F o r m a t i o n ( D a n i a n ) , and i t c o n t i n u e s i n t o the Miocene.
In
C e n t r a l America i t s e a r l i e s t known o c c u r r e n c e i s i n t h e P a l e o g e n e ( v a n d e n B o l d , 1 9 4 6 ) , and i n North America i t h a s n o t been found u n t i l t h e e a r l y EOc e n e ( W i l c o x , Howe and G a r r e t t , 1 9 3 4 ) .
I n s o u t h e r n S o u t h America it h a s s o
f a r n o t b e e n r e p o r t e d b e f o r e t h e Miocene.
T h e r e f o r e , a d i s p e r s a l from
C e n t r a l A f r i c a t o n o r t h e r n S o u t h America and from t h e r e i n n o r t h e r l y and s o u t h e r l y d i r e c t i o n s seems i n d i c a t e d .
The genus d i s a p p e a r s i n t h e Gulf
C o a s t i n t h e O l i g o c e n e , b u t a t p r e s e n t i s found i n t h e Gulf of Mexico a t d e p t h o f 60-500 m , whereas i t s e a r l i e r forms a l l seem t o have been v e r y shallow-water s p e c i e s . 2:
S i m i l a r l y t h e g e n u s Soudanella d e v e l o p s i n t h e West A f r i c a n C r e t a -
c e o u s ( A p o s t o l e s c u , 1961) and h a s b e e n r e p o r t e d from t h e A f r i c a n Miocene ( q u e s t i o n a b l y ) ( v a n den B o l d , 1 9 6 6 h ) , and Recent ( O m a t s o l a , 1972) and from t h e Mediterranean Paleocene
-
Recent.
I n t h e C a r i b b e a n it a p p e a r s f o r t h e
f i r s t t i m e i n t h e Lower Eocene (Chiptown o f J a m a i c a ) and i n A r g e n t i n a
it h a s n o t b e e n r e p o r t e d e a r l i e r t h a n Miocene ( B e c k e r , 1 9 6 4 ) .
So f a r i t
504 i s n o t known from C e n t r a l America and North America. C e n t r a l A f r i c a , t h r o u g h S o u t h America, seems p r o b a b l e .
Here d i s p e r s a l from These two examples
of d i s p e r s a l may b e e x p l a i n e d by d o m i n a t i n g e a s t - w e s t c u r r e n t s i n t h e equat o r i a l A t l a n t i c i n C r e t a c e o u s - E a r l y T e r t i a r y t i m e s (compare Berggren and
Hollister, 1975, f i g . 16).
[ c 3
B
B
s
T a b l e 4 : Range o f s e l e c t e d o s t r a c o d e s p e c i e s i n t h e l a t e P a l e o g e n e P r o v i n c i a l i s m of s p e c i e s i n d i c a t e d . and Neogene of t h e C a r i b b e a n .
505
19 18 17 16 15 14 13 12 11 10 9
8 7 6 5 4
3 2 1 17 16
15 14 13 12 I1
10 9 8 7 6 5 4
3 2
I
P
T a b l e 5:
3:
Range o f deep-water o s t r a c o d e s i n t h e T e r t i a r y o f t h e C a r i b bean.
On t h e o t h e r h a n d , t h e g e n u s T o g o i n a h a s been i d e n t i f i e d by B e r -
tels i n t h e Cretaceous of Argentina. formerly r e f e r r e d t o B r a c h y c y t h e r e
I n n o r t h e r n S o u t h America s p e c i e s ,
(van den Bold, 1 9 5 7 ) , belonging t o t h i s
g e n u s o c c u r i n t h e P a l e o c e n e , p o s s i b l y e x t e n d i n g i n t o t h e Upper Eocene ( v a n den Bold, 1 9 6 0 ) .
I n West A f r i c a i t was o r i g i n a l l y d e s c r i b e d from t h e Upper
P a l e o c e n e and L 3 w e r Eocene ( A p o s t o l e s c u , 1 9 6 1 ) .
T h e r e f o r e d i s p e r s a l seems
t o have t a k e n p l a c e i n t h e o p p o s i t e d i r e c t i o n from t h a t of B u n t o n i a and
Soudanella. 4:
The g e n u s B a s s l e r i t e s a p p e a r s t o have t a k e n t h e same p a t h as Bun-
t o n i a but a t a l a t e r time. o f N i g e r i a (Reyment, 1 9 6 6 ) .
I t s f i r s t r e p o r t e d o c c u r r e n c e i s i n t h e Eocene I n t h e whole American a r e a it f i r s t a p p e a r s
i n t h e Miocene, p r o b a b l y e a r l i e r i n n o r t h e r n S o u t h America t h a n i n t h e
506 C a r i b b e a n and North America.
So f a r it h a s n o t b e e n r e p o r t e d from s o u t h e r n
S o u t h America.
5:
P r o t o b u n t o n i a i s r e p o r t e d from t h e C r e t a c e o u s and P a l e o c e n e of
West A f r i c a (Reyment, 1 9 6 6 ) .
On t h e o t h e r s i d e o f t h e A t l a n t i c i t h a s been
r e p o r t e d from t h e C o n i a c i a n o f B r a z i l (Krommelbein, 1975) and t h e Eocene of T r i n i d a d
(van den Bold, 1 9 6 0 ) .
* I
* * * I
**
* * I
c
Mediferronean
** Table 6:
Range of o s t r a c o d e s p e c i e s i n t h e Neogene of Gabon.
6: Procythereis
.
Here w e a r e i n t h e p e c u l i a r p o s i t i o n t h a t t h e ear-
l i e s t s p e c i e s a t t r i b u t e d t o t h i s g e n u s ( i n t h e O l i g o c e n e and Miocene o f Europe and t h e C a r i b b e a n ) a r e q u e s t i o n a b l e .
I t h a s been s u g g e s t e d t h a t
t h e y may b e l o n g t o t h e g e n u s P o k o r n y e l l a ( b u t have numerous a n t e r i o r r a d i a l p o r e c a n a l s ) and c o u l d b e l o n g t o t h e r e c e n t l y e s t a b l i s h e d g e n u s P s e u d o a u r i l a o f I s h i z a k i and Kato ( 1 9 7 6 ) .
It is c e r t a i n l y peculiar t h a t these e a r l y
o c c u r r e n c e s a r e i n ( s u b ) t r o p i c a l r e g i o n s , whereas t h e Recent o c c u r r e n c e s of P r o c y t h e r e i s a r e from t h e a n t i b o r e a l p r o v i n c e s .
I f t h e f o s s i l and
r e c e n t forms r e a l l y b e l o n g t o t h e same g e n u s , t h e n t h i s must have been est a b l i s h e d on b o t h s i d e s o f t h e A t l a n t i c p r i o r t o Miocene t i m e s and s o u t h ward m i g r a t i o n t o w a r d s c o l d e r c l i m a t e s must have o c c u r r e d s i n c e t h e P l i o cene. 7:
Neononoceratina i s a common enough genus i n t h e M e d i t e r r a n e a n and
A f r i c a n Miocene-Recent
t r o p i c a l m a r i n e environment.
So f a r i t ( N . m&i-
t e r r a n e a ) h a s o n l y shown up i n Recent d e p o s i t s on t h e o t h e r s i d e of t h e A t l a n t i c , where i t a p p e a r s r e s t r i c t e d t o v e r y s h a l l b w w a t e r ( z . g . , van Morkhoven, 1 9 7 2 ) . 8:
Another c a s e w o r t h m e n t i o n i n g h e r e i s t h e d i s t r i b u t i o n of C a l l i s -
507 tocythere.
Whereas t h i s g e n u s i s common i n t h e l a t e Cenozoic of Europe
( e s p e c i a l l y M e d i t e r r a n e a n ) and h a s been r e p o r t e d from b o t h t h e Neogene and Recent d e p o s i t s of S o u t h America, i t i s c o n s p i c u o u s l y a b s e n t i n West Africa, t h e C a r i b b e a n and t h e U n i t e d S t a t e s A t l a n t i c c o a s t , e x c e p t f o r some r e c e n t There i s o n l y one o c c u r r e n c e
i n t r o d u c t i o n s i n F l o r i d a and t h e C a r i b b e a n .
r e p o r t e d i n t h e l a t e O l i g o c e n e ( ? e a r l y Miocene) o f t h e G u l f C o a s t ( P o a g ,
1972).
U
I1
E P
? ? ?
Q I
I I
I
SOUTHERN
P M
I 1
I
S.AMERlCA
0
I
E P
P
M
0
I1
I
I'
1
I
I I
? ?
,
1
.
I
3
1
?
1
P M
AMERICA
7 3
Q
?
?
I
I
I l l
I 4 1 4 1 1
?
? 7 1 ?
I
I
1
1
?
E P
CENTRAL
11444 I I
I 1
Q CARIBBEAN NORTHERN S. AMERICA
I
I I
?
0 E P
Table 7:
1
1
1
South A t l a n t i c d i s t r i b u t i o n of s e l e c t c d o s t r a c o d e genera. 1: worldwide d i s t r i b u t i o n ; 2 : M e d i t e r r a n e a n forms; 3: A f r i c a n forms; 4 : S o u t h e r n South American forms; 5 : C a r i b b e a n forms.
Some g r o u p s w i t h American d i s t r i b u t i o n . 9.
The Genus O r i o n i n a a p p e a r s t o be a n American form.
p o r t e d o c c u r r e n c e i s i n t h e Chickasawhay of t h e Gulf C o a s t .
I t s o l d e s t re-
Its present
d i s t r i b u t i o n i s from s o u t h e r n B r a z i l t o o f f t h e A t l a n t i c c o a s t of N o r t h America, and i t a l s o o c c u r s on t h e P a c i f i c s i d e . 10:
M u n s e y e l l a i s a p p a r e n t l y o f worldwide d i s t r i b u t i o n , b u t t h e ques-
t i o n a b l y a s s i g n e d M u n s e y e l l a ? p u n c t a t a g r o u p i s o n l y known from Eocene t o Miocene i n n o r t h e r n South America ( v a n den Bold, 1 9 5 8 , 1960) and from t h e Niocene of s o u t h e r n B r a z i l , which may p o i n t t o a n e x t e n s i o n o f t h e C a r i b bean z o o g e o g r a p h i c p r o v i n c e t o t h a t r e g i o n i n t h e Miocene.
11:
The genus C a t i v e l l a h a s a l s o n o t c r o s s e d t h e A t l a n t i c .
Omatsola
(1972) r e p o r t s C a t i v e l l a and N e o c a u d i t e s from Recent d e p o s i t s of t h e N i g e r delta.
However, t h i s s p e c i e s of C a t i v e l l a ( C . i x e m o j a i l
and i s v e r y s i m i l a r t o s p e c i e s o f C h r y s o c y t h e r e . a s s i g n m e n t o f s p e c i e s of P u r i a n a from West A f r i c a .
seems d i f f e r e n t
I a l s o doubt t h e c o r r e c t
The e a r l i e s t o c c u r r e n c e
of C a t i v e l l a i s i n t h e l a t e Eocene of T r i n i d a d ( v a n den B o l d , 1 9 6 0 ) .
In
t h e C a r i b b e a n i t a p p e a r s i n l a t e r s t a g e s o f t h e Miocene a s one p r o g r e s s e s f a r t h e r n o r t h ( v a n den B o l d , 1 9 7 4 a ) , and i n North America i t o n l y makes i t s a p p e a r a n c e n e a r t h e P l i o c e n e boundary.
I t h a s been r e p o r t e d from t h e Qua-
t e r n a r y o f s o u t h e r n B r a z i l and t h e A n t a r c t i c .
Here a n o r t h w a r d and s o u t h -
ward d i s p e r s a l away from t r o p i c a l America seems i n d i c a t e d .
Recent d i s t r i -
b u t i o n of t h e genus i s s c a t t e r e d ; whereas i t i s common between 1 0 and 2 0 m d e p t h i n t h e Gulf o f P a r i a , i t o c c u r s s c a t t e r e d i n t h e Gulf of Mexico between 30 and 1 0 0 m . Brackish-water genera. 12:
P e r i s s o c y t h e r i d e a i s a genus t h a t w a s supposed t o b e r e s t r i c t e d t o
t r o p i c a l and s u b t r o p i c a l America till Benson and Maddocks ( 1 9 6 4 ) found it i n t h e Knysna e s t u a r y o f S o u t h A f r i c a .
According t o Hartmann ( 1 9 7 5 ) , how-
e v e r , t h i s s p e c i e s a p p e a r s t o b e a somewhat abnormal r e p r e s e n t a t i v e .
Di-
v e r s i f i c a t i o n of t h e g e n u s a p p e a r s s t r o n g e s t i n t h e C a r i b b e a n . 13:
The d i s t r i b u t i o n of C y p r i d e i s h a s been d i s c u s s e d by s e v e r a l a u t h -
o r s b o t h on t h e A t l a n t i c and A f r i c a n s i d e of t h e A t l a n t i c o c e a n . e r n A t l a n t i c s p e c i e s appear less d i v e r s i f i e d .
The e a s t -
Except f o r C y p r i d e i s sirnilis
none o f t h e s p e c i e s a p p e a r s t o o c c u r o n b o t h s i d e s , a l t h o u g h some r a n g e o v e r a c o n s i d e r a b l e nurnber o f l a t i t u d e s , p o s s i b l y c o n n e c t e d w i t h d i s p e r s a l by m i g r a t i n g w a t e r f o w l .
14:
S u l c o s t o c y t h e r e from S o u t h A f r i c a a p p e a r s t o r e p l a c e C y p r i d e i s
l o c a l l y (Hartmann, 1 9 7 5 ) i n b r a c k i s h - w a t e r e n v i r o n m e n t ; h s o l i however, m e n t i o n s it from t h e I v o r y C o a s t .
(1975),
509 Mediterranean forms i n A f r i c a . 15:
Among t h e s e a r e Chrysocythere, Ruggieria, Carinocythereis and pos-
sibly Mutilus.
They do n o t e x t e n d a c r o s s t h e A t l a n t i c a l t h o u g h a s p e c i e s ,
t e n t a t i v e l y i d e n t i f i e d as Huggieria h a s been r e p o r t e d from Recent Caribbean
waters (van den Bold, 1966c, Teeter, 1 9 7 5 ) .
T h i s s p e c i e s , however, i s
r a t h e r d i f f e r e n t from t h e Mediterranean forms (much s t r o n g e r r e t i c u i a t i o n ) .
Carinocythereis may be r e l a t e d t o C a t i v e l l a through common a n c e s t o r s . Paijenborchellina (L. Eocene and Recent of N i g e r i a ) h a s a l s o n o t been found s o f a r on t h i s s i d e of t h e A t l a n t i c . Endemic o r p r o v i n c i a l forms. 16:
Africa:
t h e r e , Isobuntonia
Reymentia, Dahomeyella I
Evisceratocythere
I
, Sulcostocythere,
Ovocytheridea
I
a l s o Isohabrocy-
Nigeroloxoconcha.
Some of t h e s e have been r e p o r t e d from North A f r i c a ( L i b y a , A l g e r i a , E g y p t ) . 17:
Southern South America:
Wichmannella , Huantraiconella, Protocosta,
Neoveenia have n o t been r e p o r t e d from o t h e r a r e a s .
However, forms resem-
b l i n g Protocosta o c c u r i n n o r t h e r n South America and A f r i c a . and Pericythere a l s o o c c u r i n t h e Caribbean a r e a .
Mesocythere
A new s p e c i e s of Austra-
l i c y t h e r e h a s been d e s c r i b e d from t h e Miocene of s o u t h e r n B r a z i l . indicated i n text-fig.
Although
6 , it i s s t i l l t o o e a r l y t o draw any c o n c l u s i o n s .
S i m i l a r i s t h e case of Patagonocythere. 18:
Also o f i n t e r e s t is the r e s t r i c t i o n of t h e genus Ruttenella t o t h e
Eocene o f t h e s o u t h e r n Caribbean (van den Bold, 1 9 6 0 ) . 19:
P u z z l i n g i s t h e d i s t r i b u t i o n of Cobanocythere ( o r i g i n a l l y d e s c r i b -
ed from t h e P a c i f i c c o a s t o f C e n t r a l America, Hartmann, 1959) i n West A f r i ca (Hartmann, 1 9 7 5 ) , b u t a l s o mentioned by Reys from t h e Mediterranean and by Shornikov (1975) from J a p a n , w h i l e S i d d i q u i (1971) q u e s t i o n a b l y a s s i g n e d a s p e c i e s from t h e e a r l y T e r t i a r y o f P a k i s t a n t o t h i s genus. Deep-water Ostracoda. 20:
D i s t r i b u t i o n o f deep-water o s t r a c o a e s i n t h e Caribbean a r e a ( T a b l e
5) a f f o r d s some comparison w i t h t h a t i n deep-sea
cores.
Unfortunately the
s t u d y o f t h e l a t t e r h a s s o f a r been c o n f i n e d t o t h e g e n e r a Bradleya, Agren-
ocythere, Poseidonamicus and Abyssocythere (Benson, 1971, 1972)
.
In the
Caribbean, Agrenocythere i s known from Eocene t o l a t e Miocene, and Recent specimens have been found i n deep-water c o r e s i n t h e Gulf of Mexico.
In
t h e DSDP c o r e s Agrenocythere hazelae i s known from Oligocene t o e a r l y Miocene and from P l e i s t o c e n e from around 3OoS l a t i t u d e ( t e x t - f i g .
1).
Recent
specimens have been found i n t h e North A t l a n t i c , t h e Gulf o f Mexico and t h e P a c i f i c (Benson, 1 9 7 2 ) .
I t s p r e d e c e s s o r A . antiquata i s found i n t h e EO-
510 c e n e o f t h e C a r i b b e a n , t h e e a s t e r n N o r t h A t l a n t i c and I t a l y . A b y s s o c y t h e r e t r i n i d a d e n s i s i s s o f a r o n l y known i n t h e C a r i b b e a n , b u t a r e l a t e d species ( A . a t l a n t i c a ) occurs i n Quaternary cores i n t h e Central
1).
Atlantic (Text-fig.
The m a j o r i t y of s p e c i m e n s grouped under B r a d l e y a e x g r . d i c t y o n ( T a b l e
5) r e a l l y b e l o n g s t o t h a t s p e c i e s , b u t t h e y are accompanied o v e r t h e i r ent i r e s t r a t i g r a p h i c r a n g e by smaller s p e c i m e n s , which m i g h t r e p r e s e n t a F o r d i s t r i b u t i o n o f t h e s e g e n e r a t h e r e a d e r i s re-
shallow-water v a r i a n t .
f e r r e d t o Benson ( 1 9 7 1 , 1 9 7 2 )
.
So f a r n o t h i n g h a s been p u b l i s h e d on Recent r e p r e s e n t a t i v e s of A b y s -
The l a t t e r h a s been found i n water i n e x c e s s o f
s o c y p r i s and C a r d o b a i r d i a .
?GO m d e p t h i n t h e Gulf of Mexico.
Although d e e p - w a t e r s p e c i e s of K r i t h e ,
C y t h e r e l l a , A r g i l l o e c i a , B y t h o c y p r i s and B a i r d i a a r e w e l l known, b o t h from Recent and fossil m a t e r i a l , a s t u d y o f t h e i r d i s t r i b u t i o n would r e q u i r e a more d e t a i l e d i n v e s t i g a t i o n t h a n c a n be u n d e r t a k e n h e r e .
I t may b e men-
t i o n e d , however, t h a t two s p e c i e s of B a i r d i a , common t o deep-water d e p o s i t s of t h e T e r t i a r y of t h e C a r i b b e a n , a r e r e p r e s e n t e d by i d e n t i c a l o r a t l e a s t s t r o n g l y r e l a t e d forms i n t h e Gulf o f Mexico, t y p i c a l l y a t d e p t h s between I t i s i n t e r e s t i n g t o n o t e t h e s i m i l a r i t y o f Maddock's
9 0 0 and 1 2 0 0 m e t e r s .
NORTH AMERICA Borileritei (M)
Bunionia (Pi
4
I
Barder'iter ( M )
T CENTRAL
I
Bvntonia
Coiirello ( M I
(M)
Ruggieiio
A
A
(P)
MEDITERRANEAN
Mulilvr ( M I
A
Chryrocythere (M)
A(?) U P )
4
+
+ 7 Barrleritei (M) 4 Bvntanto (C)
CARIBBEAN A N D NORTHERN
*
*
Soudonello (C) Probbuntonio ( C )
Protobuntonia (C) 4
c
Cofirella (E)
SOUTHERN SOUTH AMERICA
Togolno
(P)
4
I
B ~ (MI ~ ~ . A ~ Soudonella ( M I Togo,na (C)
Cotirello (R)
I
Avrtrolicylhere ( M )
ANTARCTIC Ca'ivella
WEST AFRICA
Buntonio ( C )
Soudonella (E)
SOUTU AMERICA
Barrler~ler(E)
lR)
1
Aurtrolqthere ( R )
T e x t - f i g . 6 : A p p a r e n t d i r e c t i o n o f d i s p e r s a l o f some s e l e c t e d O s t r a (R) Recent, code g e n e r a i n A f r i c a and t h e Americas d u r i n g t h e Cenozoic. resp. Plio-Pleistocene; ( M ) MloCene; (E) Eocene; ( P ) P a l e o c e n e .
511 s p e c i e s B a i r d o p p i l a t a s p . 3 , from t h e Mozambique c h a n n e l (1860
-
2980 m ) .
I n t h e C a r i b b e a n P l e i s t o c e n e ( A t l a n t i c 71A-4) Ambocythere c a u d a t a , N o r t h A t l a n t i c ) , T r a c h y l e b e r i s bermudezi a f f . c r e b r i p u s t u l o s a
(Recent,
(identical to
Recent form from Gulf o f M e x i c o ) , B y t h o c e r a t i n a s c a b r a and Z a b y t h o c y p r i s h e t e r o d o x a were found. Benson (1975) h a s a r g u e d on t h e b a s i s o f t h e deep-sea o s t r a c o d e s , t h a t i n l a t e E o c e n e - e a r l y O l i g o c e n e t i m e s a change i n f a u n a l c o m p o s i t i o n t o o k p l a c e , which may b e c o r r e l a t e d w i t h l o w e r i n g of t e m p e r a t u r e c h a n g e s ) i n t h e deep-sea
(and o t h e r
watermass ( o r i g i n o f t h e p s y c h r o s p h e r e ) .
As t h e r e
i s a n o t a b l e l a c k o f d a t a on s h e l f f a u n a s of t h a t t i m e , no e f f e c t on shallow-water faunas can be observed.
I n T r i n i d a d , t h e o n l y p l a c e where
u p p e r P a l e o g e n e d e e p - w a t e r f a u n a s have been s t u d i e d , no s i g n i f i c a n t change c a n b e d e t e c t e d , a l t h o u g h minor changes
-
m a i n l y on t h e s p e c i e s l e v e l
-
t a k e p l a c e , b e g i n n i n g i n t h e l a t e Eocene ( T a b l e 5 ) . Conclusion Through t h e P a l e o c e n e and Eocene w e see a f a i r l y c l o s e r e l a t i o n s h i p between S o u t h American and West C e n t r a l A f r i c a n o s t r a c o d e f a u n a s .
This connection
c o u l d have been t h r o u g h f a i r l y s h a l l o w w a t e r , b u t more l i k e l y was a f f e c t e d by d i s p e r s a l o f s p e c i e s a t t a c h e d t o p l a n t m a t e r i a l d r i f t i n g m o s t l y i n e a s t -
w e s t direction.
Of i n t e r e s t h e r e i s t h e a l m o s t c o m p l e t e l a c k of r e l a t i o n -
s h i p between E a s t and West A f r i c a n f a u n a s ( D i n g l e , 1 9 7 6 ) .
A f t e r t h e Oligo-
c e n e , most d i s p e r s a l t o o k p l a c e a l o n g t h e A t l a n t i c c o a s t s i n n o r t h and south directions.
512 BIBLIOGRAPHY I n o r d e r t o keep t h e b i b l i o g r a p h y w i t h i n r e a s o n a b l e dimensions, I have o m i t t e d a l l r e f e r e n c e s t o R e c e n t f r e s h - w a t e r o s t r a c o d e s and R e c e n t Myodoc o p i d a , and p l a c e d the o n l y two on T e r t i a r y f r e s h - w a t e r o s t r a c o d e s i n brackets i n t h e b i b l i o g r a p h i c index.
T h i s i n d e x shows t h a t , w h e r e a s t h e r e
a r e e i g h t p u b l i c a t i o n s on t h e P a l e o g e n e o f C e n t r a l A f r i c a , t h e r e a r e o n l y two on t h e Neogene and f o u r on Recent o s t r a c o d e s .
F o r s o u t h e r n S o u t h Amer-
i c a t h e s e f i g u r e s a r e 9 , 7 and 4 ; i n C e n t r a l S o u t h America 0 , 0 , 8 ( a p a p e r by S a n g u i n e t t i on t h e Miocene o f t h e P e l o t a s b a s i n , s o u t h e r n B r a z i l , s h o u l d be i n p r e s s ) ; i n t h e C a r i b b e a n and n o r t h e r n S o u t h America t h e numbers a r e 1 3 , 31 and 11 r e s p e c t i v e l y .
In t h e A t l a n t i c basin i t s e l f 4 papers deal
w i t h P a l e o g e n e , Neogene and Recent o s t r a c o d e s , and s e v e n d e a l o n l y w i t h Rec e n t ones.
F i n a l l y , from t h e A n t a r c t i c p a r t o f t h e A t l a n t i c 4 p a p e r s d e s -
c r i b e R e c e n t o s t r a c o d e s , none a r e on T e r t i a r y f o r m s . dex t h e abbreviations P :
Palecgene, N:
I n the following in-
Neogene and R:
Recent have been
used. Bibliographic index AFRICA
Central:
A p o s t o l e s c u , 1961 ( P ) ; B o l d , 1966 (N), 1968c (N) (Grek o f f , 1958) ( P , N ) ; Keen, 1975 ( N , R ) ; Kogbe e t a l . , 1975 (P); M a s o l i , 1975 ( R ) ; O m a t s o l a , 1970a ( R ) , 1970b ( R ) , 1970c ( R ) , 1 9 7 1 ( R ) , 1972 ( R ) ; Reyment, 1959 ( P ) , 1960 (P), 1963 ( P ) , 1964 ( P I , 1965 ( P ) , 1966 ( P I ; Reyment and Reynent, 1959 ( P ) .
South :
Benson and Naddocks, 1964 ( R ) ; Hartmann and HartmannS c h r g d e r , 1975 ( R ) ; K l i e , 1940 ( R ) , Miiller, 1908 ( R ) .
South :
B e c k e r , 1964 ( R ) ; B e r t e l s , 1966a ( P ) , 196813 ( P ) , 1969a ( P ) , 1969b ( P ) , 1969c ( P ) , 1973 ( P ) , 1975 ( P , ? j ) ; H a l umian, 1970 (N); Ramirez, 1967 ( R ) ; Ramirez and Moguil e v s k y , 1 9 7 1 (R); R o s s i d e G a r c i a , 1966a (PI, 1966b (TJ), 1967a ( P ) , 196713 ( N ) , 1967c (N), 1 9 7 1 (N), 1972 ( 7 ) ; Whatley and C h o l i c h , 1974 ( R ) , Whatley and Moguil e v s k y , 1975 ( R ) .
SOUTH ANERICA
Central :
Brady, 1880 ( R ) ; Hartmann, 1955a ( R ) , 1955b ( R ) , 1956 ( R ) ,1957 ( R ) , 1959 ( R ) ; ( J o n e s , 1968) (N); Macedo, 1965 (R); P i n t o and O r n e l l a s , 1965 ( R ) ,1970 ( R ) .
CARIBBEAN A N D NORTHERN SOUTH A3lERICA Baker and H u l i n g s , 1966 ( R ) ; B o l d , 1946 ( P , K ) , 1950a ( P , N ) , 1950b ( P , N ) , 1957a ( P ) , 195723 ( N ) , 1958a ( R ) , 1958b ( N ) , 1 9 5 8 ~( N ) , 1960a ( P ) , 1960b ( P I t 1 9 6 1 ( P , ? : ) , 1963a (N) , 196333 ( R ) , 1963c ( P , N ) , 1963d (N), 1965a ( R ) , 1965b (P,N), 1966a (N), 1966b (N) , 1966c ( R ) ,
1966d (N), 1966e ( X ) , 1966f (N), 1966g ( N ) , 1 9 6 6 i (P, 1967b (N) t 1968a ( X ) r 196813 (N) t 196% ( N ) , 1969b ( P , N ) , 1970a ( K ) , 1970b (N), 1971a ( P , N ) ,
K ) , 1967a ( N ) ,
513 1971b ( N ) , 1 9 7 1 ~( N ) , 1972a ( P , N ) , 197213 ( N ) , 1 9 7 2 ~ (N), 1973 ( N ) , 1971c ( N ) , 1972a ( P , N ) , 197333 ( N ) , 1972c ( N ) , 1973 ( P , N ) , 1974a (N), 1974b ( N ) , 1974c ( N ) , 1975a (N) ; Brady, 1870 (R); Bronnimann and R i g a s s i , 1963 ( P ) , D r o o g e r , 1960 ( P ) , Drooger and K a a s s c h i e t e r , 1958 ( R ) 1 K e i j , 1954 (R); K l i e , 1939a ( R ) , 193923 (R) ; Lubimova and S a n c h e z , 1974 ( P , N ) ; Noordermeer a n d Wagn e r , 1969 ( P ) ; PokornG, 1968 (R); T e e t e r , 1975 ( R )
.
ATLANTIC
Benson, 1 9 6 9 , 1 9 7 1 , 1972; Benson and S y l v e s t e r - B r a d l e y , 19711 B r a d y , 1880 ( R ) , 1887 ( R ) , 1 9 1 1 ( R ) ; Hartmann, 1959a ( R ) ; Maddocks, 1969 ( R ) ; P u r i , 1967 (R), 1 9 7 1 (R)
ANT ARC T I C
.
B r a d y , 1907 (R), Neale, 1967 ( R ) , S c o t t , 1899 (R), 1912 (R). Bibliography
A p o s t o l e s c u , V . , 1961. C o n t r i b u t i o n 'a 1' g t u d e p a l g o n t o l o g i q u e ( o s t r a c o d e s ) e t s t r a t i g r a p h i q u e d e s b a s s i n s crgtacgs e t t e r t i a i r e s d e l ' A f r i q u e . Rev. I n s t . fran:. p g t r o l e e t a n n . combust. l i q . , v . 1 6 , no. 7-8, p . 779-867. B a k e r , J. H . and H u l i n g s , N . C . , 1 9 6 6 . R e c e n t Marine O s t r a c o d Assemblages o f P u e r t o R i c o . P u b l . I n s t . Mar. S c i . , T e x a s , v . 11, p . 108-125. B e c k e r , D . , 1 9 6 4 . M i c r o p a l e o n t o l o g l a d e l s u p e r p a t a g o n i e n s e de l a s l o c a l i d a d e s L a s Cuevas y Monte E n t r a n c e . Ameghiana, v. 3 , n o . 1 0 , p . 319340. Benson, R. H . , 1969. P r e l i m i n a r y R e p o r t on t h e s t u d y o f A b y s s a l O s t r a c o d a . I n : K e a l e , J . W . Tax. Morph. and E c o l . Rec. O s t r . , H u l l , O l i v e r & Boyd E d i n b u r g h , p . 475-478. 1 1971. A N e w Cenozoic Deep-sea Genus A b y s s o c y t h e r e ( C r u s t a c e a , Ost r a c o d a : T r a c h y l e b e r i d i d a e ) , w i t h D e s c r i p t i o n s o f f i v e new s p e c i e s . S m i t h s o n i a n C o n t r . P a l e o b i o l . , n o . 7 , p . 1-20. 1972. The B r a d l e y a Problem, w i t h d e s c r i p t i o n s o f two new Psychros p h e r i c O s t r a c o d e G e n e r a , A g r e n o c y t h e r e and P o s e i d o n a m i c u s ( O s t r a c o d a : C r u s t a c e a ) . S m i t h s o n i a n C o n t r . P a l e o b i o l . no. 1 2 , p . 1-138. 1975. The o r i g i n o f t h e p s y c h r o s p h e r e a s r e c o r d e d i n changes o f deep-sea o s t r a c o d e a s s e m b l a g e s . L e t h a i a , v o l . 8 , p . 69-83. Benson, R. H . and Maddocks, R. F . , 1964. R e c e n t O s t r a c o d e s o f Knysna e s t u a r y , Cape P r o v i n c e , Union o f S o u t h A f r i c a . Univ. Kansas P a l . C o n t r . , A r t . 5 , p . 1-39. Benson, R. H . and S y l v e s t e r - B r a d l e y , P . C . , 1971. Deep-sea o s t r a c o d e s and t h e t r a n s f o r m a t i o n o f o c e a n t o sea i n t h e T e t h y s . Bull. C e n t r e Rech. Pau-SNPA, v. 5 s u p p l . , p. 63-91. B e r g g r e n , W . A. and H o l l i s t e r , C . D . , 1974. P a l e o g e o g r a p h y , p a l e o b i o g e o graphy and h i s t o r y o f t h e c i r c u l a t i o n i n t h e A t l a n t i c Ocean. In: W . W . Hay ( e d i t . ) S t u d i e s i n paleo-oceanography. SOC. Econ. P a l e o n t . M i n e r a l . s p . p u b l . n o , 20, p . 126-186. B e r t e l s , A , , 1968a. H u a n t r a i c o n e l l a n . g e n . ( O s t r a c o d a , B u n t o n i i n a e ) d e l T e r t i a r i o i n f e r i o r ( D a n i a n o ) d e A r g e n t i n a . Ameghiana, V . 5 , no. 8 , p . 252-6. 1968b. M i c r o p a l e o n t o l o g T a y e s t r a t i g r a f i a d e l l i m i t e c r e t e t i c i c o t e r t i a r i o e n H u a n t r a i - c o ( P r o v i n c i a d e Neugugn). I b i d . , v. 5 , no. 8 , p. 279-295. , 1969a. E s t r a t i g r a f T a d e l l i m i t e C r e t i c i c o - T e r c i a r i o e n P a t a g o n i a septentrional. Rev. Asoc. G e o l . A r g e n t . , v. 2 4 , no. 1, p . 41-54.
-,
-,
-,
-
514 B e r t e l s , A . , 1969b. " R n c a l e b e r i d i n a e " , nueva s u b f a m i l i a ( " O s t r a c o d a , C r u s t a c e a " ) d e l l i m i t e C r e t & i c o - T e r t i a r i o de P a t a g o n i a s e p t e n t r i o n a l . Ameghiniana, v. 6 , no. 2 , p . 146-171. , 1 9 6 9 ~ . Micropaleontologia y e s t r a t i g r a f i a d e l l i m i t e CretgcicoT e r t i a r i o en Huantrai-Co ( P r o v i n c i a d e Neugugn). Ameghiniana, v . 6 , no. 4 , p. 253-280. 1973. O s t r a c o d e s o f t h e t y p e l o c a l i t y o f t h e Lower T e r t i a r y (lower Danian) Rocanian S t a g e and Roca Formation o f A r g e n t i n a . Micropal., v . 1 9 , no. 3 , p. 308-340. 1975. O s t r a c o d e e c o l o g y d u r i n g t h e u p p e r C r e t a c e o u s and Cenozoic i n A r g e n t i n a . B u l l . Amer. P a l e o n t . v . 65, no. 282, p . 317-341. Bold, W. A. van d e n , 1946. C o n t r i b u t i o n t o t h e s t u d y o f O s t r a c o d a w i t h s p e c i a l r e f e r e n c e t o t h e T e r t i a r y and C r e t a c e o u s m i c r o f a u n a o f t h e C a r i b b e a n Region. D i s s . U t r e c h t Univ., DeBussy, Amsterdam, 167 pp. 1950a. Miocene O s t r a c o d a from V e n e z u e l a . J o u r . P a l e o n t . , V . 24, no. 1, p . 76-88. , 1950b. A c h e c k l i s t o f Cuban O s t r a c o d a . I b i d . V . 24, no. 1, p. 107109. , 1957a. O s t r a c o d a from t h e P a l e o c e n e o f T r i n i d a d . M i c r o p a l . , V . 3 , no. 1, p . 1-18. , 1957b. Oligo-Miocene O s t r a c o d a from s o u t h e r n T r i n i d a d . I b i d . v. 3 , no. 3 , p. 231-254. 1958a. D i s t r i b u t i o n of f r e s h w a t e r o s t r a c o d e s i n T r i n i d a d . Ibid. v. 4 , no. 1, p . 71-74. , 1958b. O s t r a c o d a o f t h e B r a s s o F o r m a t i o n of T r i n i d a d . I b i d . v . 4 , no. 4 , p. 391-418. , 1 9 5 8 ~ . Ambocythere, a new g e n u s o f o s t r a c o d a . Ann. Mag. N a t . H i s t . , s . 1 2 , v . 1 0 , p . 801-813. , 1960a. Eocene and O l i g o c e n e O s t r a c o d a o f T r i n i d a d . M i c r o p a l . , V . 6, no. 2 , D . 145-196. Determination of o s t r a c o d e s , i n : Pessaqno, E . A , , S t r a t i graphy and m i c r o p a l e o n t o l o g y o f t h e C r e t a c e o u s and Lower T e r t i a r y o f P u e r t o Rico. M i c r o p a l . , v . 6 , no. 1, p . 87-110, O s t r a c o d a , p. 9 3 ; i n : M a t t s o n , P. H . , Geology o f t h e Mayaguez a r e a , P u e r t o Rico. B u l l . Geol. SOC. Am. V . 71, p. 310-362, O s t r a c o d a , p. 347. 1961. Some new O s t r a c o d a of t h e C a r i b b e a n T e r t i a r y . P r o c . Kon. PIederl. Akad. W e t e n s c h s e r . B . , V . 6 4 , no. 5 , p . 627-638. , 1963a. The o s t r a c o d e genus O r i o n i n a and i t s s p e c i e s . J o u r . P a l e o n t . , V . 3 7 , no. 1, p. 33-50. 1963b. Anomalous h i n g e s t r u c t u r e i n a new s p e c i e s o f C y t h e r e l l o i d e a M i c r o p a l e o n t o l o g y , V . 9 , no. 1, p . 75-78. , 1963c. O s t r a c o d s and t h e T e r t i a r y s t r a t i g r a p h y o f Guatemala. B u l l . Amer. Assoc. P e t r . G e o l . , v o l . 4 7 , no. 4 , p . 696-698. Upper Miocene and P l i o c e n e O s t r a c o d a of T r i n i d a d . Micropaleontology, v . 9 , no. 4 , p . 361-424. , 1965a. F s e u d o c e r a t i n a , a new genus o f O s t r a c o d a from t h e C a r i b b e a n . Kon. N e d e r l . Akad. Wetensch., P r o c . , s e r . B , v . 6 8 , no. 3 , 160-164. , 1965b. Middle T e r t i a r y O s t r a c o d a from n o r t h w e s t e r n P u e r t o Rico. M i c r o p a l e o n t o l o g y , v . 11, no. 4 , p . 381-414. 1966a. O s t r a c o d a from t h e Poz6n s e c t i o n , F a l c g n , Venezuela. Jour. P a l e o n t . , v. 4 0 , no. 1, p . 177-185. 196633. N e w s p e c i e s o f t h e o s t r a c o d e - g e n u s Ambocythere. Ann. Mag. -' Nat. H i s t . , s e r . 1 3 , v . 8 , no. 1, p . 1-18. 1 9 6 6 ~ . O s t r a c o d a from Colon H a r b o u r , Panama. C a r i b . J o u r . S c i . , v. 6 , no. 1 / 2 , p . 43-53. 1966d. O s t r a c o d e zones i n C a r i b b e a n Miocene. Bull. Amer. ASSOC, -1 P e t r . Geol. v. 50, no. 5 , p . 1029-1031.
-
-,
-, ,
-, -
-
-, -, -, -,
-,
515 B o l d , W . A. van d e n , 1966e. Miocene and P l i o c e n e O s t r a c o d a from N o r t h e a s t e r n Venezuela. Verh. K O n . N e d e r l . Akad. Wetensch., s e r . 1, V. 23, no. 3 , p . 1-43. , 1 9 6 6 f . Upper Miocene O s t r a c o d a from t h e T u b a r s F o r m a t i o n ( n o r t h e r n C o l o m b i a ) . M i c r o p a l e o n t o l o g y , v. 1 2 , no. 3 , p . 360-364. 1966g. O s t r a c o d a from t h e A n t i g u a F o r m a t i o n ( O l i g o c e n e , Lesser Ant i l l e s ) . J o u r . P a l e o n t . , V . 4 0 , no. 5 , p . 1233-1236. , 1966h. L e s o s t r a c o d e s du N6ogsne du Gabon. Rev. I n s t . f r a n F . du P&. V . 2 1 , no. 2 , p. 155-176. 1966i. R g p a r t i t i o n de c e r t a i n s o s t r a c o d e s dans l e T e r t i a i r e des Cara'ibes. 3 r d sess. (Bern) Comm. M e d i t . Neogene S t r a t i g r a p h y , p . 134139. 1967a. Miocene O s t r a c o d a from C o s t a R i c a . M i c r o p a l e o n t o l o g y , V . 13, no. 1. p . 75-86. , 1967b. O s t r a c o d a of t h e G a t h F o r m a t i o n , Panama. I b i d . v. 1 3 , no. 3 , p. 306-318. 1968a. D i s t r i b u t i o n of T r a c h y l e b e r i d i n a e ( O s t r a c o d a ) i n t h e Neogene of t h e C a r i b b e a n . 4 t h sess. Comm. Medit. Neog. S t r a t . , P r o c . p . 55-66 1968b. O s t r a c o d a of t h e Yague Group (Neogene) of t h e n o r t h e r n Domin i c a n R e p u b l i c . B u l l . Amer. P a l e o n t . , v . 54, no. 239, 106 pp. , 1 9 6 8 ~ . O s t r a c o d e s du NGogSne du Gabon e t d e 1 ' I t a l i e . Rev. I n s t . f r a y . P g t r . V. 23, no. 1 0 , p . 1327-1328. 1969a. Neogene O s t r a c o d e s from S o u t h e r n P u e r t o Rico. C a r i b . J o u r . S c i . , V . 9 , no. 3/4, p. 117-125. 1969b. Messinella, a new g e n u s of O s t r a c o d a i n t h e C a r i b b e a n Cenoz o i c . M i c r o p a l e o n t o l o g y , V . 1 5 , no. 4 , p. 397-400. , 1970a. The genus C o s t a ( O s t r a c o d a ) i n t h e Upper Cenozoic of t h e C a r i b b e a n Region. I b i d . V . 1 6 , no. 1, p. 61-75. , 1970b. O s t r a c o d a o f t h c Lower and Middle Miocene of S t . C r o i x , S t . M a r t i n and A n g u i l l a . C a r i b . J o u r . S c i . , v. 1 0 , no. 1 / 2 , p . 35-52. 1971a. D i s t r i b u t i o n of o s t r a c o d e s i n t h e Oligomiocene of t h e Northe r n C a r i b b e a n . T r a n s . 5 t h C a r i b . Geol. C o n f . , B u l l . no. 5 , p . 123-128. , 1971b. O s t r a c o d a of t h e C o a s t a l Group of F o r m a t i o n s of J a m a i c a . T r a n s . Gulf C o a s t ASSOC. Geol. SOC. v. 2 1 , p . 325-348. , 1 9 7 1 ~ . O s t r a c o d e a s s o c i a t i o n s , s a l i n i t y and d e p t h o f d e p o s i t i o n i n t h e Neogene of t h e C a r i b b e a n Region. B u l l . C e n t r e Rech. Pau-SNPA, 5 SUP. p . 449-460. , 1972a. O s t r s c o d o s d e ; Post-Eoceno B l V e n e z u e l a y r e g i o n e s v e c i n a s . Congr. Geol. Venezolano, V . 2 , mem. 4 , Sp. p u b l . 5 , p . 999-1063. , 1972b. C o n t r i b u t i o n o f O s t r a c o d a t o t h e c o r r e l a t i o n o f Neogene f o r m a t i o n s of t h e C a r i b b e a n Region. Mem-Trans. 6 t h C a r i b . Geol. C o n f . , p . 485-490. , 1972c. O s t r a c o d a of t h e L a Boca F o r m a t i o n , Panama C a n a l zone. M i c r o p a l e o n t o l o g y , V . 1 8 , no. 4 , p . 410-442. , 1973. D i s t r i b u t i o n of O s t r a c o d a i n t h e O l i g o c e n e and Lower and Middie Miocene of Cuba. C a r i b . J o u r . S c i . , v . 1 3 , no. 3/4, p. 145-159. , 1974a. O s t r a c o d e a s s o c i a t i o n s i n t h e C a r i b b e a n Neogene. Verh. N a t u r f . Gesell. Basel, V. 8 4 , no. 1, p. 214-221. 197423. O r n a t e B a i r d i i d a e i n t h e C a r i b b e a n . G e o s c i . Man, v . 6 , p.29 -40. , 1974c. Taxonomic s t a t u s of Cardobairdia van den B o l d , 1960, and Abyssocypris n. g e n . , two g e n e r a of deep-water o s t r a c o d e s from t h e Caribbean T e r t i a r y . I b i d . , v. 6 , p. 65-79. , 1975a. Neogene b i o s t r a t i g r a p h y ( O s t r a c o d a ) o f s o u t h e r n H i s p a n o l a . B u l l . Amer. P a l e o n t . , V . 6 6 , no. 286, p . 549-625. 1975b. O s t r a c o d e s from t h e l a t e Neogene of Cuba. I bi d., v o l . 68, no. 289, p. 121-167.
-,
-,
-, -,
-,
-
, -,
-
-,
-
-, -,
516 Bold, W. A. van d e n , 1975c. D i s t r i b u t i o n o f t h e Radimella confraqosa Group ( O s t r a c o d a , H e m i c y t h e r i n a e ) i n t h e l a t e Neogene o f t h e C a r i b b e a n . J o u r . P a l e o n t . , 49, no. 4 , p. 692-701. B r a d y , G . S . , 1870. D e s c r i p t i o n o f O s t r a c o d a , I n : D e F o l i n e t P g r i e r : L e s Fonds d e l a Mer, v . 1, p . 177-256. 1880. Report on t h e O s t r a c o d a d r e d g e d by HMS C h a l l e n g e r d u r i n g t h e y e a r s 1873-1876. R e p o r t on t h e s c i e n t i f i c r e s u l t s o f t h e voyage o f HMS C h a l l e n g e r , Zool. V . 1, p t . 3 , p . 1-184. 1887. D e s c r i p t i o n of O s t r a c o d a from t h e e x p l o r a t i o n s of t h e "Trav a i l l e u r " and t h e "Talisman". I n : De F o l i n e t P g r i e r : Les Fonds d e l a Mer, v . 4 , p. 164-226. 1907. C r u s t a c e a , P t . 5 : Ostracoda. National A n t a r c t i c Expedition. N a t . H i s t . , 1901-1904, V . 3 , p . 1-9. 1911. Kotes on m a r i n e O s t r a c o d a from Madeira. Proc. Zool. SOC., London, 1911, p . 595-601. Bronnimann, P . and R i g a s s i , D . , 1963. C o n t r i b u t i o n t o t h e g e o l o g y and pal e o n t o l o g y of t h e a r e a of t h e C i t y of La Habana, Cuba and i t s s u r roundings. E c l . Geol. H e l v . v . 56, no. 1, p . 193-480. P a l a e o g e n e o s t r a c o d s from t h e c o n t i n e n t a l s h e l f o f f D i n g l e , R . V . , 1976. N a t a l , S o u t h A f r i c a . ' T r a n s . BOY. SOC. S . A f r i c a , v o l . 4 2 , p t . 1, p . 35-79 * Drooger, C . W . , 1963. Microfauna and age of t h e B a s s e s P l a i n e s Formation of F r e n c h G u y a n a , I . and 11. o s t r a c o d a , p . 460-468; Kon. N e d e r l . Akad. Wetensch., P r o c . s e r . B . , v . 6 3 , no. 4 . Drooger, C . W . and K a a s s c h i e t e r , J . P . H . , 1958. F o r a m i n i f e r a o f t h e Orinoco-Trinidad-Paria s h e l f . Rep. Orinoco S h e l f Exp., v . 4 . Verh. Kon. N e d e r l . Akad. Wetensch., a f d . N a t u u r k . ser. 1, V . 2 2 . Ostracoda, p . 88-92. ( G r g k o f f , N . , 1958. O s t r a c o d e s du B a s s i n d e Congo. 111: T e r t i a i r e . Ann. Mus. Roy. Congo-Belge, v . 2 2 , p. 5-36.] Hartmann, Gerd, 1955. Neue m a r i n e O s t r a c o d e n d e r F a m i l i e C y p r i d a e und d e r 2001. Subfamilie Cytherideinae d e r F a m i l i e Cytheridae aus B r a s i l i e n . Anz. v . 1 5 4 . no. 5/6. D . 109-127. I 1956. Weitere m a r i n e n O s t r a c o d e n aus B r a s i l i e n , I n : B e i t r s g e z u r n e o t r o p i s c h e n F a u n a , p . 19-62 by E . T i t c h a k and H . W. Koepeke, Gustav F i s c h e r V e r l a g , Jena. 1959a. Neue O s t r a c o d e n von T e n e r i f f a . Zool. Anz. v. 1 6 2 , no. 5/6, -1 p . 160-171. , 195933. O s t r a c o d a s d e l a s agunas s a l o b r e s d e America l a t i n a y su significacign para l a investigaci6n palaeontolbgica. A c t . y Trab. p r i m e r Congr. Sudameric. z o o l o g i a ( L a P l a t a ) , V . 2 , p . 169-176. Hartmann, G. and Hartmann S c h r o d e r , G., 1975. Zoogeography and b i o l o g y of Bull. l i t t o r a l O s t r a c o d a from S o u t h A f r i c a , Angola and Mozambique. Amer. P a l e o n t . , V. 6 5 , no. 282, p. 353-367. ( J o n e s , T. R., 1860. F o s s i l E n t o m o s t r a c a from M o n t s e r r a t e ( B r a z i l ) , p . 266268 i n : A l p o r t , S., On the D i s c o v e r y of some F o s s i l Remains near B a h i a i n South America w i t h N o t e s on the F o s s i l s by S i r P. G. E g e r t o n , P r o f . J. M o r r i s and T. R. J o n e s . Q u a r t . J o u r . Geol. SOC. London, 1860, p t . 1, p. 263-268.) Keen, M . C . , 1975. Some R u g g i e r i a - l i k e o s t r a c o d s from t h e T e r t i a r y and R e c e n t o f West A f r i c a . P r o c . 5 t h A f r . C o l l . M i c r o p a l . Rev. ESP. Microp a l . , ser. 7 , no. 3 , p. 451-464. K e i j , A. J . , 1954. O s t r a c o d a : I d e n t i f i c a t i o n and d e s c r i p t i o n o f s p e c i e s i n : Van Andel and Postma: Recent s e d i m e n t s o f the Gulf, o f P a r i a . R e p o r t s O r i n o c o s h e l f Exp., v o l . 1. Verh. Kon. N e d e r l . Akad. Wetensch a f d . Natuurk. ser. 1, v . 20, no. 5 , p . 218-231.
-,
,
p
-, -,
-
517 K e i j , A. J., 1976. Note on H a v a n a r d i a and T r i e b e l i n a s p e c i e s ( C s t r a c o d a ) Kon. N e d e r l . &ad. Wetensch. P r o c . s e r . B , v o l . 7 0 , no. 1, p . 36-44, p l s . 1, 2 , 4 t e x t f i g s . K l i e , W . , 1939a. C s t r a c o d e n a u s den m a r i n e n S a l i n e n von B o n a i r e , Aruba und Curagao. C a p i t a z o o l . , V . 8 , p t . 4 , p . 1-19. 1939b. B r a c k w a s s e r o s t r a c o d e n von N o r d o s t b r a s i l i e n . Zool. J a h r b . A b t . S y s t . , V . 7 2 , p. 359-372. 1940. C s t r a c o d e n von d e r Kcste Deutsch-scdwest A f r i k a s . Beitr. Fauna E u l i t o r a l s von d . SW A f r i k a . K i e l e r M e e r e s f . , V . 3 , no. 2 , p . 404-448. Kogbe, C . A . , Mehes, K . , Le-Calvez, Y . , and G r e k o f f , N . , 1 9 7 5 , Micro-bios t r a t i g r a p h y o f lower T e r t i a r y s e d i m e n t s from t h e s o u t h - e a s t e r n f l a n k s o f t h e Iullemmeden b a s i n ( n o r t h w e s t N i g e r i a ) . P r o c . 5 t h A f r . C o l l . M i c r o p a l . , Rev. Esp. M i c r o p a l . , s e r . 7 , no. 3 , p . 523-538. Krommelbein, K a r l , 1975. Remarks on m a r i n e C r e t a c e o u s C s t r a c o d e s o f Gondwanic d i s t r i b u t i o n . P r o c . 5 t h A f r . Coll. M i c r o p a l . , Rev. Esp. M i c r o p a l . , s e r . 7 , no. 3 , p . 539-551. Macedo, A n t o n i o C . M a g i l h a e z , 1965. A s m i c r o f a u n a s d o sambaqui d e SernamE s t . Rio d e J a n e i r o . Min. Min. e E n e r g . b e t i b a e d o l i t o r a l d e Mag:, D e p t . Nac. P r o d . m i n . , d i v . G e o l . e M i n . , N o t a s p r e l i m i n a r e s e e s t u d o s no. 1 2 8 , p. 2-63. .Yaddocks, R o s a l i e F . , 1969. R e v i s i o n of R e c e n t B a i r d i i d a e ( C s t r a c o d a ) . U . S. Nat. Mus., Nat. H i s t . B u l l . , v . 295, p . 1-126. Malumian, N o r b e r t o , 1970. B i o e s t r a t i g r a f i a d e l T e r c i a r i o Marino d e l subs u e l o d e l a p r o v i n c i a de BuenoS A i r e s ( A r g e n t i n a ) . Ameghiniana, v. 7 , no. 2 , p . 173-204. M a s o l i , M . , 1975. O s t r a c o f a u n e s r g c e n t e s du p l a t e a u c o n t i n e n t a l d e l a c 8 t e d ' I v o i r e en t a n t q u ' i n d i c a t e u r s gcologiques. Proc. 5 t h Afr. C o l l . M i c r o p a l . , Rev. Esp. M i c r o p a l . , s e r . 7 , no. 3 , p. 623-633. McKenzie, K . G . , 1973. C e n o z o i c C s t r a c o d a , i n : H a l l a m A. ( e d i t . ) , A t l a s o f P a l a e o b i o g e o g r a p h y , E l s e v i e r P u b l . , p . 477-487. M i l l e r , G . W . , 1908. D i e O s t r a c o d e n d e r d e u t s c h e n S u d p o l a r - E x p e d i t i o n 1901-1903, i m A u f t r a g d e s Reichamtes d e s I n n e r e n , h e r a u s g e g e b e n von E . von D r y g a l s k i . , v o l . 1 0 , Z o o l . , no. 2 , p . 51-181. N e a l e , J. W . , 1967. An o s t r a c o d f a u n a from H a l l e y Bay, C o a t s Land, B r i t i s h B r i t . A n t a r c t . S u r v . , S c i . Rep. no. 5 8 , p . 1-50. Antarctic Territory. Noordermeer, E l l y , J . and Wagner, C. W . , 1969. P r e l i m i n a r y n o t e on t h e o s t r a c o d f a u n a o f t h e b o r i n g A l l i a n c e 28 i n Surinam (Dutch G u i a n a ) . Geol. e n M i j n b . , V . 2 8 , no. 2 , p. 163-164. Omatsola, M. E b i . , 1970a. Podocopid O s t r a c o d a from t h e Lagos Lagoon, K i geria. M i c r o p a l e o n t o l o g y , V. 1 6 , n o . 4 , p . 407-445. 1970b. Cn S t r u c t u r e and Morphologic V a r i a t i o n o f Kormal P o r e System i n R e c e n t C y t h e r i d O s t r a c o d a ( C r u s t a c e a ) , Acta Z o o l . , v . 5 1 , p. 115124. 197Oc. On t h e o c c u r r e n c e o f C y t h e r e l l i d a e ( O s t r . C r u s t . ) i n a b r a c k i s h - w a t e r Environment. B u l l . Geol. I n s t . Univ. U p p s a l a , NS, V . 2, p. 91-96. 1970d. N o t e s on t h r e e new s p e c i e s o f O s t r a c o d a from t h e N i g e r D e l t a , -1 Nigeria. I b i d . , V . 2 , p . 97-102. , 1971. C a m p y l o c y t h e r e i s , a new genus o f t h e C a m p y l o c y t h e r i n a e ( O s t r . , C r u s t . ) and i t s m u s c l e s c a r v a r i a t i o n . B u l l . C e n t r e Rech. Pau-SNPA, V. 5 , s u p p l . , p . 101-123. 1972. R e c e n t and S u b r e c e n t T r a c h y l e b e r i d i d a e a n d H e m i c y t h e r i d a e ( O s t r . C r u s t . ) from t h e w e s t e r n N i g e r d e l t a , N i g e r i a . B u l l . g e o l . I n s t . U p p s a l a , NS., v o l . e , p . 37-120. , 1972. R e c e n t and S u b r e c e n t T r a c h y l e b e r i d i d a e and H e m i c y t h e r i d a e
-, -,
-,
-, -,
518 P i n t o , I r a j l Damiani and O r n e l l a s , L i l i a P i n t o d e , 1965. A new b r a c k i s h water O s t r a c o d e C y p r i d e i s r i o g r a n d e n s i s P i n t o and O r n e l l a s from s o u t h e r n B r a z i l and i t s o n t o g e n e t i c c a r a p a c e development. E S C . Geol. Univ. do Rio Grande do Sul, p u b l . e s p . no. 8 , p . 7-23. , 1970. A new b r a c k i s h w a t e r o s t r a c o d e p e r i s s o c y t h e r i d e a k r & m e l b e i n i P i n t o and O r n e l l a s , s p . nov. from s o u t h e r n B r a z i l . I b i d . Publ. esp. no. 2 ) , p. 1-19. P i n t o , I r a j i Damiani and S a n g u i n e t t i , Yvonne, T . , 1962. A c o m p l e t e r e v i s i o n o f t h e g e n e r a B i s u l c o c y p r i s and Theriosynoecum ( O s t r a c o d a ) w i t h t h e w o r l d g e o g r a p h i c a l and s t r a t i g r a p h i c a l d i s t r i b u t i o n ( i n c l u d i n g ;*ieEacyprist E l p i d i u m t Gomphocythere and C y t h e r i d e l l a ) I bi d. publ. e s p . no. 4 , 97 pp. Pokorn?, V l a d i m i r , 1968. HavanaEdia, g. n o v . , a new genus of t h e B a i r d i i dae (Ostracoda, C r u s t . ) . V e s t . <st:. c s t . g e o l . v . 4 3 , no. 1, p . 6163. P u r i , H . S . , 1967. E c o l o g i c d i s t r i b u t i o n o f Recent O s t r a c o d a . P r o c . Symp. C r u s t a c e a , p t . 1, p . 457-495. 1971. D i s t r i b u t i o n o f o s t r a c o d e s i n t h e o c e a n s . The Micropalaeont o l o g y of Oceans, Cambr. Univ. P r e s s , p . 163-169. O s t r g c o d o s d e Lagunas d e l a P r o v i n c i a d e Buenos Ramirez, F . C . , 1967. Rev. MUS. de l a P l a t a , N.S., s e c c . , Z o o l . , v . 10; p . 5-54. Aires. Ramlrez, F. C. and Moguilevsky, A . , 1971. O s t r a c o d o s p l a n k t o n i c o s h a l l a d o s en a g u a s o c e z n i c a s f r e n t e a l a P r o v i n c i a d e Buenos A i r e s . R e s . XLI comissao o c e a n o g r a f i c a C o s t a Sul. P h y s i s , v . 30, no. 81, p . 637-665. Reyment, R. A . , 1959. D i e O s t r a c o d e n g a t t u n g P a i j e n b o r c h e l l a i m U n t e r o z a n Nigeriens. Acta Univ. Stockholm., Stockholm C o n t r . i n G e o l . , V . 3 , no. 7 , p. 139-143. 1960. N o t e s on t h e C r e t a c e o u s - T e r t i a r y boundary i n N i g e r i a . Rep. 2 1 s t S e s s . I n t . Geol. Congr. Norden, p t . 5 , s e c t . 3 , p r o c . : The C r e t a c e o u s - T e r t i a r y boundary, p . 131-135. , 1963. S t u d i e s on N i g e r i a n Upper C r e t a c e o u s and Lower T e r t i a r y Ost r a c o d a P t . 2 : D a n i a n , P a l e o c e n e and Eocene O s t r a c o d a . A c t a Univ. Stockholm. Stockholm C o n t r . i n Geol., V . 1 0 , p. 1-286. , 1964. B i o s t r a t i g r a p h i e e t m i c r o p a l 6 o n t o l o g i e du P a l g o g s n e d e l a Nigeria occidentale. Mem. B . R . G. M . , V . 28, no. 2 , p. 839-847. 1965. Q u a n t i t a t i v e morphologic v a r i a t i o n and c l a s s i f i c a t i o n o f some N i g e r i a n P a l e o c e n e C y t h e r e l l i d a e . M i c r o p a l e o n t o l o g y , V . 11, no. 4 , p. 457-465. , 1966. S t u d i e s on N i g e r i a n Upper C r e t a c e o u s and Lower T e r t i a r y Ost r a c o d a , P t . 3 : S t r a t i g r a p h i c a l , P a l e o e c o l o g i c a l and b i o m e t r i c a l conclusions. A c t a Univ. Stockholm., Stockholm C o n t r . Geol., v. 1 4 , 1 5 1 p . Reyment, R . A . and E l o f s o n , O., 1959. Zur K e n n t n i s d e r O s t r a c o d e n g a t t u n g Buntonis. I b i d . , V . 3 , no. 9 , p. 157-164. Reyment, R. A. and Reyment, E . , 1959. B a i r d i a i l a r o e n s i s S. nov. a u s dem PaleozZn N i g e r i e n s und d i e G i i l t i g k e i t d e r G a t t u n g B a i r d o p p i l a t a ( O s t r . Crust.). I b i d . , V . 3 , no. 2 , p . 59-68. Reyment, R. A. and Van V a l e n , L . , 1969. B u n t o n i a olokunudui s p . nov. ( 0 s s t r a c o d a , C r u s t a c e a ) . I b i d . N.S. v . 1, no. 3 , p. 83-94. R o s s i d e G a r c f a , E l s a , 1966a. Sobre l a p r e s e n c i a d e l g e n e r o c y t h e r i d e a en l a d e p r e s i d n d e el Sampal ( C h u b u t ) . Rev. Geol. A r g e n t . V . 2 , no. 2 , p. 118. 1966b. C o n t r i b u c i o ' n a1 c o n o c i m i e n t o de 10s o s t r & o d o s d e l a Argent i n a I : Formacidn E n t r e R i o s , d e V i c t o r i a , p r o v i n c i a d e E n t r e R i o s . I b i d . v. 21, no. 3 , p . 194-208. , 1967a. Un nuevo ge'nero d e o s t r & o d o s d e l a f a m i l i a T r a c h y l e b e r i d i dae. I b i d . v. 2 2 , no. 1, p. 95-98.
-
.
-,
-, -
-,
-
,
-
519
-t,i n1967b. a 11:
C o n t r i b u c i g n a 1 c o n o c i m i e n t o d e 10s o s t r i c o d o s d e l a ArgenO s t r i c o d o s d e l cord6n l i t o r a l Loma d e Tajamar. Ibid. V. 22, no. 3, p . 203-208. -1 1969. Algunos o s t r i c o d o s d e l E n t e r r i e n s e d e P a r a n i , p r o v i n c i a de E n t r e Rios. I b i d . V . 24, no. 3, p. 267-280. , 1971. O s t r a c o d e s du Miocsne d e l a Rgpublique A r g e n t i n a . A c t . 4e Coll. A f r . M i c r o p a l . , p. 391-431. 1972. C u v i l l i e r i n a , nuevo g e n e r o de o s t r i c o d o s . Rev. Esp. Microp a l . , v . 4 , no. I, p. 23-26. S c o t t , Th., 1899. R e p o r t on t h e m a r i n e and f r e s h w a t e r c r u s t a c e a from F r a n z J o s e p h Land, c o l l e c t e d by M r . W . S . Bruce of t h e Jackson-Harmsworth Expedition. Linn. SOC. J o u r n . Zool. London, v. 27, p. 60-126. 1912. The E n t o m o s t r a c a o f t h e S c o t t i s h N a t i o n a l A n t a r c t i c Expedition. T r a n s . Roy. SOC. E d i n b u r g h , v . 4 8 , p t . 3 , p. 521-600. Teeter, J. W . , 1975. D i s t r i b u t i o n of Holocene O s t r a c o d a from Belize. In: Kenneth F. Wantland and Walter C. Pusey 111: B e l i z e S h e l f - C a r b o n a t e S e d i m e n t s , C l a s t i c s e d i m e n t s and Ecology. Amer. Assoc. P e t r . G e o l , S t u d i e s i n Geology, N o . 2 , p. 400-499. Whatley, R. C. and C h o l i c h , Teresa d e l Carmen, 1974, A new Q u a t e r n a r y ost r a c o d g e n u s from A r g e n t i n a . P a l a e o n t o l o g y , v. 1 7 , no. 3, p. 669-684. W h a t l e y , R. C. and Moguilevsky, A., 1975. The f a m i l y L e p t o c y t h e r i d a e i n A r g e n t i n e waters. B u l l . Amer. P a l e o n t . V. 6 5 , no. 2 8 2 , p. 501-521.
-,
-,
This Page Intentionally Left Blank
521
C E N O Z O I C RADIOLARIANS O F THE ATLANTIC BASIN AND XPAGINS
R i c h a r d E . C a s e y a n d K e n n e t h J. M c M i l l e n R i c e U n i v e r s i t y , ousto on, T e x a s
ABSTRACT Living r a d i o l a r i a n s i n the A t l a n t i c a r e i n d i c a t i v e of spec i f i c water masses. Radiolarian skeletons occur only i n the u p p e r f e w c e n t i m e t e r s of l a t e N e o g e n e s e d i m e n t s o v e r most o f t h e mid a n d l o w e r l a t i t u d e s o f t h e A t l a n t i c . T h i s i s i n cont r a s t t o common o c c u r r e n c e s o f r a d i o l a r i a n s i n t h e s e d i m e n t s o f t h e s e l a t i t u d e s d u r i n g much of t h e P a l e o g e n e a n d e a r l y Neogene. P a l e o g e n e t o e a r l y Neogene r a d i o l a r i a n b i o s t r a t i g r a p h ies a r e cosmopolitan due t o t h e p a n - t r o p i c a l o c e a n i c connection. I s o l a t i o n o f t h e A t l a n t i c s t a r t i n g i n Mid-Miocene res u l t e d i n c h a n g e s i n s i l i c e o u s s e d i m e n t a t i o n and t h e d e v e l o p ment o f p r o v i n c i a l r a d i o l a r i a n f a u n a s . The d e v e l o p m e n t o f a new c o s m o p o l i t a n r a d i o l a r i a n z o n a t i o n b a s e d on t r o p i c a l s u b m e r g e n t f o r m s may a i d i n i n t e r o c e a n i c warm w a t e r c o r r e l a t i o n s . R ~ S U M ~
Les r a d i o l a i r e s a c t u e l s d e 1 ' A t l a n t i q u e c a r a c t g r i s e n t des masses d ' e a u dgtermin6es. L e s s q u e l e t t e s d e r a d i o l a i r e s se t r o u v e n t seulement dans les g u e l q u e s c e n t i m g t r e s s u p g r i e u r s d e s s g d i m e n t s d u Ne'ogsne s u p 6 r i e ~ rd a n s l a p l u p a r t d e s l a t i t u d e s moyennes e t i n f g r i e u r e s d e 1 ' A t l a n t i q u e . C e c i f o r m e un c o n t r a s t e avec l e u r abondance r e l a t i v e dans les s6diments d e ces l a t i t u d e s d u r a n t l a p l u s g r a n d e p a r t i e d u P a l g o g b n e e t du NCogsne I n f 6 r i e u r . Les b i o s t r a t i g r a p h i e s des r a d i o l a i r e s e n t r e l e P a l C o g s n e e t l e N6og;ne I n f 6 r i e u r s o n t c o s m o p o l i t g s 'a c a u s e d e l a communication oce'anigue p a n - t r o p i c a l e . A partir du m i l i e u du Mio c e n e l ' i s o l e m e n t d e 1 ' A t l a n t i q u e c a u s e d e s changements d a n s l e s g d i m e n t a t i o n d e l a s i l i c e e t provoque l e dgveloppement d e f a u n e s p r o v i n c i a l e s d e r a d i o l a i r e s . Le d 6 v e l o p p e m e n t , p o u r l e s r a d i o l a i r e s , u n e b i o s t r a t i g r a p h i e nouv e l l e e t c o s m o p o l i t a i r e base'e s u r les formes t r o p i c a l e s s u b mergges a i d e r a p e u t - 8 t r e 5 de'terminer d e s c o r r g l a t i o n s i n t e r ocCaniques e n t r e les r 6 g i o n s t r o p i c a l e s .
522
R A D I O L A R I A N PRESERVATION I N THE FTLANTIC THROUGHOUT THE CENOZOIC
FND ITS IMPLICATIONS The P a l e o g e n e and e a r l y Neogene P t l a n t i c a p p e a r s t o be f a v o r a b l e t o r a d i o l a r i a n p r e s e r v a t i o n a s a r e t h e o t h e r major ocean b a s i n s of t h a t t i m e .
F major change i n r a d i o l a r i a n p r e s e r v a -
t i o n o c c u r r e d d u r i n g t h e mid-Miocene a l a t a Zone t i m e ) .
( o r a t about Dorcadospyris
From t h e mid-Miocene t o t h e p r e s e n t r a d i o -
l a r i a n s h a v e b e e n o n l y r a r e l y and s p o r a d i c a l l y p r e s e r v e d i n F t l a n t i c b a s i n p r o p e r , and i t s m a r g i n a l b a s i n s , from a b o u t 4 0 d e g r e e s n o r t h , a l t h o u g h p o l e w a r d of e a c h of t h e s e l a t i t u d e s r a d i o l a r i a n s a r e commonly b e i n g p r e s e r v e d t o d a y a s t h e y h a v e b e e n t h r o u g h o u t much of t h e C e n o z o i c .
This p a t t e r n i s u n l i k e
t h e P a c i f i c p a t t e r n t h a t a p p e a r s t o h a v e remained r e l a t i v e l y constant throughout t h e Cenozoic with r a d i o l a r i a n oozes forming underneath t h e e q u a t o r i a l divergences,
radiolarian-diatom
oozes
f o r m i n g p o l e w a r d of t h e p o l a r c o n v e r g e n c e s and r e l a t i v e l y h i g h radiolarian contents
( s o m e t i m e s becoming r a d i o l a r i a n o o z e s )
u n d e r n e a t h t h e e a s t e r n and w e s t e r n b o u n d a r y c u r r e n t s y s t e m s . F major change s e e n i n P a c i f i c r a d i o l a r i a n s e d i m e n t a t i o n ap-
p e a r s t o be t h e i n c r e a s e i n r a d i o l a r i a n
(and s i l i c e o u s forms i n
g e n e r a l ) s e d i m e n t s b e g i n n i n g i n t h e mid-Miocene
( a t t h e same
t i m e the r a d i o l a r i a n s w e r e dropping out of t h e p i c t u r e i n t h e A t l a n t i c ) and c o n t i n u i n g b u t d e c r e a s i n g somewhat i n m a g n i t u d e t o the present.
K o b a y a s h i ( 1 9 4 4 ) h a s n o t i c e d an a l t e r n a t i o n o f
r a d i o l a r i a n r o c k s i n t h a t when t h e y e x h i b i t a g r e a t d e v e l o p ment i n t h e e a s t e r n F s i a t i c r e g i o n , t i m e e q u i v a l e n t r o c k s i n t h e A u s t r a l i a n r e g i o n from P a l e o z o i c t o T e r t i a r y a r e r a d i o l a r i a n poor.
T h e r e f o r e t h e e x p l a n a t i o n of t h e d e c l i n e o f r a -
d i o l a r i a n s e d i m e n t a t i o n i n t h e P t l a n t i c and t h e i n t e n s i f i c a t i o n of r a d i o l a r i a n s e d i m e n t a t i o n i n t h e P a c i f i c
(probably a t t h e
e x p e n s e of r a d i o l a r i a n s e d i m e n t a t i o n i n t h e F t l a n t i c ) d u r i n g t h e Neogene may w e l l h a v e h a d e a r l i e r c o u n t e r p a r t s . A l t h o u g h r a d i o l a r i a n s a r e e s s e n t i a l l y a b s e n t from Neogene m i d and low l a t i t u d e A t l a n t i c s e d i m e n t s t h e y a r e q u i t e common
and i n many c a s e s t h e d o m i n a n t b i o g e n i c e l e m e n t i n many P a l e o -
Figure 1
TECTONIC
A N D OCEANIC EVENTS I N THE ATLANTIC A N D T H t - I R POSSIBLE EFFECTS ~ ON RADIOLARIANS _ A_ N D S I L I C E O U S SEDIMENTATION PUSSIBLE EFFECT
EVEN1
1.
Development o f d i a t o m s and rapid increase i n diversit y and "abundance" d u r i n q t h e Cenozoic
2.
Separation o f Australia and A n t a r c t i c a about L a t e P a l e o g e n e ( a b o u t 3 5 m.y.a.) (Kennett e t a l . . 1972)
3.
Closure of e a s t e r n Tethys (Africa-Arabia joined t o S . W . A s i a ) i n e a r l y Mioc e n e a b o u t 1 8 t o 20 m.y.a. and c l o s u r e o f w e s t e r n tethys (Europe-Africa join a t Gibraltar) i n mid-Miocene a b o u t 12-14 m i l l i o n y e a r s ago ( B e r g g r e n a n d Van C o u v e r i n g , 1 9 7 4 ) and e l e v a t i o n o f Panamanian b l o c k t o a b o u t 1500 m e t e r s (Bandy and Casey, 1 9 7 3 )
4.
E l e v a t i o n o f t h e Panamanian Block t o effective s i l l , s t o p p i n g communicat i o n between t h e e q u a t o r i a l A t l a n t i c and P a c i f i c completely a t about the M i o c e n e - P l i o c e n e bounda r y ( 4 t o 5 m.y.a.)
1.
R a d i o l a r i a n s a d a p t t o development o f d i a t o m s and c o m p e t i t i v e pressure f o r s i l i c a by u s i n q l e s s s i l i c a i n t e s t .
2.
D e v e l o p m e n t o f a circum-Antarctic c u r r e n t r e s u l t i n g i n t h e r m a l i s o l a t i o n o f A n t a r c t i c a and i n c r e a s e d g l a c i a t i o n i n A n t a r c t i c a . P a l e o q e n e " A n t a r c t i c B o t t o m W a t e r " f o r m e d and f l o w s i n t o A t l a n t i c through newly-formed pas5 i n t h e Rio Grande-Walvis Ridge c h a n g i n g t h e m i d and l o w l a t i t u d e A t l a n t i c f r o m a c a r b o n a t e s i n k o r l a y o o n a l - t y p e b a s i n ( t h a t i t had been d u r i n g most o f t h e Paleogene) t o a s i l i c a s i n k o r e s t u a r i n e - t y p e b a s i n ( w i t h b o t h s i l i c a and c a r b o n a t e b e i n g d e p o s i t e d ) s i m i l a r t o t h e p r e s e n t day P a c i f i c . T h i s p e r s i s t e d u n t i l e a r l y Miocene.
3.
A t l a n t i c i s o l a t e d f r o m e q u a t o r i a l I n d i a n Ocean a n d c o n s i d e r a b l y r e s t r i c t e d from the equatorial Pacific, resultinq i n the develo p m e n t o f Neogene w a t e r m a s s r e q i m e s and r a d i o l a r i a n f a u n a s e n demic t o t h e s e w a t e r masses ( s u c h a s t h o s e r e f e r r e d t o i n t h e l i v i n g radiolarians i n t h i s paper). P e r i o d s o f g l a c i a t i o n due t o d e v e l o p m e n t o f m e r i d i a n a l c i r c u l a t i o n and p e r i o d s o f p r o d u c t i o n o f Mediterranean intermediate water resulting i n saline waters r e a c h i n g t h e A n t a r c t i c and l o w e r i n q t h e f r e e z i n g p o i n t d e p r e s s i o n a n d r e s u l t i n g i n " i n t e r o l a c i a l s " and p e r i o d s o f no F e n e r a t i o n o f M e d i t e r r a n e a n I n t e r m e d i a t e Water r e s u l t i n g i n M e d i t e r r a n e a n e v a p o r i t e 8 and " f r e s h e n i n g " o f A n t a r c t i c w a t e r s and r e s u l t i n g " q l a c i a l s " . T h i s p e r i o d i s t h e p e r i o d of change f r o m an P . t l a n t i c s i l i c a s i n k ( e s t u a r i n e t y p e b a s i n ) t o a c a r b o n a t e b a s i n ( l a g o o n a l t y p e ) w h i c h has p e r s i s t e d t o t h e p r e s ent. (Ryan, e t a l . 1 9 7 4 )
4.
Development o f p r o v i n c i a l r d d i o l a r i d n faunas i n t h e l o w l d t i t u d e oceans w i t h " l e a k i n q " o f l o w l a t i t u d e P a c i f i c and I n d i a n O c e a n f a u n a s i n t o t h e A t l a n t i c a r o u n d t h e Cape o f Good Hope and t h e r e t e n t i o n o f a r e l i c t r a d i o l a r i a n f a u n a i n t h e equat o r i a l A t l a n t i c and i t s m a r g i n a l seas ( G u l f o f M e x i c o and Caribbean). T h e a l m o s t c o m p l e t e c l o s u r e a t Panama i n t e n s i f i e s t h e G u l f S t r e a m , c a r r y i n g warm w a t e r t o h i q h l a t i t u d e s , a l l o w i s l a n d h o p p i n g o f v e r t e b r a t e faunas between N o r t h and South America, and r e s u l t s i n i n c r e a s e d p r e c i p i t a t i o n and t h e d e v e l opment o f n o r t h e r n h e m i s p h e r e g l a c i a t i o n a t a b o u t 3 m.y.a. I n c r e a s e d g l a c i a t i o n i n t h e P l e i s t o c e n e may r e s u l t f r o m t h e m o r e c o m p l e t e r e s t r i c t i o n t h r o u g h Panama a t t h a t t i m e .
524
g e n e and e a r l i e s t Neogene s e d i m e n t s of t h e same a r e a .
Our r e -
s e a r c h on t h i s s u b j e c t i s s t i l l i n i t s e a r l y s t a g e s , however w i t h t h e previous d i s c u s s i o n a s background t h e f o l l o w i n g para5raph p r e s e n t s a p o s s i b l e e x p l a n a t i o n f o r t h e Cenozoic s t r a t i g r a p h i c and g e o g r a p h i c d i s t r i b u t i o n o f r a d i o l a r i a n s i n t h e Atlantic. R a d i o l a r i a n s e d i m e n t s a p p e a r t o h a v e b e e n a common c o n s t i t u e n t o f t h e w o r l d o c e a n s e d i m e n t s of t h e P a l e o g e n e and e a r l y Miocene i n c l u d i n g t h e A t l a n t i c .
I n t h e mid-Miocene
( a t about
D o r c a l o s p y r i s a l a t a Zone t i m e ) t h e s e d i m e n t d e p o s i t e d i n t h e mid and low l a t i t u d e A t l a n t i c became e s s e n t i a l l y d e v o i d of r a d i o l a r i a n s e x c e p t for s p o r a d i c o c c u r r e n c e s and some " n e a r s h o r e " ( C a l v e r t F o r m a t i o n ) and m a r g i n a l b a s i n ( E u r o p e a n stratotypes) occurrences.
The main r e a s o n s f o r t h i s c h a n g e ap-
pear t o b e i s o l a t i o n of t h e A t l a n t i c B a s i n , p a l e o c l i m a t i c c h a n g e s , t h e c h a n g e of t h e A t l a n t i c B a s i n from a n " e s t u a r i n e t y p e " b a s i n t o a " l a g o o n a l t y p e " b a s i n ( i n B e r g e r ' s terms, B e r g e r , 1 9 7 0 ) and t h e r a p i d d e v e l o p m e n t o f d i a t o m s and r e s u l t a n t " c o m p e t i t i o n " o f d i a t o m s and r a d i o l a r i a n s f o r a v a i l a b l e ( H a r p e r and K n o l l , 1 9 7 5 ) .
silica
The p r o p o s e d e v e n t s r e s p o n s i b l e f o r o r c o r r e l a t e d and r e l a t e d to t h i s c h a n g e i n much of t h e P t l a n t i c from a s i l i c a s i n k b a s i n and a p r o p o s e d h i s t o r y f o r t h e A t l a n t i c r e l a t e d t o r a d i o l a r i a n h i s t o r y a r e l i s t e d on F i g u r e 1
C o n d i t i o n s f o r optimum
r a d i o l a r i a n p r e s e r v a t i o n and optimum r a d i o l a r i a n d i s s o l u t i o n i n s e d i m e n t s a r e s u g g e s t e d a l o n g w i t h p r e s e n t d a y and p a s t e x amples o f
Figure 2
C E N O Z O I C RADIOLARIAN BIOSTRATIGRAPHY O F THE A T L A N T I C
R i e d e l and S a n f i l i p p o h a v e d e v e l o p e d a "warm w a t e r " r a d i o l a r i a n b i o s t r a t i g r a p h y f o r t h e Cenozoic u s i n g m a t e r i a l mainly from P a c i f i c Deep S e a D r i l l i n g P r o j e c t c o r e s filippo, in press).
( R i e d e l and San-
T h i s z o n a t i o n a p p e a r s t o work q u i t e w e l l
i n "warm w a t e r " P a l e o g e n e and e a r l y Neogene A t l a n t i c s e d i m e n t s .
Figure 2
CONDITIONS FOR OPTIMUM RADIOIARIAN PRESERVATION AND OPTIMUM RADIOLARIAN DISSOLUTION
OPTIMUM PRESERVATION
OPTIMUM DISSOLUTION
Iiow Latitudes
Low Latitudes 1. Low Productivity
1.
2.
High Productivity (Upwelling areas, under boundary currents and equatorial divergences) Low Dissolution Potential of waters (Estuarine type waters)
(within mid-latitude qyres) 2.
High dissolution potential of waters (laqoonal type waters)
3.
Rapid radiolarian sedimentation (fecal pellet sedimentation)
3.
Low radiolarian sedimentation (individual test sedimentation)
4.
Relatively low benthonic activity per unit time
4.
Relatively hiqh benthonic activity per unit time
5.
Low dissolution potential in sediments (high silica content in sediments and interstital waters plus low pH)
5.
Hiqh dissolution potential in sediments (low silica content in sediments and interstital waters plus high pH)
6.
High rate of terriqenous sedimentation
6.
JXw rate of terriqenous sedimentation
Hiqh Latitude 1. Medium productivity (away from polar diveqrmces)
High Latitudes 1. High productivity (developnent of centric diatoms to exclusion of shallow radiolarians (at polar divergences) or low productivity)
2.
Rapid radiolarian sedimentation (fecal pellet sedimentation)
2.
Low radiolarian sedimentation (individual test sedimentation)
3.
Relatively l o w benthonic activity per unit time
3.
Relatively high benthonic activity per unit time
4.
Low dissolution potential in sediments (high silica content in sediments and interstitial water plus low PH)
4.
High dissolution potential in sediments (low silica content in sediments and interstital waters plus high pH)
5.
High rate of terrigenous sedimentation
5.
Low
rate of terrigenous sedimentation
cn N cn
5 26
P o s t e a r l y Neogene
( o r more s p e c i f i c a l l y p o s t a b o u t D o r c a d o s p y -
r i s a l a t a Zone) s e d i m e n t s c o n t a i n i n g s i g n i f i c a n t r a d i o l a r i a n s
~~
f o r b i o s t r a t i g r a p h i c p u r p o s e s a r e u s u a l l y l a c k i n g . The few occ u r r e n c e s o f " r a d i o l a r i a n r i c h s e d i m e n t s " , i n c l u d i n g t h o s e from t h e l a t e Neogene s t r a t o t y p e l o c a l i t i e s
( R i e d e l and S a n f i l i p p o ,
i n p r e s s ) a r e d i f f i c u l t t o f i t i n t o R i e d e l and S a n f i l i p p o ' s zonation.
The r e a s o n f o r t h i s d i f f i c u l t y i s most l i k e l y d u e t o
t h e " i s o l a t i o n " o f t h e A t l a n t i c and p a c i f i c - I n d i a n Ocean f a u n a s a s mentioned p r e v i o u s l y i n t h i s c h a p t e r .
A s mentioned e a r l i e r
w e s u g g e s t t h a t t h e f i r s t o c c u r r e n c e d a t u m p l a n e s o f R i e d e l and S a n f i l i p p o ' s Z o n a t i o n ( p o s t Mid-Miocene) may be r e l i a b l e mark-
e r s a l t h o u g h t h e y may l a g i n t i m e somewhat b e h i n d t h e i r a p p e a r a n c e i n t h e P a c i f i c - I n d i a n Ocean.
Nigrini
(1971) developed a
Quaternary r a d i o l a r i a n zonation for the e q u a t o r i a l P a c i f i c . Some m e m b e r s s h e u s e d a s Q u a t e r n a r y f i r s t o c c u r r e n c e d a t u m s s u c h a s B u c c i n o s p h a e r a i n v a g i n a t a H a e c k e l and Amphirhopalum y p s i l o n H a e c k e l o c c u r i n t h e l i v i n g p l a n k t o n o f t h e low l a t i t u d e A t l a n t i c and t h e i r f i r s t o c c u r r e n c e s m i g h t a l s o be u s e d i n Atlantic.
However t h e l a s t o c c u r r e n c e d a t u m s a p p e a r t o be
u n r e l i a b l e d u e t o t h e " o n e way" l e a k a g e m e n t i o n e d e a r l i e r . finding of l i v i n g Spongaster pentas
The
( R i e d e l and S a n f i l i p p o ) and
r e l a t e d f o r m s i n p l a n k t o n tows and s u r f a c e s e d i m e n t s o f t h e low l a t i t u d e F t l a n t i c and i t s m a r g i n a l s e a s a t t e s t s t o t h i s u n r e l i a b i l i t y of l a s t o c c u r r e n c e datums i n t h e A t l a n t i c . Hays
( 1 9 6 5 ) , Hays and Opdyke ( 1 9 6 7 ) , Bandy e t a 1 ( 1 9 7 1 )
have developed r a d i o l a r i a n z o n a t i o n s f o r t h e A n t a r c t i c which s h o u l d be u s a b l e f o r t h e A n t a r c t i c and s o u t h e r n F t l a n t i c p o r t i o n s of t h e A t l a n t i c .
T h e y e r and Hammond ( 1 9 7 4 ) h a v e r e l a t e d
t h e r a d i o l a r i a n s t r a t i g r a p h y of R i e d e l and S a n f i l i p p o ( 1 9 7 1 ) w i t h m a g n e t o s t r a t i g r a p h y f o r t h e Neogene i n t h e e q u a t o r i a l Pacific.
Casey
( i n p r e s s b ) a n d J o h n s o n and K n o l l ( 1 9 7 5 ) h a v e
r e l a t e d N i g r i n i ' s zonation t o t h e paleomagnetic s c a l e i n cores from t h e e q u a t o r i a l p a c i f i c .
These r a d i o l a r i a n z o n a t i o n s , s i g -
n i f i c a n t d a t u m p l a n e s and t h e i r r e l a t i o n s t o m a g n e t o s t r a t i g r a p h y a r e i l l u s t r a t e d on F i g u r e s 9 and 1 0 .
The g e n e r a l outcome
527
o f t h i s c o r r e l a t i o n s u g g e s t s t h a t t h e P a l e o g e n e and e a r l y Neog e n e o f t h e "warm w a t e r " A t l a n t i c c a n be r e l i a b l y zoned u s i n g R i e d e l and S a n f i l i p p o ' s Z o n a t i o n .
The warm w a t e r mid and l a t e
Neogene o f t h e A t l a n t i c e x h i b i t s p r o v i n c i a l i s m and t h e h i g h l a t i t u d e S o u t h A t l a n t i c c a n be r e l i a b l y zoned w i t h i n t h e l a t e Neogene. P l a n k t o n t o w s and H o l o c e n e s e d i m e n t s a m p l e s c o l l e c t e d i n t h e t r o p i c a l P t l a n t i c , G u l f o f Mexico and C a r i b b e a n h a v e y i e l d ed r a d i o l a r i a n s p r e v i o u s l y b e l i e v e d t o h a v e b e e n e x t i n c t , and i n f a c t u s e d i n C e n o z o i c "warm-water" r a d i o l a r i a n b i o s t r a t i g raphic zonations.
R a d i o l a r i a n s c o l l e c t e d i n p l a n k t o n tows and
s t a i n e d w i t h Rose B e n g a l i n c l u d e S p o n g a s t e r p e n t a s , S p o n g a s t e r b e r m i n g h a m i , and " c i r c u l a r " and " e l l i p t i c a l " s p o n g o d i s c i d s . The e v o l u t i o n o f S p o n g a s t e r p e n t a s from S p o n g a s t e r b e r m i n g hami o c c u r r e d about 4 . 5 m i l l i o n y e a r s ago i n t h e t r o p i c a l Pacific
( T h e y e r and Hammond, 1 9 7 4 ) and i s u s e d t o d e f i n e t h e
b a s e of t h e S p o n g a s t e r p e n t a s Zone ( R i e d e l and S a n f i l i p p o , i n press).
S p o n g a s t e r b e r m i n g h a m i a p p a r e n t l y became e x t i n c t
s h o r t l y a f t e r t h e e v o l u t i o n of S p o n g a s t e r p e n t a s , and S p o n g a s -
t e r p e n t a s became -~ Pacific
e x t i n c t about 3.6 m i l l i o n y e a r s ago i n t h e
(Casey, i n p r e s s b ) .
Specimens of
" c i r c u l a r " and
" e l l i p t i c a l " s p o n g o d i s c i d s b e l i e v e d t o be t h e a n c e s t o r s of S p o n g a s t e r berminghami have a l s o been c o l l e c t e d i n t h e p l a n k t o n . These s p e c i e s r e p r e s e n t a r e l i c t r a d i o l a r i a n fauna i n t h e t r o p i c a l A t l a n t i c , G u l f o f Mexico and C a r i b b e a n . sence
Their pre-
s u g g e s t s some i n t e r e s t i n g c o n s e q u e n c e s o f b o t h b i o s t r a -
t i g r a p h i c and p a l e o o c e a n o g r a p h i c s i g n i f i c a n c e .
of
biostratig-
r a p h i c s i g n i f i c a n c e i s t h e c o n c l u s i o n t h a t t h e g e o l o g i c and g e o g r a p h i c r a n g e s of some o f t h e s p e c i e s u s e d i n R i e d e l and S a n f i l i p p o ' s zonation a r e p r o v i n c i a l (they have suggested t h i s themselves)
( S a n f i l i p p o and R i e d e l , 1 9 7 4 ) .
This p r o v i n c i a l i t y
i s a r e a l p r o b l e m b e c a u s e t h e l a t e Neogene p a r t o f t h e i r zonat i o n was m a i n l y d e v e l o p e d u s i n g t r o p i c a l P a c i f i c c o r e s , and t h e findings here suggest t h a t t h e radiolarian biostratigraphy
528
(and perhaps o t h e r m i c r o f o s s i l b i o s t r a t i g r a p h i e s ) i n t h e A t l a n t i c , and t h e s t r a t o t y p e l o c a l i t i e s o f t h e l a t e Neogene i n E u r o p e s h o u l d be q u i t e d i f f e r e n t from t h e "warm-water'' P a c i f i c z o n a t i o n of R i e d e l and S a n f i l i p p o ( R i e d e l and S a n f i l i p p o , i n press).
C o r r e l a t i o n a t t e m p t s o f t h e P a c i f i c and E u r o p e a n s t r a -
totype r s d i o l a r i a n assemblages have m e t with l i m i t e d success, p r o b a b l y due i n p a r t t o t h e problem of p r o v i n c i a l i t y h e r e i n mentioned
.
T h i s p r o v i n c i a l i t y f o r t h e l a t e Neogene h a s n o t b e e n o b v i -
ous b e f o r e d u e t o t h e f a c t t h a t t h e s e d i m e n t s o f t h e l o w - l a t i t u d e A t l a n t i c and i t s m a r g i n a l s e a s a r e e s s e n t i a l l y v o i d o f r a d i o l a r i a n s p o s t mid-Miocene.
The u p p e r few c e n t i m e t e r s o f
H o l o c e n e s e d i m e n t s i n t h e G u l f o f M e x i c o and C a r i b b e a n d o c o n t a i n r a d i o l a r i a n s and among them S p o n g a s t e r p e n t a s , S p o n g a s __ t e r berminghami,
and " c i r c u l a r " and " e l l i p t i c a l " s p o n g o d i s c i d s .
The p a l e o o c e a n o g r a p h i c s i g n i f i c a n c e i s p e r h a p s e v e n more important.
The A t l a n t i c and p a c i f i c a p p e a r t o e x h i b i t more
o r l e s s " c o s m o p o l i t a n " , "warm-water" r a d i o l a r i a n b i o s t r a t i g r a p h i e s u p t o a t l e a s t mid-Miocene. p o s t mid-Miocene
(See Figure 3 ) .
Sometime
t h e r e appears t o have been a divergence i n
r a d i o l a r i a n f a u n a s and a d e v e l o p m e n t o f g r e a t e r p r o v i n c i a l i s m . The r e a s o n s f o r t h i s d i v e r g e n c e a r e a p p a r e n t l y r e l a t e d t o g e o g r a p h i c and c l i m a t i c i s o l a t i o n and r e s u l t a n t a l l o p a t r i c s p e c i a t i o n and d i f f e r e n t i a l g e o l o g i c r a n g e s o f t h e s e i s o l a t e d populations. The a u t h o r s o f t h i s p a p e r b e l i e v e t h a t t h e g e o g r a p h i c i s o l a t i o n o f t h e t r o p i c a l P a c i f i c and P t l a n t i c was d u e t o t h e e v e n t s l i s t e d on F i g u r e 1 and c u l m i n a t i n g ir. t h e u p l i f t of t h e Panamanian Block d u r i n g t h e Miocene
(Bandy and C a s e y , 1 9 7 3 ) t o
" e f f e c t i v e s i l l " a t about 4 . 5 m i l l i o n y e a r s ago.
Isolation is
placed a t about 4 . 5 m i l l i o n y e a r s ago f o r p r i o r t o t h i s t i m e t h e S p o n g a s t e r f a u n a s o f t h e G u l f o f Mexico and C a r i b b e a n r e -
semble t h o s e o f t h e P a c i f i z b u t d i v e r g e s h o r t l y t h e r e a f t e r . A t 4 . 5 m i l l i o n y e a r s ago, o r a t about t h e Miocene-Pliocene
529
RIEDEL AND SANFILIPPO (IN PRESS) EPOCHS
ZONES -
QUAT .
LAMPROCYRTIS HAYS I PTEROCANIUM PRISMATIUM SPONGASTER PENTAS STICHOCORYS PEREGRINA
PLIOCENE
W
' 0
w
p
L. NEOHETEROPOROS TO L. HAYS1 L.O. STICHOCORYS PEREGRINA S. BERMINGHAMI TO S. PENTAS S. DELMONTENSIS TO S. PEREGRINA O?IMATARTUS L.O. OMMATARTUS PENULTIMUS HUGHES1 OMMATARTUS C. PETTERSSONI TO ANTEPENULTIMUS 0. HUGHES1 CANNARTUS F.O. C. PETTERSSONI PETTERSSONI
--
ALATA o CALOCYCLETTA Y COSTATA STICHOCORYS WOLFFII STICHOCORYS 8 DELMONTENSIS CYRTOCAPSELLA TETRAPERA LYCHNOCANOMA ELONGATA DORCADOSPYRIS OLIGOCENE ATEUCHUS THEOCYRTIS TUBEROSA THYSOCYRTIS BROMIA PODOCYRTIS GOETHEANA PODOCYRTIS C HALARA PODOCYRTIS -. MITRA 0 PODOCYRTIS AMPLA w --- THYRSOCYRTIS TRIANCANTHA THEOCAMPE W WONGOLFIERI u THEOCOTYLE CRYPTOCEPHALA ii CRYPTOCEPHALA PHORMOCYRTIS STRIATA STRIATA BURYELLA CLINATA BEKOMA PALEOCENE BIDARFENSIS V
2
3
~
3
RADIOLARIAN EVENTS
THEYER AND HAMMOND (1974) ESTIMATED AGE (M.Y.) AND POLARITY PCSITIO?' 1.6-1.7 top Of Olduval Y 2.5-2.6 latest Gauss fl
4.5-4.6 Gilbert "C2" event L/ 6.2-6.3 Epoch 6 , above Jevent "a" 8.6-8.7 base of Epoch 8
L/
10.7-10.8
top
Of
'
Epoch 11
I/
11.3-11.4 base of Epoch 11
15.5-15.6 late Epoch 16 J D. ALATA F. 0.p Y COSTATA F.O. STICHOCORYS WOLFFII L.O. THEOCYRTIS ANNOSA F.O. C. TETRAFERA
F.O. L. ELONGATA TRISTYLOSPYRIS TRICEROS TO D. ATEUCHUS LITHOCYCLIA ARISTOTELIS GROUP TO L. ANGUSTA F.O. CARPOCANISTRUM AZYX F.O. PODOCYRTIS GOETHEANA GOETHEANA PODOCYRTIS MITRA TO P. CHALARA PODOCYRTIS SINUOSA TO P. MITRA PODOCYRTIS PHYXIS TO P. AMPLA F.O. EUSYRINGIUM LANGENA LAGENA F .O. THEOCAMPE MONGOLFIER l THEOCOTYLE CRYPTOCEPHALA NIGRINIAE TO T. CRYPTOCEPHALA CRYPTOCEPTHALA F.O. THEOCORYS ANACLASTA BURYELLA TETRADICA TO B. CLINATA F.O. BEKOMA BIDARFENSIS
1
530 b o u n d a r y , t h e s i l l d e p t h o f t h e Panamanian B l o c k would h a v e b e e n a b o u t 500 meters (Bandy and C a s e y , 1 9 7 3 ) .
Therefore,
the
i s o l a t i o n may w e l l be t w o f o l d ; r e s t r i c t e d c i r c u l a t i o n d u e t o t h e e m e r g e n c e o f t h e Panamanian B l o c k , and c o o l i n g t h a t r e s u l t e d i n t h e i n i t i a t i o n and d e v e l o p m e n t o f Neogene g l a c i a t i o n s
and w a t e r mass r e g i m e s
(Casey, 1973).
Our c u r r e n t r e s e a r c h s u g g e s t s t h a t w a t e r mass r e g i m e s and r a d i o l a r i a n faunas s i m i l a r t o t o d a y ' s w e r e e s t a b l i s h e d by midMiocene and t h a t A t l a n t i c and P a c i f i c warm-water
faunas have
b e e n " i s o l a t e d " from one a n o t h e r s i n c e a b o u t t h e b a s e o f t h e S p o n g a s t e r p e n t a s Z o n e , or a b o u t 4 . 5 m i l l i o n y e a r s a g o , o r about t h e Miocene-Pliocene boundary. and " o n e way" i s o l a t i o n .
This i s o l a t i o n is a leaky
The P t l a n t i c s t i l l ( s i n c e 4 . 5 m i l l i o n
y e a r s a g o ) r e c e i v e s n e w l y e v o l v e d warm w a t e r f o r m s from t h e Pacific-Indian
Ocean f a u n a s v i a t h e f i l t e r b r i d g e o f t h e Cape 0
o f Good Hope ( w a t e r s w i t h t e m p e r a t u r e a s h i g h a s 2 5 C come round t h e Cape p r e s e n t l y and a t warmer i n t e r v a l s i n t h e p a s t warmer f a u n a s s h o u l d h a v e m i g r a t e d i n t o t h e P t l a n t i c )
.
However
warm w a t e r r a d i o l a r i a n f a u n a s e v o l v i n g i n t h e A t l a n t i c , p o s t c l o s u r e of Panama, c a n n o t m i g r a t e i n t o t h e P a c i f i c a r o u n d t h e T i e r r a d e l Fuego. T h e r e f o r e i t a p p e a r s t h e "warm w a t e r " z o n a t i o n o f R i e d e l and S a n f i l i p p o w i l l work i n p a r t i n t h e A t l a n t i c , e s p e c i a l l y f i r s t o c c u r r e n c e s may l a g a b i t i n t i m e h a v i n g t o a w a i t e s t a b lishment i n t h e A t l a n t i c , m o s t l i k e l y being e s t a b l i s h e d .during w o r l d - w i d e warm i n t e r v a l s . The R-mode c l u s t e r o f l i v e r a d i o l a r i a n s from t h e S o u t h Texas outer continental shelf
( Figure 2 )
r a d i o l a r i a n s from t h e o t h e r s
separates the r e l i c t
( t h e y a r e n o t a s s o c i a t e d w i t h any
s e a s o n and o n l y a s s o c i a t e a t a low s i m i l a r i t y l e v e l w i t h anything).
S p o n g a s t e r p e n t a s a t t a c h e s a t a low ( a n d p r o b a b l y i n -
s i g n i f i c a n t ) l e v e l w i t h t h e w i n t e r g r o u p which i s i n t e r e s t i n g f o r i t is w i t h i n t h e winter group t h a t associates.
Spongaster c r u c i f e r u s
However S p o n g a s t e r c r u c i f e r u s a s s o c i a t e s a t a
531 " h i g h l e v e l " w i t h a few o t h e r s and a g a i n t h i s h i g h l e v e l i s d u e
t o few o c c u r r e n c e s s o t h i s may be t h r o w n o u t w i t h more s a m p l ing.
S p o n g a s t e r ? p e n t a s , and t h e " c i r c u l a r " and " e l l i p t i c a l "
s p o n g a s t e r s a l l c l u s t e r o u t t o g e t h e r between t h e s p r i n g upwelli n g (SU) and summer ( S ) r a d i o l a r i a n a s s e m b l a g e s . W e b e l i e v e t h a t t h i s "throwing out
I'
of t h e r a d i o l a r i a n sea-
s o n a l c l u s t e r g r o u p s means t h a t e i t h e r t h e r e l i c t r a d i o l a r i a n s c a n g e t a l o n g w i t h a n y g r o u p ( w h i c h would be a way t o s u r v i v e ) o r t h a t t h e y have an u n s p e c i a l i z e d n i c h e (can e a t a v a r i e t y of n a n o p h y t o p l a n k t o n or a r e d e t r i t u s f e e d e r s o r c o n t a i n s y m b i o t i c z o o x a n t h e l l a e ) and h a v e b e e n a b l e t o s u r v i v e a s t h e r e s t o f t h e p o p u l a t i o n s h a v e Lvolved
"around them".
p l a n k t o n tows
t a k e n i n t h e low l a t i t u d e A t l a n t i c i n P p r i l of 1976 r e v e a l e d t h a t t h e spongasters (Spongaster t e t r a s ,
S . p e n t a s , and t h e
" c i r c u l a r " and " e l l i p t i c a l " s p o n g a s t e r s ) a p p e a r r e d i n f r e s h l i v e unstained material.
W e suggest t h a t t h i s red colour is
d u e to t h e p r e s e n c e o f s y m b i o t i c a l g a e and s u g g e s t s a v e r y u n s p e c i a l i z e d and p e r h a p s a v e r y c o n s e r v a t i v e n i c h e . A s h a s b e e n n o t e d many r a d i o l a r i a n datum p l a n e s and zona-
t i o n s a r e e i t h e r p r o v i n c i a l , s u c h a s R i e d e l and S a n f i l i p p o ' s "warm w a t e r " C e n o z o i c , and N i g r i n i ' s e q u a t o r i a l Q u a t e r n a r y z o n a t i o n s , o r a r e t i m e t r a n s g r e s s i v e , s u c h a s h a s b e e n shown by petrushevskaya (1972). Neogene o f t h e F t l a n t i c .
This is especially t r u e f o r the l a t e Our s t u d i e s s u g g e s t t h a t a t l e a s t
some r a d i o l a r i a n s e x h i b i t f a i r l y c o n s i s t e n t d a t u m p l a n e s o f a more o r l e s s c o s m o p o l i t a n n a t u r e and h a s b e e n s u g g e s t e d b y Casey (Casey, i n p r e s s b ) . i n g p o l a r and/or
These w e r e a p p a r e n t l y shallow l i v -
t e m p e r a t e f o r m s t h a t were " e n d e m i c " t o d i v i n g
w a t e r m a s s e s and e x h i b i t t r o p i c a l s u b m e r g e n c e .
Living radio-
l a r i a n s e x h i b i t i n g t h i s type of d i s t r i b u t i o n have been desc r i b e d b y C a s e y ( 1 9 7 1 b and in press, a).
Living radiolarians
e x h i b i t i n g t h i s t y p e of d i s t r i b u t i o n i n c l u d e S p o n g o p y l e o s c u l o s a D r e y e r , and S i p h o c a m p e e r u c o s a H a e c k e l w h i c h a p p e a r t o be e n d e m i c t o t h e d i v i n g C e n t r a l W a t e r M a s s e s o f t h e P a c i f i c ,
532 C y r t o p e r a l a g u n c u l a H a e c k e l and C o r n u t e l l a p r o f u n d a E h r e n b e r g w h i c h a p p e a r t o be e n d e m i c t o t h e d i v i n g I n t e r m e d i a t e W a t e r Masses o f t h e P a c i f i c and P e r i p y r a m i s c i r c u m t e x t a H a e c k e l w h i c h a p p e a r s t o o c c u r i n b o t h C e n t r a l and I n t e r m e d i a t e W a t e r Masses
( C a s e y , 1 9 7 1 and i n p r e s s , a ) .
F o s s i l r a d i o l a r i a n s be-
l i e v e d t o h a v e e x h i b i t e d t h i s t y p e o f d i s t r i b u t i o n a r e Lamproc y c l a s h e t e r o p o r o s Haus p r o b a b l y e x h i b i t i n g an e n d e m i c p a t t e r n t o t h e d i v i n g C e n t r a l Water Masses.
? P r u n o p y l e t i t a n Campbell
and C l a r k and ?Lychnocanium g r a n d e C a m p b e l l and C l a r k a r e bel i e v e d t o b e d i v i n g I n t e r m e d i a t e W a t e r Mass f o r m s and E u c y r t i dium c a l v e r t e n s e S . S . M a r t i n a p p e a r s t o h a v e o c c u r r e d i n b o t h C e n t r a l and I n t e r m e d i a t e W a t e r M a s s e s d u e t o i t s r e l a t i v e l y common o c c u r r e n c e i n V-24-59
and i t s o c c u r r e n c e a s f a r s o u t h
a s p. t i t a n i n W e a v e r ' s c o r e s ( 1 9 7 3 ) .
T h e s e f o s s i l forms may
w e l l p r o v e t o e x h i b i t " c o s m o p o l i t a n " datum p l a n e s and a b i o s t r a t i g r a p h i c z o n a t i o n b a s e d on r a d i o l a r i a n s o f t h i s t y p e may be t h e b e s t cosmopolitan b i o s t r a t i g r a p h i c zonation possible. These f o s s i l s p e c i e s a r e c o n s i d e r e d t o e x h i b i t t h i s t r o p i c a l submergent p a t t e r n of d i s t r i b u t i o n b e c a u s e t h e i r o c c u r r e n c e i n f o s s i l sediments mirrors t h e occurrences of the l i v i n g t r o p i c a l s u b m e r g e n t forms i n r e c e n t s e d i m e n t s ( b e i n g d o m i n a n t i n high l a t i t u t d e s but r a r e i n t r o p i c a l sediments).
Their
c o n s i s t e n c y a s u s e f u l datum p l a n e s i s s u g g e s t e d b y c o m p a r i n g t h e i r e x t i n c t i o n s and e v o l u t i o n s i n h i g h
l o w ( c o r e V-24-59)
( c o r e E-14-8)
and
l a t i t u d e s with magnetostratigraphy (as
shown on F i g u r e 4 ) .
L a m p r o c y c l a s h e t e r o p o r o s and p e r h a p s
E u c r y t i d i u m c a l v e r t e n s e S.S.
l a s t o c c u r a t t h e O l d u v a i or
G i l s a e v e n t , and Lychnocanium g r a n d e and P r u n o p y l e t i t a n c o n s i s t e n t l y l a s t occur w i t h i n t h e upper Gauss; whereas Lamprocyclas h e t e r o p o r o s a p p a r e n t l y f i r s t o c c u r s i n a primit i v e form i n t h e G i l b e r t .
U s i n g t h i s same l i n e o f r e a s o n i n g
i t is possible t o suggest t h a t Theocyrtis redonodoensis Campbell and C l a r k e x h i b i t e d a b i p o l a r p a t t e r n o f d i s t r i b u t i o n i n s t e a d of a t r o p i c a l submergent p a t t e r n f o r i t a p p e a r s a t
533 h i g h and mid l z t i t u d e s ( t h e A n t a r c t i c and s o u t h e r n C a l i f o r n i a ) d u r i n g t h e l a t e Neogene b u t n o t a t low l a t i t u d e s ( s u c h a s c o r e V-24-59). T h e s e i d e a s a r e s t i l l i n an e m b r y o n i c s t a g e o f d e v e l o p m e n t and more work w i l l h a v e t o be d o n e t o c o n f i r m o r d e n y t h e i r However w e s u g g e s t t h a t t h e f u t u r e u s e of t h i s
validity.
" c o s m o p o l i t a n r a d i o l a r i a n z o n a t i o n " may be o f g r e a t h e l p i n c o r r e l a t i n g p r o v i n c i a l r a d i o l a r i a and o t h e r m i c r o f o s s i l zonations. A few s e l e c t e d e x a m p l e s o f r a d i o l a r i a n s u s e d i n t h e v a r i o u s
z o n a t i o n s m e n t i o n e d a r e i l l u s t r a t e d on p l a t e 2 a l o n g w i t h comments i n t h e legend a s t o what z o n a t i o n s t h e y a p p l y .
ACKNOWLEDGEMENTS
R e s e a r c h on R e c e n t r a d i o l a r i a n e c o l o g y and d i s t r i b u t i o n i n t h e w a t e r column and s e d i m e n t s h a s b e e n s u p p o r t e d m a i n l y b y t h e O c e a n o g r a p h i c S e c t i o n , N a t i o n a l S c i e n c e F o u n d a t i o n , NSF G r a n t DES 74-21805 and i n p a r t b y t h e Bureau o f Land Management C o n t r a c t #08550-CTS-17.
R e s e a r c h on t h e C e n o z o i c h i s t o r y
o f t h e A t l a n t i c and t h e p a l e o o c e a n o g r a p h y and b i o s t r a t i g r a p h y and e v o l u t i o n o f r a d i o l a r i a n s h a s b e e n s u p p o r t e d b y a g r a n t from t h e P e t r o l e u m R e s e a r c h Fund o f t h e P m e r i c a n C h e m i c a l S o c i e t y , PRF #8657-AC
2 t o w h i c h t h e a u t h o r s a r e most g r a t e f u l .
The a u t h o r s a l s o a c k n o w l e d g e t h e e d i t o r i a l and t y p i n g a s s i s t a n c e of B . H a w k i n s , t y p i n g a s s i s t a n c e o f J . C i s n e r o s , t h e r e search a s s i s t a n c e of M. t r a n s l a t i o n b y J.-C.
Bauer and
J.
G e v i r t z , and a b s t r a c t
D e B r e m a e c k e r , a l l o f t h e Geology D e p a r t -
ment, Rice University.
Lamont-Doherty G e o l o g i c a l O b s e r v a t o r y
and F l o r i d a S t a t e U n i v e r s i t y k i n d l y p r o v i d e d c o r e m a t e r i a l . C u r a t i n g f a c i l i t i e s a t Lamont a r e s u p p o r t e d b y N S F g r a n t GF 35454 and ONR c o n t r a c t N00012-67-~-0108-0004,
and a n NSF-
O f f i c e of P o l a r P r o g r a m s g r a n t GV-27549 t o F l o r i d a S t a t e University
.
534
REFERENCES Bandy, 0. L., and Casey, R. E . , 1973. R e f l e c t o r h o r i z o n s and p a l e o b a t h y m e t r i c h i s t o r y , e a s t e r n Panama. Geol. SOC. Amer. B u l l . , v . 8 4 , p. 3081-3086. Bandy, 0. L . , c a s e y , R. E . , and Wright, R. C., 1971. L a t e Neogene p l a n k t o n i c z o n a t i o n , m a g n e t i c r e v e r s a l s , and r a d i o m e t r i c d a t e s , A n t a r c t i c t o t h e t r o p i c s . In Reid, J. L., ( E d . ) , A n t a r c t i c Oceanography 1, A n t a r c t i c Research S e r i e s , v . 15 : Washington (American G e o p h y s i c a l U n i o n ) , p.1-26. B e r g e r , W. H . , 1968, R a d i o l a r i a n s k e l e t o n s : s o l u t i o n a t d e p t h : S c i e n c e , v . 1 5 9 , p. 1237-1238. , 1970, Biogenous deep-sea s e d i m e n t s : f r a c t i o n by deep-sea c i r c u l a t i o n . Geol. SOC. Amer. B u l l . , v . 8 1 , no. 5 , p. 1385-1402. Berggren, W. A., and Van C o w e r i n g , J. A., 1974. The L a t e Neosene, Amsterdam ( E l s e v i e r S c i e n t i f i c P u b l i s h i n g c o . ) , p . 1-216. Casey, R. E , , 1971a. D i s t r i b u t i o n of p o l y c y s t i n e r a d i o l a r i a n s i n t h e o c e a n s i n r e l a t i o n t o p h y s i c a l and c h e m i c a l c o n d i In The M i c r o p a l e o n t o l o g y of Oceans. B. M. F u n n e l 1 tions. and W. R. R i e d e l ( E d s . ) . cambridge (Cambridge Univ. Press). 151. , 1971b. R a d i o l a r i a n s a s i n d i c a t o r s o f p a s t and p r e s e n t w a t e r masses. In The M i c r o p a l e o n t o l o g y of Oceans. B. M. F u n n e l 1 and W. R. R i e d e l ( E d s . ) . Cambridge (cambridge Univ. P r e s s ) . 331. , 1973, R a d i o l a r i a n e v i d e n c e f o r t h e i n i t i a t i o n and development of neogene g l a c i a t i o n s and t h e neogene w a t e r mass regimes: ( d i s c u s s i o n p a p e r i n ) Geol. SOC. Amer. M e e t , , D a l l a s , Texas, Geol. Soc. Am. Ann. Meetinq, Dallas, 1973, A b s t r a c t s , p. 570-571. , i n p r e s s . a , The e c o l o g y and d i s t r i b u t i o n of recent Radiolaria. g Ramsay, A.T.S. (Ed.) Oceanic micropaleontology. , i n p r e s s . b . L a t e Neogene r a d i o l a r i a n b i o s t r a t i y raphy r e l a t e d t o m a g n e t o s t r a t i g r a p h y , P o l a r t o T r o p i c s : 0. L. Bandy Memorial Volume. C i f e l l i , R., and S a c h s , K. N . , 1966. Abundance r e l a t i o n s h i p s o f p l a n k t o n i c F o r a m i n i f e r a and R a d i o l a r i a . Deep-sea R e s . , V. 1 3 , p. 751-753, Edmund, M. and Anderson,G. C . , 1970. On t h e s t r u c t u r e of North A t l a n t i c Deep Water: Deep-sea R e s . , v . 18, p . 127138 Fanning, K O A., and Schink,D. R., 1969. I n t e r a c t i o n Of marine s e d i m e n t s w i t h d i s s o l v e d s i l i c a : Limnol. and Oceanogr., V . 1 4 , p . 59-68. G o l l , R. M. and B j d r k l u n d , K O R., 1971. R a d i o l a r i a i n s u r f a c e s e d i m e n t s o f t h e North A t l a n t i c Ocean, M i c r o p a l e o n t . 1 7 , ( 4 ) , 434-457.
-
J.
535
Goll, R. M. and Bjdrklund, K. R., 1974. Radiolaria in surface sediments of the South Atlantic; Micropaleontology, v. 20, no. 1, p . 38-75. Grant, A. B., 1968. Atlas of oceanographic sections, Davis Strait-Labrador Basin-Denmark Strait-Newfoundland Basin: 1965-1967; Atlantic Ocean Lab, Bedford Inst. Rept., AOL v. 68, no. 5, 80 p . Harper, H, E., Jr. and Knoll, A. H., 1975. Silica, diatoms, and Cenozoic radiolarian evolution. Geology, vol. 3, no. 4,
175-177.
Hays., J. D., 1965. Radiolaria and late Tertiary and Quaternary history of Antarctic Seas. Biology of Antarctic Seas 11. Am. Geophys. Union, Antarctic Research Ser. 5, 125. Hays, J. D. and Opdyke, N. D., 1967, Antarctic Radiolaria, Magnetic Reversals, and Climatic change. Science 158, (3804),
1001.
Johnson, T. C., 1974. The dissolution of siliceous microfossils in surface sediments of the eastern tropical Pacific. Deep-sea Res., v. 21, p . 851-864. Johnson, D. A., and Knoll, A. H., 1975. Absolute ages of Quaternary radiolarian datum levels in the equatorial Pacific. Quaternary Research 5, p . 99-110. Kennett, J. P., Burns, R. E., Andrews, J.E., Churkin, H. Jr., Davis, T o A., Dumitrica, P., Edwards, A. R., Galehouse, J.S., Packham, G. H,, and van der Lingen, G o J., 1972, Australian-Antarctic Continental Drift, Palaeocirculation Changes and Oligocene Deep Sea Erosion. Nature, vol. 239, p. 51-55.
Kobayashi, T o , 1944. Reciprocal development of radiolarian rocks as between Asiatic and Australian sides. Proc. Imp. Acad. Tokyo, vol. 21, no. 4, p . 234-238. McMillen, K. J., 1975. Quaternary lebensspuren and their relationship to depositional environments in the Caribbean Sea, the Gulf of Mexico, and the eastern and central North Pacific, M. A. thesis, Rice University, Houston. , 1976. Ecology, distribution and preservation of Polycystine Radiolaria in the Gulf of Mexico and caribbean Sea, Ph. D. thesis, Rice University, Houston. Nigrini, C. A., 1971, Radiolarian zones in the Quaternary of the Equatorial Pacific Ocean. In Funnell, B. M. and Reidd W. R. (Eds.) The Micropaleontology of Oceans: cambridge (Cambridge Univ. Press), p . 443-461. Petrushevskaya, M. G o , 1971. Spumellarian and nassellarian Radiolaria in the plankton and bottom sediments of the Central Pacific. In The Micropalaeontology of Oceans (B.M. Funnell and W.R. Riedel - Eds.) Cambridge University Cambridge, 309-317. , 1972. Biostratigraphy of deep-water Quaternary deposits from radiolarian analysis. Oceanology, v. 12, no. 1, 57-70. Riedel, W.R. and Sanfilippo, A., 1971. Radiolaria. In
.
536
Winterer, E. L., Riedel, W. R., et al. Initial Reports of the Deep Sea Drilling Project, Volume 7: Washington ( U . S . Government Printing Office), p. 1529. Riedel, W, R, and Sanfilippo, A., in press, Cenozoic Radiolaria. In Ramsay, A.T.S, (Ed.) Oceanic micropalaeontology. Ryan, W.B.F., Cita, M.B., Dreyfus Rawson, M., Burckle, L. H., and Saito, T . 1974. A paleomagnetic assignment of Neogene stage boundaries and the development of isochronous datum planes between the Mediterranean, the Pacific and Indian Oceans in order to investigate the response of the World Ocean to the Mediterranean "salinity crisis". Riv. Ital. Paleont., vol. 80, pp. 631-688. Sanfilippo, A., Burckle, L.H,, Martini, E., and Riedel, W.R,, 1973, Radiolarians, diatoms, silicoflagellates and calcareous nannofossils in the Mediterranean Neogene: Micropaleontology, v. 19, no. 2, p. 209-234. Tbeyer, F., and Hanunond, S.R., 1974, Paleomagnetic polarity sequence and radiolarian zones, Brunhes to polarity epoch 20: Earth Planet. Sci. Lett., v. 22, p . 307-319. Weaver, F.M., 1973, Pliocene paleoclimatic and paleoglacial history of East Antarctica recorded in deep sea piston cores: Sedimentology Research Laboratory, Department of Geology, Florida State Univ., Contrib. 36.
537
-
-I-
:
:;;. -
S
-
V- 2 4 - 5 9
E::
v = 2
c
B
A;
W
,100
U
e L 3 L
1
'
F Collorphaerc tuberora
2 ***
ci
L A n t h o c y r t i d i u m angulare
%-
I500
L P t e r o c a n i u m p r i r m a t i u m *-• & Lamprocyclar h e t e r o p o r o r & Eucyrtidium calrertense & Cyrtopera S.S.
L?Prunopyle t i t a n + +
F -
+ * *.
'J
6?Lychnocanium grande 6 Stichoeoryr peregrina & P t e r o c o r y r rplendenc
I
Y
-
c v)
=
& O m m i t a r t u s p e n u l t i m u s .**
.lo00 & C y r t t c a p r e t l a t e t r a p e r i a c c ,
am I
=:
L Spongaster p e n t a r
r= 4
3
I-
z z
laguncula
A
hmgoplegmr antirct icui t warm water forms
-- _ _ - -
I1
- w a r m water forms ------_---_____
L'
A c a n t h o r p h a e r a sp.
3
0
-
d
r:
I Buccinorphaerr inraginata
n
:I
-
-
'Z
nd-
ANTARCTIC CORE E- 1 4 - 8
4 I Spangarter tetras 6 Ommatartus t e t r a t h a1 a Onmatartur'hugheri 6 IP t eprroi sc m a na itri nu r n I . S .
& Lamprocyclar h e t e r o po r o s
0 v)
.
L P t e r o c a n i u m t rilobum
L Eucyrtidium c a l r t r t e n ~ I F Cyrtopera Iagunculi Oermospyrir s pongiora 6 Helotholur rema Prunopyle t i tan 6 L y c h n o c a n i u m grand# 6 P t e r o c o r y r rplendenr
Ororcena ( d i g i t a t e l 6 0. c a r o l a e Cyrtocapsella tetrapera (L T h e o c y r t i r redondoensir
v)
L
R
Y
E
g r o v
a -
z z
=;"
F i g . 4. R a d i o l a r i a n b i o s t r a t i y r a p h y .
T r i c e r a r p y r i s rp.
L Ommatocampe h u g h c s i & C a n a r t i s c u r marylan
538
PLATE 1 R a d i o l a r i a n s i n d i c a t i v e of s p e c i f i c w a t e r masses from t h e Cari b b e a n S e a , Gulf of Mexico, and A t l a n t i c Ocean. The b l a c k b a r e q u a l s a b o u t 1 0 0 m i c r o n s . The l e t t e r p i n d i c a t e s specimens from p l a n k t o n t o w s , t h e l e t t e r s i n d i c a t e s i n d i v i d u a l s from sediment samples. Figures 1 t o 5 S u r f a c e Water Figure 1 Pterocorys zancleus (Muller) s; Figure 2 Pterocanium p r a e t e x t u m p r a e t e x t u m ( E h r e n b e r g ) s ; F i g u r e 3 Sponga s t e r t e t r a s t e t r a s Ehrenberg s ; F i g u r e 4 Choenicosphaera murrayana Haeckel p w i t h p r o t o p l a s m ; F i g u r e 5 E u c h i t o n i a e l e g a n s (Ehrenberg ) s
-
---
S u r f a c e Water, r e l i c t f a u n a - F i g u r e s 6 t o 7 F i g u r e 6 " C i r c u l a r " s p o n g o d i s c i d s; F i g u r e 7 S p o n g a s t e r p e n t a s R i e d e l and S a n f i l i p p o s S u b t r o p i c a l Underwater - F i g u r e s 8 t o 1 2 F i g u r e 8 C l a t h r o c a n i u m diadema s; F i g u r e 9 L i t h e l i u s minor J o r g e n s o n m S on o t r o c h u s g l a c i a l i s Popofsky s ; F i g u r e 11 Amphirhopalum*on Haeckel s ; F i g u r e 1 2 9lochlamidium a s t e r i s c u s Haeckel p
*-
A n t a r c t i c I n t e r m e d i a t e Water - F i g u r e s 1 3 t o 1 7 F i g u r e 1 3 Actinomma s p . s ; F i g u r e 1 4 C u b o t h o l u s c . f . b i c u s Haeckel s; F i g u r e 1 5 Carpocanium evacuatum Haeckel s ; F i g u r e 1 6 T h o l o n i d p ; F i g u r e 1 7 Dictyophimus c r i s i a e Ehrenberg s
-
N o r t h A t l a n t i c Deep Water Figures 18 t o 20 F i g u r e 18 Spongopyle o s c u l o s a Dreyer s ; F i g u r e 1 9 P t e r o c o r y s b i c o r n i s Popofsky s ; F i g u r e 2 0 C l a t h r o c o r y s s p . p PLATE 2
R a d i o l a r i a n s used a s datums i n R i e d e l and S a n f i l i p p o s "warm w a t e r " Cenozoic P a c i f i c Z o n a t i o n ( F i g u r e s 1, 2 , 3 , 4 , 5 and 8 ) , N i g r i n i ' s Quaternary E q u a t o r i a l P a c i f i c Zonation ( F i g u r e s 6 and 7 ) and C a s e y ' s Developing T r o p i c a l Submergent Cosmopolitan Zonation ( F i g u r e s 9 through 1 5 ) . F i g u r e 1 P t e r o c a n i u m p r i s m a t i u m R i e d e l , Core V-24-59, 1111 c m . F i r s t o c c u r r e n c e datum f o r b a s e of S p o n g a s t e r p e n t a s Zone of R i e d e l and S a n f i l i p p o ( i n p r e s s ) F i g u r e 2 Ommatartus? h u g h e s i (Campbell and C l a r k ) , Core V-24-59, 1 1 9 1 c m . F i g u r e 3 Ommatartus h u g h e s i (Campbell and C l a r k ) , Core E14-8, 1750-1752 c m . L a s t o c c u r r e n c e u s e d i n R i e d e l and S a n f i l i p p o ( i n p r e s s ) a s datum f o r b a s e of Ommatartus penu l t i m u s Zone. F i g u r e 4 Ommatartus e n u l t i m u s ( R i e d e l ) , Core V-24-59, 1 2 4 1 cm. m i h o w e r p o r t i o n of t h e Ommatart u s p e n u l t i m u s Zone o f R i e d e l and S a n f i l i p p o ( i n p r e s s ) F i g u r e 5 C y r t o c a p s e l l a t e t r a p e r a H a e c k e l , Core E-14-8, 1750-1752 c m . F i r s t o c c u r r e n c e d e f i n e s t h e bottom of t h e new C y r t o c a p s e l l a t e t r a p e r a Zone of R i e d e l and S a n f i l i p p o
-
-
539
( i n press) F i g u r e 6 Amphirhopalum y p s i l o n H a e c k e l , Core V-24-59, ? c m . T h i s form of t h e s p e c i e s i s i n d i c a t i v e of t h e Amphirhopalum y p s i l o n Assemblage Zone (Zone 3 ) of N i g r i n i ( 1 9 7 1 ) F i g u r e 7 C o l l o s p h a e r a t u b e r o s a H a e c k e l , Core V - 2 4 - 5 9 , ? cm. D e f i n e s t h e bottom of C o l l o s p h a e r a t u b e r o s a C o n c u r r e n t Range Zone (Zone 2 ) of N i g r i n i ( 1 9 7 1 ) F i g u r e 8 S t i c h o c o r y s p e r e g r i n a ( R i e d e l ) , Core V-24-59, 1008 c m . E v o l u t i o n a r y bottom (more t h a n 50% of 5. p e r e g r i n a when compared t o a n c e s t o r 2. d e l m o n t e n s e ) d e f i n e s t h e b a s e of t h e S t i c h o c o r y s p e r e g r i n a Zone ( R i e d e l and S a n f i l i p p o , i n p r e s s ) , l a s t o c c u r r e n c e d e f i n e s bottom of P t e r o c a n i u m p r i s m a t i u m Zone ( R i e d e l and S a n f i l i p p o , i n p r e s s ) . Appears t o be u s e f u l o v e r wide l a t i t u d i n a l a r e a . F i g u r e 9 E u c y r t i d i u m c a l v e r t e n s e M a r t i n , Core #-14-8, 17501752 crn. L a s t o c c u r r e n c e a t P l i o c e n e - P l e i s t o c e n e boundary i n A n t a r c t i c (Hays, 1965) (Bandy g 1 9 7 1 ) . A probably t r o p i c a l submergent " c o s m o p o l i t a n " s p e c i e s (Casey , i n p r e s s b) F i g u r e 1 0 T h e o c y r t i s r e d o n d o e n s i s Campbell and C l a r k , Core E-14-8, 1728-1730 c m . Appears t o e x h i b i t a b i p o l a r r a t h e r t h a n t r o p i c a l submergent d i s t r i b u t i o n a l p a t t e r n ( C a s e y , i n press b) F i g u r e 11 ? P r u n o p y l e t i t a n Campbell and C l a r k , Core E.-14-8, 809 c m . Found i n t h e t r o p i c s , c o n s i d e r e d a t r o p i c a l subm e r g e n t f o r m , i t s l a s t o c c u r r e n c e i s i n d i c a t i v e of u p p e r Gauss m a g n e t i c t i m e o v e r l a r g e g e o g r a p h i c a l a r e a (Casey, i n press b) F i g u r e 1 2 P r u n o p y l e t i t a n Campbell and C l a r k , E x p e r i m e n t a l Mohole, EM-8-15, 242-245 c m . The h i g h e r l a t i t u d e " s u b s p e c i e s " of t h e low l a t i t u d e t r o p i c a l submergent " s u b s p e c i e s " i l l u s t r a t e d i n F i g u r e 11 of t h i s p l a t e . F i g u r e 1 3 Oroscena w i t h d i g i t a t e s p i n e s , Core E-14-8, 17501752 c m . May b e t r o p i c a l submergent " c o s m o p o l i t a n " s p e c i e s whose l a s t o c c u r r e n c e i s i n d i c a t i v e of t h e u p p e r G i l b e r t . May b e good f o r " r e d c l a y " b i o s t r a t i g r a p h y . F i g u r e 1 4 Lychnocanium g r a n d e Campbell and C l a r k , Core E 14-8, 1750-1752 c m . C o n s i d e r e d t r o p i c a l submergent form commonly l a s t o c c u r r i n g w i t h P r u n o p y l e t i t a n i n t h e u p p e r Gauss. F i g u r e 1 5 Larnprocylas h e t e r o p o r o s Hays, Core V - 2 4 - 5 9 , 809 c m . C o n s i d e r e d t r o p i c a l submergent whose r a n g e i s a p p a r e n t l y t h e r a n g e of t h e P l i o c e n e .
e.
540
PLA1-E 1
541
PLATE 2
542
Discussion D . Habib: When you s a y t h e c o l d - w a t e r f o r m s d i v e , a r e t h e y diving a l i v e o r a r e t h e tests being transported? Casey: Y e s , t h e y ' r e d i v i n g a l i v e i These organisms w i l l t a k e t h e p r o t o p l a s m i c s t a i n i n p r e s e r v e d p l a n k t o n tows from t h e s e d e e p e r w a t e r s . NOW, of c o u r s e t h e s e w a t e r m a s s e s h a v e t a k e n h u n d r e d s of y e a r s t o g e t from o n e p l a c e t o a n o t h e r , s o i t ' s t h e i r g r e a t g r e a t g r e a t g r a n d f a t h e r t h a t was i n s h a l l o w water. D r . I . G . Sohn: What a n i m a l s e a t r a d i o l a r i a n s ? Casey: W e l l t h e y ' r e n o t t r e m e n d o u s l y a b u n d a n t i n t h e w a t e r column, t h e r a d i o l a r i a n s , so I would t h i n k t h a t a n y k i n d of a f i l t e r f e e d e r would b e i m p o r t a n t i n e a t i n g them. You do f i n d them i n f e c a l p e l l e t s of c o p e p o d s and t h a t s o r t o f thing. I t appears t h a t r a d i o l a r i a n s a r e important i n t e r m e d i a r i e s i n t h e o p e n o c e a n f o o d web b e t w e e n n a n n o p l a n k t o n and o t h e r m i c r o p l a n k t o n and m e s o p l a n k t o n . Sohn: The r e a s o n f o r t h e q u e s t i o n i s y o u r s t a t e m e n t t h a t when you p u t r a d i o l a r i a n s i n a c i d and t a k e them o u t t h e y d i s s o l v e . S e v e r a l y e a r s ago I c o l l a b o r a t e d w i t h K o r n i c k e r i n a s t u d y where w e f e d l i v e o s t r a c o d e s t o f i s h and some o f t h e v a l v e s d e v e l o p e d h o l e s . Could d i g e s t i v e a c i d remove a p r o t e c t i v e coating thus causing t h e r a d i o l a r i a n t o dissolve? C a s e y : T h i s m i g h t be: w e ' v e c o n s i d e r e d t h i s . One of t h e t h i n g s t h a t ' s i n t e r e s t i n g a b o u t many o f t h e c o p e p o d s , o n e of t h e main f e e d e r s on r a d i o l a r i a n s , i s t h a t i f t h e y g e t a c h a n c e t h e y ' l l a c t a s g l u t t o n s , s a y when t h e y ' r e g r a z i n g o n d i a t o m s , and t h e f e c a l p e l l e t s w i l l a c t u a l l y h a v e t h e p r o t o p l a s m s t i l l i n t h e d i a t o m s , e v e n t h e c h l o r o p h y l l , so t h e y ' l l j u s t g o r i g h t t h r o u g h . So a l t h o u g h many of them may b e d i s s o l v e d i n t h i s m a n n e r , o r b e e t c h e d and r e a d y t o d i s s o l v e , p r o b a b l y many w i l l come t h r o u g h . We t h i n k , t h e r e f o r e , t h a t i n g e n e r a l b e i n g e a t e n by s o m e t h i n g would b e a n a i d , b e c a u s e y o u ' v e g o t a s a n d - s i z e d p a r t i c l e now, and i t s g o i n g t o f a l l t h r o u g h t h e w a t e r i n a c o u p l e d a y s . Based o n o u r s e t t l i n g r a t e s t u d i e s o n t h e r a d i o l a r i a n s w e know t h a t i n average oceanic depths t h e average r a d i o l a r i a n f a l l i n g ind i v i d u a l l y t h r o u g h t h e w a t e r column t a k e s a b o u t a y e a r t o So i t ' s t h e c o u p l e d a y s compared t o a y e a r h i t bottom. t h a t would h e l p o u t . D r . W. A . Berggren: Do you b e l i e v e f e c a l p e l l e t s e d i m e n t a t i o n i s i m p o r t a n t i n forming r a d i o l a r i a n oozes? Casey: Y e s , i n g e n e r a l w e t h i n k t h a t t h e r a d i o l a r i a n s are b e t t e r p r e s e r v e d u n d e r p l a c e s where t h e y a r e e a t e n and t h e n r e a c h t h e bottom i n f e c a l p e l l e t s . I n o t h e r words, f e c a l p e l l e t s e d i m e n t a t i o n i s o n e of t h e main ways w e t h i n k w e can g e t r a d i o l a r i a n occurrences i n the sediments. In areas where t h e r a d i o l a r i a n s j u s t d i e i n d i v i d u a l l y and f a l l t h r o u g h t h e w a t e r column t h o s e a r e t h e p l a c e s w h e r e t h e y w i l l u s u a l l y j u s t d i s s o l v e away. But t h e whole q u e s t i o n of r a d i o l a r i a n s e d i m e n t a t i o n i s o n e weak s p o t i n r a d i o l a r i a n r e s e a r c h . W e may see v e r y f r a g i l e c o l l o s p h a e r i d s i n t h e b o t t o m s e d i m e n t s and e v e n i n s e d i m e n t s where o t l - e r r a d i o l a r i a n s h a v e d i s s o l ved o u t . I n t h e Gulf of Mexico and C a r i b b e a n i n g e n e r a l r a d i o l a r i a n s o c c u r i n t h e u p p e r 1 0 c m and t h e n t h e y ' r e j u s t gone a s though t h e r e ' s p e r h a p s a d i s s o l u t i o n h o r i z o n r i s i n g t h r o u g h t h e s e d i m e n t . I t may b e t h a t 1 0 c m i s t h e d e p t h o f Dr.
543
a c t i v e b u r r o w i n g by b e n t h i c o r g a n i s m s . I i m a g i n e t h e d i s a p p e a r a n c e of y o u r s p e c i e s SpongBerggren: a s t e r p e n t a s i s no d o u b t a s s o c i a t e d w i t h t h e e l e v a t i o m t h e I s t h m u s of Panama, b u t when you s a y y o u ' r e n o t s u r e a b o u t how t o d a t e i t I t h i n k you c a n d a t e i t p r e t t y a c c u r a t e l y . You c a n d o i t w i t h F o r a m i n i f e r a . Casey: About 3 . 5 m i l l i o n y e a r s a g o . B e r g g r e n : Y e s , b u t t h e r e a s o n you c a n d o i t i s t h e r e a r e cert a i n t a x a which a p p e a r i n b o t h o c e a n s which e v o l v e d a b o u t 4 m i l l i o n y e a r s a g o . T h e r e ' s a n o t h e r t a x o n which evolved a r o u n d 3 . 5 m i l l i o n y e a r s a g o and i t ' s p r e s e n t l y i n t h e A t l a n t i c b u t n e v e r o c c u r s i n t h e P a c i f i c , s o i t had t o happen The g e n u s P u l l e n i a t i n a which somewhere between t h a t t i m e . h a s l i t t l e l e f t and r i g h t c o i l i n g w i g g l e s i n t h e A t l a n t i c and P a c i f i c t h r o u g h t h e P l i o c e n e d i s a p p e a r s i n t h e A t l a n t i c a t e x a c t l y 3 . 5 m i l l i o n y e a r s ago. I t ' s no l o n g e r p r e s e n t b u t i t c o n t i n u e s w i g g l i n g l e f t and r i g h t i n t h e P a c i f i c r i g h t up t o t h e p r e s e n t d a y . I t reappears i n t h e A t l a n t i c a t 2.3 m i l l i o n y e a r s and w e d o n ' t know why t h a t happens b u t it p r o b a b l y came a r o u n d t h e s o u t h p a r t of t h e Agulhas C u r r e n t a g a i n somehow o r o t h e r . But I t h i n k y o u r S p o n g a s t e r p e n t a s Zone t h e u p p e r l i m i t of which i s , w h a t , 4 m i l l i o n y e a r s from t h e m a g n e t i c s you h a v e o n a s c a l e t h e r e ? C a s e y : Died o u t b e f o r e t h e end o f t h e Zone. B e r g g r e n : Y e s , it i s p r o b a b l y r e l a t e d i n some way t o t h e c h a n g i n g g e o m e t r y o f t h e I s t h m u s of Panama b u t t h a t e v e n t can be dated p r e t t y accurately. I t h i n k s o , b u t I was c o n c e r n e d a b o u t s u g g e s t i n g t h a t Casey: a t 3 . 5 m i l l i o n y e a r s a g o a n a n i m a l c o u l d walk a c r o s s t h e I s t h m u s . The e l e v a t i n g I s t h m u s h a s t o b e a lumpy s o r t of t h i n g , and t h e r e may h a v e b e e n a n e f f e c t i v e s i l l t h a t s t o p ed t h e c o n n e c t i o n of r a d i o l a r i a n p o p u l a t i o n s ; p e r h a p s s t o p ed i t f o r o n e g r o u p e a r l i e r t h a n i t d i d f o r a n o t h e r ( a s h a l l o w - l i v i n g group v s . a d e e p - l i v i n g g r o u p ) . D r . S . G a r t n e r : I'll a d d r e s s t h i s p o i n t b o t h t o you Dick and t o B i l l B e r g g r e n , and l e t m e s a y f i r s t o f a l l I talce e x c e p t t i o n t o y o u r v i e w w i t h r e s p e c t t o t h e I s t h m u s of Panama. About s i x o r s e v e n y e a r s a g o w e d i d a s t u d y of B l a k e P l a t e a u s e d i m e n t s and e s t a b l i s h e d t h a t t h e s e d i m e n t a r y r e g i m e p r e v a i l i n g t h e r e a t t h e p r e s e n t t i m e , which i s l a r g e l y cont r o l l e d by t h e Gulf S t r e a m , was e s t a b l i s h e d s o m e t h i n g i n e x c e s s of 5 m i l l i o n y e a r s a g o . T h i s would i n d i c a t e t h a t t h e The Gulf S t r e a m would h a v e had i t s p r e s e n t c o n f i g u r a t i o n . Gulf S t r e a m would b e d e p e n d e n t on t h e I s t h m u s of Panama, and i f t h e l a t t e r d i d n o t come i n t o e x i s t e n c e u n t i l 3 . 5 m i l l i o n y e a r s a g o , n e c e s s a r i l y t h e Gulf S t r e a m would h a v e b e e n d i f ferent a t that time. But t h e Gulf S t r e a m d i d e s t a b l i s h i t s p r e s e n t c o n f i g u r a t i o n a t l e a s t o v e r t h e B l a k e P l a t e a u i n exc e s s o f 5 m i l l i o n y e a r s a g o . Now t h e e v i d e n c e t h a t you have p r e s e n t e d abd t h a t B i l l p r e s e n t e d i s s u g g e s t i v e t h a t i t m i g h t h a v e happened s a y 3 . 5 m i l l i o n y e a r s a g o a t p r e s u m a b l y about t h e t i m e t h a t Antarctic g l a c i a t i o n occurred. I t seems t o m e t h a t y o u ' r e n o t s e p a r a t i n g t h e s e two f a c t o r s . It c o u l d b e t h a t what you a r e l o o k i n g a t i s t h e i n f l u e n c e of A n t a r c t i c g l a c i a t i o n a l l by i t s e l f b e i n g f e l t 3 . 5 m i l l i o n y e a r s a g o 3nd t h a t t h e I s t h m u s d i d i n f a c t come i n t o e x i s t e n c e som t i m e b e f o r e t h a t . Now p e r h a p s I m i s u n d e r s t o o d e i t h e r o r b o t h of you. Do you h a v e any p o s i t i v e e v i d e n c e t h a t s u g g e s t s t h a t t h e I s t h u s had t o b e i n e x i s t e n c e o r
...
544
c o u l d have n o t been i n e x i s t e n c e u n t i l 3 . 5 m i l l i o n y e a r s ago? Casey: Ln a p a p e r t h a t Bandy and I p u b l i s h e d i n 1 9 7 3 , m a i n l y on b e n t h o n i c f o r a m s , t h e e l e v a t i o n of i t i s shown. What I ' m t h i n k i n g i s t h a t a l l of t h e s e c a n b e c o r r e l a t e d w i t h t h e e l e v a t i o n of t h e I s t h m u s d u r i n g t h e Miocene, s a y a b o u t t h e t i m e t h a t t h e main e l e v a t i o n t o o k p l a c e . I think t h a t various t h i n g s happened b e c a u s e of t h e e l e v a t i o n : t h e s h u t t i n g o f f of t h e f a u n a s a s it r e a c h e d e f f e c t i v e s i l l d e p t h , e f f e c t i v e sills for faunal isolation. I t m i g h t have a l s o r e a c h e d some e f f e c t i v e s i l l d e p t h e a r l i e r t o d i v e r t and t o i n t e n s i f y t h e Gulf Stream b e c a u s e t h e l a t t e r i s a v e r y b i g and d e e p s o r t of t h i n g . So i t may b e t h a t i t was w i a e enough and d e e p enough so t h a t a m a j o r body of w a t e r was d i v e r t e d n o r t h , bef o r e t h e shallow water r a d i o l a r i a n i s o l a t i o n s took place. I d o n ' t t h i n k t h e y a l l have t o happen a t t h e same t i m e . The s o r t of sewuence I would l i k e t o see i s t h a t i t comes u p , i t s t a r t s t o d i v e r t t h e Gulf Stream and t h e n t h i s i n t e n s i f i e s , w i t h more e l e v a t i o n , t h e warm w a t e r pushed n o r t h c a u s i n g c o o l i n g i n Ewing-Donn s e n s e , and r a d i o l a r i a n p a l e o t e m p e r a t u r e c u r v e s t h a t I ' v e done s u g g e s t t h a t i t d o e s g e t c o l d a t s a y 4 m i l l i o n y e a r s ago o r t h e r e a b o u t s . But t h e n you g e t i n t o P l e i s t o c e n e g l a c i a t i o n s and t h e y ' r e e v e n c o l d e r : and t h a t m i g h t be due t o e v e n a more c o m p l e t e i s o l a t i o n and d e f l e c t i o n o f w a t e r by t h e c o m p l e t e emergence of Panama. Land f a u n a s c o u l d move between N o r t h and S o u t h America by hopping p r i o r t o t h i s p e r h a p s . I t d o e s n ' t h a v e t o be a c o m p l e t e and t o t a l emergence a t a n i n s t a n t . I t h i n k it came up and t h e n I t h i n k t h e p o i n t s you b r i n g up a r e i m p o r t a n t o n e s . Then y o u ' v e g o t t o s t a r t wondering a b o u t e u s t a t i c r i s e s and f a l l s of s e a l e v e l , e t c .
545
ATLANTIC C E N O Z O I C SILICOFLAGELLATES, POTENTIAL FOR BIOSTRATIGRAPHIC AND PALEOECOLOGIC STUDIES Richard E . Casey, R i c e U n i v e r s i t y , Houston, Texas
F BSTRACT S i l i c o f l a g e l l a t e s have been used only occasionally i n t h e A t l a n t i c r e g i o n m a i n l y d u e t o p o o r p r e s e r v a t i o n i n much o f t h i s r e g i o n . However, s i l i c o f l a g e l l a t e s d o p o s s e s s p o t e n t i a l f o r b i o s t r a t i g r a p h i c and p a l e o e c o l o g i c s t u d i e s i n t h e A t l a n tic. R ~ S U M ~
L e s s i l i c o f l a g e l l 6 s n ' o n t 6 t 6 u t i l i s g s que rarement dans l a r 6 g i o n a t l a n t i g u e , principalement 2 cause d e l e u r mgdiocre & t a t de conservation dans c e t t e rggion. Nganmoins l e s s i l i c o f l a g e l l g s p o s s s d e n t un p o t e n t i e l ' a l ' u t i l i s a t i o n p o u r d e s Qtudes biostratigraphiques e t palgogcologiques dans 1'Atlant ique. INTRODUCTION
S i l i c o € l a g e l l a t e s e v o l v e d i n t h e C r e t a c e o u s and h a v e b e e n r e p r e s e n t e d b y a b o u t 12 g e n e r a and 50 s p e c i e s b e l o n g i n g t o two f a m i l i e s .
About 6 t o 12 s p e c i e s and t h e i r v a r i e t i e s ( p e r -
h a p s amounting t o 60 e n t i t i e s ) b e l o n g i n g t o a b o u t 3 t o 6 genera
( m a i n l y t h e g e n e r a D i c t y o c h a , D i s t e p h a n u s and Mesocena)
and o n e f a m i l y l i v e i n t h e p r e s e n t - d a y s e a s .
Silicoflagel-
l a t e s h a v e b e e n u s e d i n b i o s t r a t i g r a p h i c and p a l e o t e m p e r a t u r e s t u d i e s m a i n l y i n t h e P a c i f i c and A n t a r c t i c .
Silicoflagel-
l a t e s t u d i e s i n t h e A t l a n t i c and i t s m a r g i n s h a v e b e e n few mainly b e c a u s e of t h e poor p r e s e r v a t i o n of t h e s e f o s s i l s i n most o f t h e A t l a n t i c t h r o u g h o u t m o s t o f t h e C e n o z o i c .
Silico-
f l a g e l l a t e s a r e u s u a l l y o n l y common i n r o c k s and s e d i m e n t s c o n t a i n i n g l a r g e amounts o f b i o g e n i c s i l i c a ( L i p p s , 1 9 7 0 ) and t h e r e f o r e a r e s u b j e c t t o some of t h e same p r e s e r v a t i o n p r o b -
l e m s a s a r e t h e r a d i o l a r i a n s i n t h e P t l a n t i c ( s e e C a s e y and
546
McMillen i n t h i s v o l u m e ) .
The r e l a t i v e l y low d i v e r s i t y of
s i l i c o f l a g e l l a t e s h a s and w i l l l i m i t t h e i r u s e f u l n e s s i n b i o s t r a t i g r a p h i c and p a l e o e c o l o g i c s t u d i e s : h o w e v e r , t h e f o l l o w ing s e c t i o n s d e a l with t h e i r p o t e n t i a l f o r such u s e
(especially
f o r u s e i n t h e A t l a n t i c ) , and w i t h a s h o r t r e v i e w of i m p o r t a n t silicoflagellate literature.
SILICOFLAGELLATE
BIOSTRATIGRAPHY
AND T H E I R POTENTIAL
USE I N THE A T L A N T I C
Hanna
(1928) suggested t h a t s i l i c o f l a g e l l a t e s were excel-
lent biostratigraphic indices. 25 years
However, o n l y w i t h i n t h e l a s t
( a n d e s p e c i a l l y s i n c e t h e s t a r t of t h e Deep S e a D r i l l -
ing P r o j e c t ) h a s t h i s group been s e r i o u s l y considered f o r biostratigraphic correlation.
Mandra
(1960) r e c o g n i z e d s i l i c o -
f l a g e l l a t e c o m p l e x e s c h a r a c t e r i s t i c o f L a t e E o c e n e , Miocene and Pliocene ages using s t a t i s t i c a l a n a l y s i s .
Papp (1959) n o t e d
t h a t s i l i c o f l a g e l l a t e s can b e e a s i l y used t o d i s t i n g u i s h Paleog e n e from Neogene d e p o s i t s . The p r e s e n c e of t h e s i l i c o f l a g e l l a t e g e n u s Lyramula a p p e a r s t o b e i n d i c a t i v e o f u p p e r C r e t a c e o u s and d o e s n o t p e n e t r a t e i n t o t h e Cenozoic.
The g e n e r a C o r b i s e m a , N a v i c u l o p s i s and
P a r a d i c t y o c h a a p p e a r t o b e i n d i c a t i v e of t h e P a l e o g e n e , wherea s t h e g e n e r a C a n n o p i l u s and R o c e l l a a p p e a r t o b e r e s t r i c t e d t o t h e Miocene a l o n g w i t h 5 o t h e r Miocene g e n e r a
(Loeblich et
a l , 1 9 6 8 ) w i t h t h e l a t e Neogene b e i n g d o m i n a t e d b y t h e g e n e r a D i c t y o c h a , Mesocena and D i s t e p h a n u s . M a r t i n i (1971) set u p t h e f i r s t s i l i c o f l a g e l l a t e zonation d e r i v e d from c o n t i n u o u s s e q u e n c e s . and S w e d i s h Deep-sea Bukry ( 1 9 7 5 ) 'I
T h e s e w e r e from DSDP s i t e s
c o r e 7 6 from t h e e q u a t o r i a l P a c i f i c .
i n a l l u d i n g t o the s i l i c o f l a g e l l a t e zonation
ph i 10soph i e s " s t a t e s t h a t
'I
s i 1i c o f 1a g e 11a t e zon a t i o n , un 1i k e
t h e o n e employed f o r c o c c o l i t h s t h a t r e l i e s e x c l u s i v e l y on o c c u r r e n c e , r e q u i r e s q u a n t i t a t i v e d a t a and more f l e x i b l e
547
definition.
Bukry c o n t i n u e s s t a t i n g " c o n s i s t e n c y and abun-
'I
d a n c e o f s p e c i e s a r e g e n e r a l l y more s i g n i f i c a n t i n s i l i c o f l a g e l l a t e z o n a t i o n t h a n a r e a b s o l u t e f i r s t and l a s t o c c u r r e n c e s
--
t h e a p p a r e n t s k e l e t a l p l a s t i c i t y of s i l i c o f l a g e l l a t e s p e c i e s
r e f l e c t i n g high ecological responsiveness,
also contributes t o
a n e e d f o r l o w e r e d r e l i a n c e on f i r s t o r l a s t o c c u r r e n c e s , i n f a v o r o f f u l l - a s s e m b l a g e a n a l y s i s and c o s m o p o l i t a n - t r e n d determination
--
p a r t o f t h i s n e c e s s a r y c a u t i o n r e s u l t s from
t h e present lack of m u l t i p l e successions preventing determinat i o n o f t h e r e g i o n a l v e r s u s c o s m o p o l i t a n c h a r a c t e r o f assemblages.
I'
Of s p e c i a l i n t e r e s t i n Bukry
(1975)
i s h i s comparison of
t r o p i c a l , c o s m o p o l i t a n and n o n t r o p i c a l s i l i c o f l a g e l l a t e zonations.
From t h i s work and a r a p i d s u r v e y o f o t h e r s i l i c o -
f l a g e l l a t e b i o s t r a t i g r a p h i c w o r k s a c o s m o p o l i t a n z o n a t i o n app e a r s t o f u n c t i o n w e l l f o r t h e P a l e o g e n e t o mid-Miocene. mid-Miocene
Post
t h e s i l i c o f l a g e l l a t e f l o r a s and b i o s t r a t i g r a p h y
d i v e r g e i n t o t r o p i c a l and n o n t r o p i c a l c o m p o n e n t s .
It i s sug-
g e s t e d t h a t t h i s d i v e r g e n c e i s c a u s e d b y t h e same f a c t o r s i n v o l v e d i n t h e s i m i l a r d i v e r g e n c e i n r a d i o l a r i a n f a u n a s and biostratigraphy
at
t h e same t i m e
( r e f e r t o paper b y Casey
and McMillen i n t h i s volume f o r d e t a i l e d d i s c u s s i o n and t o t h e i r f i g u r e on t e c t o n i c and o c e a n i c e v e n t s and t h e i r e f f e c t s on C e n o z o i c s i l i c e o u s s e d i m e n t a t i o n and r a d i o l a r i a n s i n t h e Ftlantic).
I t is t h e r e f o r e s u g g e s t e d t h a t t h e s i l i c o f l a g e l -
l a t e z o n a t i o n o f Bukry ( 1 9 7 5 )
may be u s e d i n most o f t h e
A t l a n t i c f o r t h e P a l e o g e n e and e a r l y Neogene b u t t h a t t h e s i l i c o f l a g e l l a t e f l o r a s o f t h e A t l a n t i c i n t h e mid and l a t e Neogene e x h i b i t p r o v i n c i a l i s m ( a s d o t h e f l o r a s r e f e r r e d t o d u r i n g t h i s t i m e i n Bukry ( 1 9 7 5 1 ,
and t h e A t l a n t i c r a d i o l a r i -
an f a u n a s d i s c u s s e d i n t h e c h a p t e r b y C a s e y and McMillen i n t h i s volume).
M a r t i n i ( S a n f i l i p p o e t a l , 1973) attempted t o
z o n e p o r t i o n s o f t h e M e d i t e r r a n e a n Neogene u s i n g h i s Neogene s i l i c o f l a g e l l a t e zonation established i n t h e equatorial Pacific
548 (Martini, 1971).
M a r t i n i was o n l y a b l e t o p l a c e h i s s a m p l e s o f
M e d i t e r r a n e a n i n l a t e Neogene t e n t a t i v e l y b u t h i s s a m p l e s from mid-Miocene
( a b o u t t h e D o r c a d o s p y r i s a l a t a l o n e o f R i e d e l and
S a n f i l i p p o , i n p r e s s ) and o l d e r w e r e zoned w i t h a p p a r e n t e a s e . H e r e t h e n i t might b e p o s s i b l e t o s u g g e s t t h a t t h e development o f
a
d e t a i l e d warm w a t e r P t l a n t i c Neogene s i l i c o f l a g e l l a t e zona-
t i o n i s i n o r d e r i f enough m a t e r i a l i n c o n t i n u o u s enough s e c t i o n s from mid and l o w l a t i t u d e s c a n b e f o u n d .
SILICOFLFGELLATE PFLEOECOLOGIC STUDIES FND T H E I R POTENTIAL USE I N THE FTLANTIC
Colom ( 1 9 5 2 ) a t t e m p t e d t o r e c o n s t r u c t p a l e o o c e a n o g r a p h i c condition i n Aquitanian-Burdigalian deposits i n Spain.
More
r e c e n t l y a t t e m p t s h a v e b e e n made a t u s i n g s i l i c o f l a g e l l a t e s t o determine paleotemperatures.
Mandra and Mandra
(1969) des-
c r i b e d a t e c h n i q u e u s i n g t h e r a t i o s of D i c t y o c h a (warm) t o Distephanus (cold) t o determine paleotemperatures i n t h e T e r t i a r y of t h e Antarctic.
Weaver and C i e s i e l s k i
(1974) suggest
c o o l i n g s o f 8 t o 10°C i n t h e F n t a r c t i c P l i o c e n e .
E a r l i e r work
on t h i s i n t e r v a l from c o r e s i n t h e S o u t h e r n Ocean u s i n g r a t i o s 0
o f r a d i o l a r i a n s s u g g e s t e d a d r o p of a b o u t 1 5 C (Bandy e t a l . , 1971).
A p p a r e n t l y s i l i c o f l a g e l l a t e s c a n be u s e d f o r p a l e o -
t e m p e r a t u r e a n a l y s i s i n warm w a t e r s a l s o a s h a s b e e n d o n e b y M a r t i n i ( 1 9 7 1 ) f o r t h e Neogene o f t h e e q u a t o r i a l P a c i f i c . S i l i c o f l a g e l l a t e s h a v e n o t b e e n r e l a t e d t o w a t e r mass d i s t r i b u t i o n s a s h a v e many o f t h e o t h e r s h e l l e d n a n n o p l a n k t o n i c and m i c r o p l a n k t o n i c g r o u p s , b u t a r e v i e w o f t h e d i s t r i b u t i o n s o f r e c e n t and f o s s i l s p e c i e s p u b l i s h e d b y G l e z e r ( 1 9 7 0 ) s u g g e s t s t h a t t h e s e s u r f a c e l i v i n g forms e x h i b i t b o t h c o s m o p o l i t a n and p r o v i n c i a l d i s t r i b u t i o n a l p a t t e r n s .
For e x a m p l e D i s t e p h -
anus speculum (Ehrenberg) a p p e a r s t o occur i n a l l oceans b u t becomes r a r e n e a r s h o r e w h e r e s a l i n i t i e s a p p r o a c h 1 0 p p t .
In
c o n t r a s t D i s t e p h a n u s f i b u l a E h r e n b e r g a p p e a r s t o be a b l e t o
549
e x i s t a t t h e s e lower s a l i n i t i e s ( G l e z e r , 1 9 7 0 ) .
Therefore
s i l i c o f l a g e l l a t e s may be u s e f u l p a l e o o c e a n o g r a p h i c i n d i c e s not only f o r paleotemperature but paleowater-mass, paleosalini t y and o t h e r s t u d i e s a s w e l l .
IMPORTANT SILICOFLAGELLATE LITERATURE W I T H NOTES ON CONTENTS
M o n o g r a p h i c w o r k s on s i l i c o f l a g e l l a t e s i n c l u d e :
Glezer ' s
( 1 9 7 0 ) d e a l i n g w i t h b i o l o g y , s y s t e m a t i c s , d i s t r i b u t i o n and e c o l o g y and b i o s t r a t i g r a p h i c and g e o l o g i c o c c u r r e n c e s , a must f o r s i l i c o f l a g e l l a t e w o r k : L o e b l i c h ' s e t a l . ( 1 9 6 8 ) w h i c h i s an a n n o t a t e d i n d e x o f f o s s i l and r e c e n t s i l i c o f l a g e l l a t e s and e b r i d i a n s w i t h d e s c r i p t i o n s and i l l u s t r a t i o n s a n o t h e r m u s t ; and L i n g ' s ( 1 9 7 2 ) work on u p p e r C r e t a c e o u s and C e n o z o i c s i l i c o f l a g e l l a t e s and e b r i d i a n s . Some i m p o r t a n t r e c e n t p u b l i c a t i o n s c o n c e r n e d w i t h s i l i c o f l a g e l l a t e z o n a t e s i n c l u d e : Mandra
( 1 9 5 1 , 1954 and 1 9 6 0 ) on
C a l i f o r n i a m a t e r i a l ; S t r a d n e r ( 1 9 6 1 ) on O l i g o c e n e s i l i c o f l a g e l l a t e s from A u s t r i a ; Bachmann, P a p p and S t r a d n e r ( 1 9 6 3 ) on T e r t i a r y s i l i c o f l a g e l l a t e s o f t h e V i e n n a b a s i n ; Bachmann and I c h i k a w a ( 1 9 6 2 ) on t h e s i l i c o f l a g e l l a t e s o f t h e Neogene of Japan;
Zhuze ( 1 9 4 9 , 1 9 5 1 , 1 9 5 5 ) on Upper C r e t a c e o u s and
P a l e o g e n e o f t h e e a s t e r n s l o p e of t h e U r a l s and t h e West S i b e r i a n p l a i n : G l e z e r ( 1 9 7 0 ) ; and e s p e c i a l l y t h e w o r k s publ i s h e d i n t h e Deep S e a D r i l l i n g P r o j e c t t o o numerous t o ment i o n h e r e except f o r t h e paper by Dumitrica
( 1 9 7 2 ) on t h e
Mediterranean. Some i m p o r t a n t r e c e n t p u b l i c a t i o n s c o n c e r n e d w i t h s i l i c o f l a g e l l a t e e c o l o g y , p a l e o e c o l o g y and p a l e o o c e a n o g r a p h y i n c l u d e : Mandra and Mandra ( 1 9 6 9 ) , J e n d r z e j e w s k i and Z , a r i l l o ( 1 9 7 1 ) , Martini (1971), Ciesielski
and Weaver
( 1 9 7 3 ) and Weaver and
C i e s i e l s k i ( 1 9 7 3 and 1 9 7 4 ) on s i l i c o f l a g e l l a t e p a l e o t e m p e r a tures.
G l e z e r ( 1 9 7 0 ) g i v e s c o n s i d e r a b l e d e t a i l on t h e d i s t r i -
b u t i o n and e c o l o g y o f l i v i n g s p e c i e s and r e f e r s t o many
550
v a l u a b l e r e f e r e n c e s on t h e s e a s p e c t s . Bukry i s t h e most p r o l i f i c s i l i c o f l a g e l l a t e b i o s t r a t i g r a p h -
e r w i d e l y p u b l i s h e d i n t h e Deep S e a D r i l l i n g P r o j e c t v o l u m e s . I n h i s DSDP Leg 34 p a p e r
(Bukry, 1976) h e g i v e s a b r i e f b u t
e x c e l l e n t discussion of s i l i c o f l a g e l l a t e s t r a t i g r a p h y , paleoe c o l o g y and t e r m i n o l o g y o f s i l i c o f l a g e l l a t e m o r p h o l o g y p l u s some i m p o r t a n t r e f e r e n c e s n o t g i v e n h e r e .
ACKNOWLEDGEMENTS R e s e a r c h o n th.Es p a p e r has b e e n s u p p o r t e d By t h e O c e a n o g r a p h i c S e c t i o n , N a t i o n a l S c i e n c e F o u n d a t i o n , NSF G r a n t DES 74-21805 and b y a g r a n t from t h e P e t r o l e u m R e s e a r c h Fund o f t h e P m e r i c a n C h e m i c a l S o c i e t y , PRF #8657-PC grateful.
2 t o which t h e a u t h o r
is
most
The a u t h o r s a l s o a c k n o w l e d g e t h e e d i t o r i a l and t y p -
i n g a s s i s t a n c e o f B . Hawkins,
t y p i n g a s s i s t a n c e of J. C i s n e r o s ,
and a b s t r a c t t r a n s l a t i o n b y J . C . DeBremaecker,
a l l of t h e
Geology D e p a r t m e n t , R i c e U n i v e r s i t y .
REFERENCES Bachmann, F . and I c h i k a w a , W . , 1 9 6 2 . The s i l i c o f l a g e l l i d e s i n t h e Wakura B e d s , Nanao C i t y , P r e f e c t u r e I s h i k a w a , J a p a n : Kanazawa U n i v . S c i . R e p t . , v 8 , p . 1 6 1 . Bachmann, A., P a p p , A., and S k r . a d n e r , H . , 1963. M i k r o p a l a o n t o l o g i s c h e S t u d i e n i m " B a d e n e r T e g e l " von F r a t t i n g s d o r f N . O . M i t t . g e o l . G e s . W i e n , v . 5 6 , n o . 1, p . 117-210. Bandy, 0 . L . , C a s e y , R . E . , and W r i g h t , R. C . , 1 9 7 1 . L a t e N e o g e n e p l a n k t o n i c z o n a t i o n , m a g n e t i c r e v e r s a l s , and r a d i o metric dates, Antarctic t o the tropics. R e i d , J. L . , ( E d . ) , A n t a r c t i c O c e a n o g r a p h y 1, F n t a r c t i c R e s e a r c h S e r i e s , v. 1 5 : W a s h i n g t o n (American G e o p h y s i c a l U n i o n ) , p. 1-26. B u k r y , D., 1 9 7 5 . S i l i c o f l a g e l l a t e and c o c c o l i t h s t r a t i g r a p h y , Deep S e a D r i l l i n g P r o j e c t Leg 29. Kennett, J. P . , H o u t z , R . E . , et a l . , I n i t i a l R e p o r t s o f t h e Deep S e a D r i l l i n g P r o j e c t , Volume 2 9 : W a s h i n g t o n (U.S. Government P r i n t i n g O f f i c e ) , p. 845-872. _ , 1 9 7 6 . S i l i c o f l a g e l l a t e and c o c c o l i t h s t r a t i g r a p h y , s o u t h e a s t e r n p a c i f i c Ocean, Deep S e a D r i l l i n g P r o j e c t Leg 34. In Y e a t , R . S . , H a r t , S. R., et &., I n i t i a l R e p o r t s of t h e Deep S e a D r i l l i n g P r o j e c t , Volume 3 4 : W a s h i n g t o n (U.S. Government P r i n t i n g O f f i c e ) , p . 715-735.
=
551 C i e s i e l s k i , P . F. and W e a v e r , F . M., 1 9 7 3 . S o u t h e r n Ocean P l i o c e n e p a l e o t e m p e r a t u r e s b a s e d on s i l i c o f l a g e l l a t e s from d e e p - s e a c o r e s : A n t a r c t i c J . U . S . , v. 8 , n o . 5 , p. 295297. Colom, G . , 1 9 5 2 . F q u i t a n - b u r d i g a l i e n Diatom D e p o s i t s o f t h e North B e t i c S t r a i t . J . P a l e o n t . , v . 2 6 , n o . 6 , p. 867-885. D u m i t r i c a , P . , 1 9 7 2 . Miocene and Q u a t e r n a r y s i l i c o f l a g e l l a t e s i n s e d i m e n t s from t h e M e d i d i t e r r a n e a n S e a . In Ryan, W . B . F . , H S U , K. J . g &., I n i t i a l R e p o r t s o f t h e Deep S e a D r i l l i n g P r o j e c t , Volume 1 3 : W a s h i n g t o n ( U . S . Government P r i n t i n g O f f i c e ) , p . 902. G l e z e r , 2 . I . , 1 9 6 6 . Silicoflagellatophyceae. In G o l l e r b a k h , M. M. ( E d . ) , C r y p t o g a m i c p l a n t s o f t h e U.S.S.R.: Fkad. Nauk SSSR, V. A. Komarova B o t . I n s t . ( T r a n s l a t e d from Russ i a n b y I s r a e l Program f o r S c i e n t i f i c T r a n s l a t i o n s L t d . , J e r u s a l e m , 1 9 7 0 ) , v . 7 , p. 1-363. H a n n a , G . D . 1 9 2 8 , S i l i c o f l a g e l l a t a e from C r e t a c e o u s o f C a l i p a l e o n t . , v o l . 1, n o . 4 , p. 259-264. fornia, P . and Z a r i l l o , G . A . , 1 9 7 1 . Late PleistoJendrzejewski, c e n e p a l e o t e m p e r a t u r e s : S i l i c o f l a g e l l a t e s and f o r a m i n i f e r a l frequency changes i n a S u b a n t a r c t i c deep-sea c o r e : F n t a r c t i c J. U.S., v . 6 , p . 178-179. L i n g , H . Y., 1 9 7 2 . Upper C r e t a c e o u s and C e n o z o i c s i l i c o f l a g e l l a t e s and e b r i d i a n s : A m . P a l e o n t o l . B u l l . v . 6 2 , p . 1 3 5 229. L i p p s , J. H . , 1 9 7 0 . E c o l o g y and e v o l u t i o n o f s i l i c o f l a g e l lates. P r o c . N o r t h Fmer. P a l e o . Conv. S e p t . 1 9 6 9 . PFRTG. p . 965-993. L o e b l i c h , F . R . , 1 1 1 , L o e b l i c h , L . A . , T a p p a n , H . , and L o e b l i c h , F . R . , J r . , 1 9 6 8 . A n n o t a t e d i n d e x o f f o s s i l and R e c e n t s i l i c o f l a g e l l a t e s and e b r i d i a n s w i t h d e s c r i p t i o n s and ill u s t r a t i o n s of v a l i d l y proposed t a x a : Geol. Soc. Am., M e m . 1 0 6 , 319 p. Mandra, Y. T . , 1 9 5 1 . P r e l i m i n a r y s t r a t i g r a p h i c r e p o r t on some C a l i f o r n i a Eocene S i l i c o f l a g e l l a t e s . B u l l . G e o l . S O C . Am., P a r t 2 , v . 6 2 , n o . 1 2 , p. 1 5 2 3 . , 1 9 5 4 . S i l i c o f l a g e l l a t a , a new t o o l f o r t h e Geologist. B u l l . G e O l . S O C . Am., P a r t 2 , v . 6 5 , no. 1 2 , p. 1396. ~ o s s i ls i l i c o f l a g e l l a t e s from C a l i f o r n i a , - ~ _ _ ,_ 1 9 6 0 . U.S.F. r e p t . o f T w e n t y - F i r s t S e s s i o n I n t e r n . G e o l . Cong r e s s , p a r t 6 , Proc. Sect. 6 , Pre-Quatern. Micropaleontol., p . 77-89. Mandra, Y . T . and Mandra, H . , 1 9 6 9 . S i l i c o f l a g e l l a t e s : F new t o o l f o r t h e s t u d y of F n t a r c t i c T e r t i a r y c l i m a t e s : F n t a r c t i c J . U.S., v . 4 , p. 172-174. M a r t i n i , E . , 1 9 7 1 . Neogene s i l i c o f l a g e l l a t e s from t h e e q u a t o r i a l Pacific. I n W i n t e r e r , E . L . , R i e d e l , W. R . , e t a l . , I n i t i a l R e p o r t s o f t h e Deep S e a D r i l l i n g P r o j e c t Volume VII: W a s h i n g t o n (U.S. Government P r i n t i n g O f f i c e ) , p. 1695-1708.
J.
J.
552
Papp, A., 1 9 5 9 . Handbuch der stratigraphischen Geologie. vol. 3 no. 1, p. 3 2 8 . Riedel, W. R. and Sanfilippo, .A., in press. Cenozoic Radiolaria. 111 Ramsay, A.T.S. (Ed.), Oceanic micropalaeontology Sanfilippo, A , , B u c k l e , L. H., Martini, E., and Riedel, W. R., 1973. Radiolarians, diatoms, silicoflagellates and calcareous nannofossils in the Mediterranean Neogene: Micropaleontology, v. 1 9 , p. 2 0 9 . Stradner, H., 1 9 6 1 . Uber fossile Silicoflagelliden und die Moglich-keit ihrer Verwendung in der Erdolstratigraphie: Erdbl und Kohle, v. 1 4 , no. 2 . , p. 8 7 - 9 2 . Weaver, F. M., and Ciesielski, P. F., 1 9 7 3 . Pliocene paleoclimatic history recorded in antarctic deep sea cores. Programs: Geological Society of America, Abstracts& w
-
P. 8 5 6 - 8 5 7 .
, 1 9 7 4 . Pliocene paleotemperatures and regional correlations, southern ocean. Ant. Jour. of the United States, v. 9, no. 5, p. 2 5 1 - 2 5 3 . Zhuze, A. P., 1 9 4 9 . New Diatoms and Silicoflagellates from Upper Cretaceous Argillaceous Sands of the Basin of bol' shoi Aktai River (Eastern Slope of the Northern Urals). -Botanicheskie Materialy Otdela Sporovykh Rastenii Botanicheskogo Instituta AN SSSR, 6 ( 1 / 6 ) : 6 5 - 7 8 , Moskva-Leningrad. (in Russian) , 1 9 5 1 . Diatoms and Silicoflagellates of Upper Cretaceous of the Northern Urals. -- Botanicheskie Materialy Otdela Sporovykh Rastenii Botanicheskogo Instituta, AN SSSR, Vol. 7 : 4 2 - 6 5 , Moskva - Leninqrad. (in Russian) , 1 9 5 5 . Silicoflagellates of the Paleogene. -Botanicheskie Materialy Otdela Sporovykh Rastenii Bota nicheskogo Instituta, AN SSSR, Vol. 10: 7 7 - 8 1 . Moskva Leningrad. (in Russian)
553
N o r t h American M i c r o t e k t i t e s , R a d i o l a r i a n E x t i n c t i o n s and t h e Age o f t h e Eocene-Oligocene Boundary B . P . Glass and M i c h a e l J , Z w a r t , Geology D e p a r t m e n t , Univ-
e r s i t y o f D e l a w a r e , Newark, DE 19711 Abstract N o r t h American m i c r o t e k t i t e s have b e e n f o u n d i n one G u l f o f Mexico and two C a r i b b e a n c o r e s . The m i c r o t e k t i t e l a y e r o c c u r s i n u p p e r Eocene s e d i m e n t and c o i n c i d e s w i t h t h e a p p a r e n t e x t i n c t i o n o f f i v e r a d i o l a r i a n s p e c i e s , Based on f i s s i o n t r a c k d a t i n g of t h e m i c r o t e k t i t e s and K - A r and f i s s i o n - t r a c k d a t i n g of N o r t h American t e k t i t e s , t h e N o r t h American m i c r o t e k t i t e s have a n a b s o l i j t e a g e o f - 3 4 m . y . , i n d i c a t i n g a n a g e of l e s s t h a n 35 m a y . f o r t h e Eocene-Oligocene b o u n d a r y .
R&sumk L e s m i c r o t e c h t i t e s d e l ' A m & r i q u e du Nord o n t & t &
t r o u v k e s d a n s un c o e u r du G o l f e d u Mexique e t d a n s deux du CaraPbe.
L a c o u c h e m i c r o t e c h t i t e s e t r o u v e d a n s les d&phots
d e l'hoc'ene s u p i r i e u r e e t c d i n c i d e avec l ' e x t i n c t i o n d e c i n q espkces r a d i o l a r i e n n e s
.
BasLs s u r f i s s i o n - t r a c k d a t i n g , d e s
m i c r o t e c h t i t e s , e t les K-Ar
e t fission-track dating des
t e c h t i t e s d e l ' A m & r i q u e du Nord, l e s m i c r o t e c h t i t e s d e 1'AmLrique du Nord o n t un s g e a b s o l u e d e 3 4 m . a . , ce q u i i n d i q u e u n ^age d e moins d e 3 5 m . a .
pour l a l i m i t e ioc'ene-
oligockne.
Introduction T e k t i t e s a r e s m a l l , g e n e r a l l y 2-4 cm d i a m e t e r , j e t b l a c k
t o t r a n s l u c e n t g r e e n g l a s s b o d i e s found a t s e v e r a l l o c a l i t i e s , r e f e r r e d t o as s t r e w n f i e l d s , on t h e e a r t h ' s s u r f a c e . T e k t i t e s a r e similar t o obsidian, but can be distinguished from t e r r e s t r i a l i g n e o u s g l a s s e s by t h e i r c h e m i s t r y ( t e k t i t e s have h i g h e r MgO and l e s s Na2O and H20 t h a n t e r r e s t r i a l i g n e o u s g l a s s e s w i t h s i m i l a r S i 0 2 c o n t e n t s ) and g e n e r a l l a c k o f c r y s t a l l i n e inclusions. There a r e f o u r g e n e r a l l y a c c e p t e d t e k t i t e s t r e w n f i e l d s : A u s t r a l a s i a n ( A u s t r a l i a , I n d o c h i n a and N o r t h e r n
554
Philippines), Ivory Coast of Africa, Czechoslovakian and North American (Texas and Georgia). The tektites from these strewnfields have ages of approximately 0 , 7 , 1.1, 14.7 and 35 m.y., respectively (O'Keefe, 1976). Microscopic tektites (
555
c o r r e l a t i o n t h a t aided us i n finding t h e m i c r o t e k t i t e l a y e r i n t h e two DSDP c o r e s , The p u r p o s e o f t h i s p a p e r i s t o d i s c u s s t h e s t r a t i g r a p h i c and a b s o l u t e a g e of t h e N o r t h American m i c r o t e k t i t e l a y e r and t o d i s c u s s t h e i m p l i c a t i o n s of t h i s d a t a c o n c e r n i n g t h e a g e of t h e Eocene-Oligocene boundary. Core Loca-cions and D e s c r i p t i o n s P i s t o n c o r e RC9-58 i s from t h e Venezuelan B a s i n (140 33.4' N. L a t . , 70° 48.61W. L o n g . ) . I t i s a p p r o x i m a t e l y
490 cm l o n g and i s composed of s i l i c e o u s and c a l c a r e o u s l u t i t e . It c o n t a i n s , i n a d d i t i o n t o R a d i o l a r i a and c a l c a r e o u s n a n n o f o s s i l s , n e g l i g i b l e amounts of b e n t h i c c a l c a r e o u s and a r e n a c e o u s F o r a m i n i f e r a , b a d l y c o r r o d e d p l a n k t o n i c Foramini f e r a and f i s h t e e t h , R a d i o l a r i a a r e w e l l - p r e s e r v e d and p r e dominant i n t h e c o a r s e f r a c t i o n g r e a t e r t h a n 38 u m . Interm i t t e n t , c o n s p i c u o u s a s h f a l l s , b l u r r e d by burrow m o t t l i n g , o c c u r a t - 1 6 0 , 310 and 470 cm. E x c e p t f o r a s l i g h t d a r k e n i n g e f f e c t a t t h e a s h l e v e l s , t h e s e d i m e n t i s homogeneous p a l e o r a n g e ( 10YR8/2) t h r o u g h o u t , S i t e 94 of t h e DSDP i s l o c a t e d on t h e c o n t i n e n t a l s l o p e of t h e Yucatan p l a t f o r m ( 2 4 0 31.64.l N . L a t . , 880 2 8 . 1 6 i W . L o n g . ) . Core 15, b a r r e l s 3 and 4 , c o n t a i n s t h e m i c r o t e k t i t e l a y e r . T h i s s e c t i o n o f t h e c o r e i s d e s c r i b e d as a s t r o n g l y burrowed s o f t F o r a m i n i f e r a - r i c h n a n n o f o s s i l c h a l k o f g r e e n i s h w h i t e (5G9/1) c o l o r , c o n t a i n i n g v o l c a n i c a s h (Worzel e t a l . ,
1973)
I
S i t e 149 i s l o c a t e d i n t h e c e n t r a l Venezuelan B a s i n
(15O 0 6 . 2 5 ' N . L a t . , 69'
21.85
W . Long.) n o t f a r from where
c o r e RC9-58 was t a k e n . Core 31 c o n t a i n s t h e m i c r o t e k t i t e l a y e r and i s d e s c r i b e d as a crumbly s e m i - i n d u r a t e d r i c h l y c a l c a r e o u s r a d i o l a r i a n ooze of g r a y i s h o r a n g e (10YR7/4) t o d a r k y e l l o w i s h o r a n g e (10YR6/6) c o l o r ( E d g a r e t a l . , 1 9 7 3 ) . S p a r s e v o l c a n i c g l a s s and p l a g i o c l a s e were f o u n d t h r o u g h o u t , a l o n g w i t h d i s s e m i n a t e d pumice p e b b l e s up t o 1 . 5 cm i n d i a m e t e r . A few m i c r o t e k t i t e s were found i n t h e washed r e s i d u e of t h e c o r e c a t c h e r ( D o n n e l l y and Chao, 1 9 7 3 ) . The M i c r o t e k t i t e Layer The m i c r o t e k t i t e s i n c o r e RC9-58 o c c u r i n a s i n g l e w e l l -
5 56
d e f i n e d l a y e r w i t h o v e r 90% of t h e r e c o v e r e d m i c r o t e k t i t e s (-6000 w i t h d i a m e t e r s g r e a t e r t h a n 125 a m ) coming f r o m a 304.0 cm t h i c k zone a t a d e p t h o f w 2 5 O cm i n t h e c o r e ( F i g . 2 ) . Nany o f t h e a r e n a c e o u s F o r a m i n i f e r a r e c o v e r e d from t h i s c o r e c o n t a i n e d m i c r o t e k t i t e s i n t h e i r t e s t s ( B a k e r and Glass,
1974). A t s i t e 149 a few m i c r o t e k t i t e s were found a t t h e b o t t o m o f c o r e 30, b u t most were r e c o v e r e d f r o m c o r e 31, w i t h t h e
peak abundance o c c u r r i n g a t t h e t o p of t h e c o r e . U n f o r t u n a t e l y t h e DSDP i n i t i a l r e p o r t shows a g a p o f - 7 m e t e r s b e tween t h e two c o r e s ; t h e a c t u a l l e n g t h of m i s s i n g s e d i m e n t i s n o t known. M i c r o t e k t i t e s o c c u r i n c o r e 1 5 , s e c t i o n s 3 and 4, o f s i t e 94. The peak abundance i s a t - 4 1 6 . 1 8 m subbottom d e p t n i n s e c t i o n 3 , b u t a s m a l l e r peak i s a t 417.64 m subbottom d e p t h i n s e c t i o n 4 ( F i g . 3 ) . The d o u b l e peak i n m i c r o t e k t i t e abundance c a n n o t b e e x p l a i n e d , I f i t documents two s e p a r a t e e v e n t s , b o t h would b e e x p e c t e d t o o c c u r i n t h e o t h e r c o r e s , where o n l y one peak i s o b s e r v e d . I t i s s u s p e c t e d t h a t t h e d o u b l e peak i s a n a r t i f a c t o f t h e d r i l l i n g method u s e d , S t r a t i g r a p h i c Age of t h e M i c r o t e k t i t e Layer E x c e p t f o r some m i x i n g o f Eocene and Recent s e d i m e n t s a t t h e v e r y t o p , c o r e RC9-58 i s e n t i r e l y Eocene i n a g e (L. Burc k l e , p e r s o n a l c o m m u n i c a t i o n ) . The m i c r o t e k t i t e - r i c h l a y e r was i d e n t i f i e d a s m i d d l e Upper Eocene i n a g e b a s e d on Radiol a r i a ( R . Goll, p e r s o n a l c o m m u n i c a t i o n ) . F l o r e n t i n M a u r r a s s e , of t h e F l o r i d a I n t e r n a t i o n a l U n i v e r s i t y , worked o u t a d e t a i l e d r a d i o l a r i a n s t r a t i g r a p h y for t h i s c o r e , M a u r r a s s e a l s o a s s i g n e d a l a t e s t Eocene a g e t o t h e m i c r o t e k t i t e l a y e r b a s e d on R a d i o l a r i a ( M a u r r a s s e and G l a s s , i n p r e s s ) . C o r r e l a t i o n o f t h e m i c r o t e k t i t e d a t a w i t h t h e r a d i o l a r i a n s t r a t i g r a p h y shows t h a t t h e peak abundance o f t h e m i c r o t e k t i t e s o c c u r s a t a l e v e l i n t h e c o r e where a t l e a s t four s p e c i e s o f R a d i o l a r i a , T h y r s o c y r t i s bromia E h r e n b e r g , 1873, 2. t e t r a c a n t h a ( E h r e n b e r g ) R i e d e l and Sanf i l i p - o o , 1970, 2. t r i a c a n t h a ( E h r e n b e r g ) R i e d e l and S a n f i l i p p o , 1 9 7 0 , and C a l o c y c l a s t u r r i s E h r e n b e r g , 1873 become s u d d e n l y e x t i n c t o r e r r a t i c ( F i g . 2 ) ( M a u r r a s s e and Glass, i n p r e s s ) ,
The m i c r o t e k t i t e l a y e r i n t h e c o r e s
557
from DSDP sites 94 and 149 also appears to coincide with the extinction or sharp decrease in abundance of these four species and a fifth species, Thyrsocyrth finalis Ehrenberg, 1873 (Zwart, unpublished Master’s Thesis) (Figs. 3 & 4). Core 15 of site 94, containing the microtektite layer(s), was assigned a late Eocene age in the Initial Reports of the Deep Sea Drilling Project, V o l . X (Worzel et al., 1973). Core 14 is also assigned a late Eocene age, Core 13, which ends-29 m above the major microtektite layer, is assigned an early Oligocene age. Although based on widely spaced samples, in many cases, the section of core containing the microtektite layer is assigned to the Isthmolithus recurvus or Sphenolithus pseudoradians calcareous nannofossil Zone (Hay, 1973), the Discoaster barbadiensis Zone (Bukry, 1973) , the Thysocurtis bromia radiolarian Zone (Foreman, 1973; Sanfilippo and Riedel, 1973), the Globiaerapsis semiinvoluta or Globorotalia xroazulensis foraminiferal Zone (NcNeely, 1973),and the Pl5 foraminiferal Zone (Worzel et al., 1973). The microtektite layer at the top of core 31, DSDP site 149, also occurs in sediment identified as late Eocene in the Initial Reports of the Deep Sea Drilling Prolect, V o l . XV (Edgar et al., 1973). Core 30, directly above core 31, was assigned a middle Oligocene age (Edgar et al., 1973). Core 31 is assigned t o the Discoaster saipanensis ( ? ) Zone (Hay and Beaudry, 1973), and the Thyrsocyrtis bromia and/or Podocyrtis goetheana radiolarian Zone (Riedel and Sanfilippo, 1973) Thus, based on studies by numerous investigators involving Radiolaria, Foraminifera and nannofossils, the microtektite layer in all three cores occurs in sediments of late Eocene age. The biostratigraphic age of the rtorth American microtektite layer is, theref ore, well established. I
Absolute Age of the North American Microtektites The microtektites from core RC9-58 have been dated using the fission-track method by D. Storzer and G. A. Wagner of the Max-Planck-Institut in Heidelberg, Germany, They obtained an age, after correcting for annealing effects, of 34.6 F 4.2 m.y. (Glass et al., 1973) The large error is
558
a t t r i b u t e d t o bad c o u n t i n g s t a t i s t i c s due t o t h e small s u r f a c e a r e a a v a i l a b l e f o r c o u n t i n g . However, t h e m i c r o t e k t i t e s i n c o r e RC9-58 and t h e two DSDP c o r e s a r e p a r t of t h e N o r t h Ameri c a n t e k t i t e s t r e w n f i e l d as i n d i c a t e d by t h e i r a p p e a r a n c e , g e o g r a p h i c a l l o c a t i o n , s t r a t i g r a p h i c and a b s o l u t e a g e , p e t r o g r a p h y , p h y s i c a l p r o p e r t i e s and m a j o r element c h e m i s t r y (Glass e t a l . , 1973; Z w a r t , u n p u b l i s h e d Master's T h e s i s ) . F u r t h e r , t h e N o r t h American t e k t i t e s have b e e n d a t e d b y b o t h f i s s i o n - t r a c k and K - A r methods, Ten K - A r a g e s for N o r t h American t e k t i t e s o b t a i n e d b y Z a h r i n g e r (1963) and G e n t n e r e t a l . (1969) have a n a v e r a g e o f 34.2 m,y. w i t h a s t a n d a r d d e v i a t i o n o f 0.48. Seven published f i s s i o n - t r a c k a g e s ( F l e i s c h e r and P r i c e , 1964; F l e i s c h e r d., 1965; S t o r z e r and Wagner, 1971; G a r l i c k e t a l . , 1971; S t o r z e r e t a l . , 1973) have a n a v e r a g e o f 34.6 m.y. w i t h a s t a n d a r d d e v i a t i o n o f 0 . 4 8 , C o n s i d e r i n g a l l o f t h e above d a t a we would a s s i g n a n a g e of 34.4 2 0 . 5 m a y . f o r t h e a g e of t h e N o r t h American t e k t i t e s and m i c r o t e k t i t e s . Thus, t h e m i c r o t e k t i t e l a y e r provides a well-dated chronostratigraphic layer, Age of t h e Eocene-Oligocene Boundary Evernden and Evernden ( 1 9 7 0 ) i n d i c a t e d a n a g e of a b o u t
36-37 m . y. for t h e Ducheunian-Chardomian boundary which i s e q u i v a l e n t t o t h e Eocene-Oligocene b o u n d a r y . B e r g g r e n ( 1 9 7 2 ) a s s i g n e d a s i m i l a r a g e o f 37.5 may. for t h e Eocene-Oligocene b o u n d a r y , which r e l i e s i n p a r t , a t l e a s t , on a K - A r g l a u c o n i t e d a t e o f 37.5 2 3 m.y. (Odin e t a l , , 1969) for t h e s a n d s of N e e r r e p e n ( B e l g i a n B a s i n ) , which i s l a t e s t Eocene o r e a r l i e s t O l i g o c e n e i n a g e , The N o r t h American m i c r o t e k t i t e l a y e r , however, o c c u r s i n u p p e r Eocene s e d i m e n t s based on r a d i o l a r i a n , f o r a m i n i f e r a 1 and n a n n o f o s s i l d a t a for t h r e e c o r e s and h a s a n a b s o l u t e a g e o f 3 4 . 4 2 0.5 m.y. b a s e d on f i s s i o n - t r a c k and K - A r a g e s o f t h e N o r t h American t e k t i t e s and m i c r o t e k t i t e s . I n c o n c l u s i o n , t h e r e f o r e , t h e m i c r o t e k t i t e d a t a i n d i c a t e t h a t t h e Eocene-Oligocene boundary must b e l e s s t h a n 35 m.y. which i s 2-3 m i l l i o n y e a r s younger t h a n t h e a g e a s s i g n e d t o t h i s boundary by B e r g g r e n ( 1 9 7 3 ) , b u t i s w i t h i n t h e l i m i t s o f a c c u r a c y o f t h e K - A r a g e of t h e s a n d s o f
559
N e e r r e p e n g i v e n b y Odin e t a l . ( 1 9 6 9 ) . Acknowledgements J . A . Glass h e l p e d p r e p a r e t h e m a n u s c r i p t .
helped with r a d i o l a r i a n i d e n t i f i c a t i o n .
F , Maurrasse R e s e a r c h was s u p -
p o r t e d b y NSF g r a n t OCE72-01439 A C 4 , References B a k e r , R . N . , and Glass, B . P . , 1974, M i c r o t e k t i t e s a s t e s t components o f C a r i b b e a n a r e n a c e o u s F o r a m i n i f e r a , Microp a l e o n t o l o g y , v o l . 20, p p . 231-235. B a r n e s , V.E., and B a r n e s , M . A . , e d i t o r s , 1973, T e k t i t e s . I n : R . W . F a i r b r i d g e ( S e r i e s E d i t o r ) , Benchmark P a p e r s i n Geology, Dowden, H u t c h i n s o n and R o s s , S t r o u d s b u r g , P a .
,
445 PP* B e r g g r e n , W . A . , 1972, A Cenozoic t i m e s c a l e - some i m p l i c a t i o n s f o r r e g i o n a l g e o l o g y and p a l e o b i o g e o g r a p h y , L e t h a i a , v o l . 5 , p p . 195-215. Bukry, D . , 1973, C o c c o l i t h S t r a t i g r a p h y . I n : W o r z e l , J . L . , B r y a n t , W . , e t a l . , 1973, I n i t i a l R e p o r t s o f t h e Deep S e a D r i l l i n g P r o , i e c t , v o l . 1 0 , Washington (U.S. Government P r i n t i n g O f f i c e ) , pp. 385-406. D o n n e l l y , T . W . and Chao, E . C . T . , 1973, M i c r o t e k t i t e s of l a t e Eocene Age f r o m t h e e a s t e r n C a r i b b e a n S e a . I n : E d g a r , N.T., S a u n d e r s , J.B., e t a l . , 1973, I n i t i a l R e p o r t s f t h e Deep S e a D r i l l i n n Pro.i@, v o l . 15, Washington (U.S. Government P r i n t i n g O f f i c e ) , p p . 1031-1037. E d g a r , N . T . , S a u n d e r s , J.B., e t a l . , 1973, I n i t i a l R e p o r t s of t h e Deep S e a D r i l l i n 9 P r o j e c t , v o l . 15, Washington ( U . S . Government P r i n t i n g O f f i c e ) , 1137 p p . E v e r n d e n , J . F . and E v e r n d e n , R.K.S. , 1 9 7 0 , The Cenozoic t i m e s c a l e , G e o l o g i c a l S o c i e t y of America, S p e c i a l P a p e r 124,
7 1 PP. F l e i s c h e r , R . L . , and P r i c e , P . B . ,
1964, F i s s i o n - t r a c k e v i d e n c e
f o r t h e s i m u l t a n e o u s o r i g i n of t e k t i t e s and o t h e r n a t u r a l g l a s s e s . Geochimica e t Cosmochimica A c t a , v o l . 28,
P P . 755-760,
560
F l e i s c h e r , R . L . , P r i c e , P . B . , and W a l k e r , R . N . ,
1965, On t h e
s i m u l t a n e o u s o r i g i n of t e k t i t e s and o t h e r n a t u r a l g l a s s e s . Geochimica e t Cosmochimica A c t a , v o l . 29, p p . 161166, Foreman, E S P , , 1973, R a d i o l a r i a of Leg 10 w i t h s y s t e m a t i c s and r a n g e s f o r t h e f a m i l i e s Amphipyndacidae, A r t o s t r o b i i d a e , and T h e o p e r i d a e . I n : W o r z e l , J . L . , B r y a n t , W , , e t a l . , 1973, I n i t i a l R e p o r t s of t h e Deep S e a D r i l l i n g P r o , j e c t , v o l . 1 0 , Washington (U.S. Government P r i n t i n g O f f i c e ) , pp. 407-473. G a r l i c k , G . D . , TJaeser, C . W . , and O ’ N e i l , J . R . , 1971, A Cuban T e k t i t e . Geochimica e t Cosmochimica A c t a , v o l , 35, PP. 731-734. G e n t n e r , W., S t o r z e r , D . , and Wagner, G . A . , 1969, New f i s s i o n s r a c k a g e s of t e k t i t e s and r e l a t e d g l a s s e s . Geochimica e t Cosmochimica A c t a , v o l . 33, pp. 1075-1081. Glass, a . P . , 1969a, Chemical c o m p o s i t i o n of I v o r y C o a s t m i c r o t e k t i t e s , Geochimica e t Cosmochimica A c t a , v o l . 33, p p , 1135-1147.
G l a s s , 3.P., 1969b, Reworking of d e e p - s e a s e d i m e n t s a s i n d i c a t e d b y t h e v e r t i c a l d i s p e r s i o n o f t h e A u s t r a l a s i a n and I v o r y C o a s t m i c r o t e k t i t e h o r i z o n s , E a r t h and P l a n e t a r y S c i e n c e L e t t e r s , v o l . 6 , p p . 409-415. Glass, B . P . , 1 9 7 2 , A u s t r a l a s i a n M i c r o t e k t i t e s i n d e e p - s e a s e d i m e n t s . I n : Hayes, D . E . , ( e d i t o r ) , A n t a r c t i c Oceanology 11: The A u s t r a l a s i a n - N e w Zealand S e c t o r , A n t a r c t i c R e s e a r c h S e r i e s , v o l . 1 9 , Washington (American Geophysi c a l U n i o n ) , p p . 335-348. Glass, 3 . P . , 1975, Geomagnetic r e v e r s a l s a n d t e k t i t e s , I n : F i s h e r , R . N . , F u l l e r , N . , Schmidt, V.A. and Wasilewski, P . J . ( E d i t o r s ) . P r o c e e d i n g s o f t h e T a k e s i Kagata Conf e r e n c e , J u n e 3-4, 1974, G r e e n b e l t , Maryland (Goddard S p a c e F l i g h t C e n t e r ) , p p . 225-229. G l a s s , B . P . , and Heezen, B . C . , 1 9 6 7 , T e k t i t e s and g e o m a g n e t i c r e v e r s a l s . S c i e n t i f i c American, v o l . 2 1 7 , p p . 32-38.
56 1
Glass, B . P . , B a k e r , R . N . , S t o r z e r , D . and Wagner, G.A., 1 9 7 3 , North American m i c r o t e k t i t e s from t h e C a r i b b e a n S e a and t h e i r f i s s i o n t r a c k a g e . E a r t h and P l a n e t a r y S c i e n c e L e t t e r s , v o l . 1 9 , pp. 184-192. Hay, W . W . , 1973, P r e l i m i n a r y d a t i n g b y f o s s i l c a l c a r e o u s n a n n o p l a n k t o n , Deep S e a D r i l l i n g P r o j e c t : Leg 1 0 . I n : Worzel, J . L . , B r y a n t , W., e t a l . , 1973, I n i t i a l R e p o r t s
of t h e
Deep S e a D r i l l i n g Pro:,ect, v o l . 1 0 , Washington ( U . S . Government P r i n t i n g O f f i c e ) , p p , 375-383.
Hay, W . W . , and S e a u d r y , F . M . , 1973, C a l c a r e o u s n a n n o f o s s i l s Leg 15, Deep S e a D r i l l i n g P r o j e c t , I n : E d g a r , N . T . , S a u n d e r s , J . B . , e t a l , , 1973, I n i t i a l R e p o r t s of t h e Deep S e a D r i l l i n g P r o . i e c t , v o l . 15, Washington ( U . S . Government P r i n t i n g O f f i c e ) , p p . 625-683. N a u r r a s s e , F . and Glass, B . P . , g r a p h y and N o r t h American c o r e RC9-58: I m p l i c a t i o n s l a r i a n c h r o n o l o g y and t h e
-
I n press, Radiolarian stratim i c r o t e k t i t e s i n Caribbean c o n c e r n i n g l a t e Eocene r a d i o a g e o f t h e Eocene-Oligocene
boundary. Proceedings o f t h e 7 t h Caribbean G e o l o g i c a l C o n f e r e n c e , J u l y , 1974, Guadeloupe. Mcn’eely, B . W . , 1973, B i o s t r a t i g r a p h y o f t h e Nesozoic and P a l e o g e n e p e l a g i c s e d i m e n t s of t h e Campeche Embankment a r e a . I n : W o r z e l , J . L . , B r y a n t , W . , e t a l . , 1973, I n i t i a l R e p o r t s o f t h e Deep S e a D r i l l i n g P r o j e c t , v o l , 1 0 , Washi n g t o n ( U . S . Government P r i n t i n g O f f i c e ) , p p , 679-695. Odin, C . S . , G u l i n c k , M . , B o d e l l e , J . , and Lay, C . , 1 9 6 9 , G 6 o c h r o n o l o g i e d e n i v e a u x g i a u c o n i e u x t e r t i a r e s du b a s i n de B e l g i q u e ( m i t h o d e p o t a s s i u m - a r g o n ) , C . R . Somm S g a n c e s S O C . Ggol. F r a n c e , v o l . 6 , p , 198. G’Keefe, J . A . , 1 9 7 6 , T e k t i t e s and t h e i r o r i g i n , New York ( E l s e v i e r ) , 254 p p . Riedel, W.R.,
and S a n f i l i p p o , A , , 1973, Cenozoic R a d i o l a r i a
from t h e C a r i b b e a n , Deep S e a D r i l l i n g P r o j e c t , Leg 15. I n : E d g a r , N . T . , S a u n d e r s , J . B . , e t a l , , 1973, I n i t i a l R e p o r t s of t h e Deep S e a D r i l l i n g P r o . i e c t , v o l .
15,
Washington ( U . S . Government P r i n t i n g O f f i c e ) , pp. 705-
751.
562
Sanfilippo, A , , and Riedel, W.R., 1973, Cenozoic Radiolaria (Exclusive of Theoperids, Artostrobids and Amphipyndacids) from the Gulf of Mexico, Deep Sea Drilling Project Leg 10. In: Worzel, J.L., Bryant, W , , et al., 1973, Initial Reports of the Deep Sea Drilling Project, vol. 10, Washington (U.S. Government Printing Office), pp. 475-611. Storzer, D., and Wagner, G.A., 1971, Fission track ages of North American tektites. Earth and Planetary Science Letters, v o l . lo, pp. 435-444. Storzer, D., Wagner, G.A., and King, E.A., 1973, Fission track ages and stratigraphic occurrence of Georgia tektites. Journal of Geophysical Research, v o l . 78, pp. 4915-4919. Worzel, J.L., Bryant, W., et al., 1973, Initial Reports of the Deep Sea Drilling Project, vol. 10, Washington (U.S. Government Printing Office) , 748 pp. Zwart, M.J., 1976, North American microtektites from Deep Sea Drilling Project Cores (Unpublished Master‘s Thesis, University of Delaware, Newark, DE) , Zwart, M.J. and Glass, B.P., 1975, North American microtektites from DSDP cores: associated radiolarian extinctions and paleomagnetic stratigraphy. American Geophysical Union, Transactions, v o l . 56, p. 385 (abstract). Zghringer, J . , 1963, K-Ar measurements of tektites. In: Radioactive Dating. Proceedings Symposium, Athens, Nov. 19-23, 1962, International Atomic Energy Agency, Vienna, pp. 289-305.
0
0
00
0
Fig. 1. Tektite strewnfields. Deep-sea cores searched f o r Australasian, I v o r y Coast and North American microtektites indicated by circles, triangles and squares, respectively. Cores found to contain microtektites indicated by solid figures.
564
R C9-58
Mi CR OT EKTlT ES/grn 1
0
2
3
' 7 (x100)
Lz W
I
Fig. 2.
V e r t i c a l d i s t r i b u t i o n o f m i c r o t e k t i t e s and abundance of f i v e R a d i o l a r i a s p e c i e s i n c o r e RC9-58.
DSDP
SITE 9 4
MICROTEKTITES/IO crn3 0
2
4
6
8 1 0 1 2
415
+ 416 W
I
n 417+
418
f
m 10%
Fig.
3.
m 50%
m 5%
m 20%
m 5%
V e r t i c a l d i s t r i b u t i o n of m i c r o t e k t i t e s and abundance
o f f i v e R a d i o l a r i a s p e c i e s i n s e c t i o n s 3 and 4 o f c o r e 15, DSDP s i t e 94. Depth i s f r o m s e d i m e n t - w a t e r i n t e r f a c e .
DSDP
SITE 149
MICROTEKTITES/10cm3 5
0
10
1s
20
(XlOO)
1
I I
-----
271J
10%
Fig.
4.
20%
20%
10%
5%
V e r t i c a l d i s t r i b u t i o n of m i c r . o t e k t i t e s and abundance
o f f i v e R a d i o l a r i a s p e c i e s i n s e c t i o n 2 , c o r e 30 and s e c t i o n
1, c o r e 31, DSDP s i t e terface.
149. Depth is from sediment-water i n cn m cn
Discussion Dr.
B. K .
Holdsworth:
T h e r e seems some d o u b t w h e t h e r t h e
T h y r s o c y r t i s s p p . e x t i n c t i o n l e v e l i n deep-sea c o r e s r e a l l y
i s t h e Eocene/Oligocene boundary. I t would c o n v e n t i o n a l l y be t a k e n a t t h e b a s e of t h e t u b e r o s a Zone - which i s a h i g h e r
.;
level. Dr.
B. P. Glass:
I d i d n o t mean t o imply t h a t t h e r a d i o l a r i a n
e x t i n c t i o n s o c c u r r e d a t t h e Eocene/Oligocene boundary.
In
t h e DSDP r e p o r t s , t h e e x t i n c t i o n s d o o c c u r below t h e Eocene/ O l i g o c e n e boundary.
T h i s means t h a t t h e m i c r o t e k t i t e s g i v e
an upper l i m i t t o t h e a g e of t h e Eocene/Oligocene boundary. Dr.
F . Michael:
How a r e t h e e x t i n c t i o n of t h e R a d i o l a r i a and
t h e m i c r o t e k t i t e zone r e l a t e d t o p a l e o m a g n e t i c r e v e r s a l s ? Glass:
The A u s t r a l i a n and I v o r y C o a s t m i c r o t e k t i t e s a p p e a r t o
be a s s o c i a t e d w i t h t h e Brunhes/Matuyama boundary and J a r a m i l l o event, respectively.
The N o r t h American m i c r o t e k t i t e s d o n o t
appear t o be associated with a r e v e r s a l .
The s e c t i o n s of
cores containing t h e m i c r o t e k t i t e l a y e r appear l y normally magnetized.
t o be e n t i r e -
However, i t ' s v e r y d i f f i c u l t t o g e t
good p a l e o m a g n e t i c s t r a t i g r a p h y from t h e s e e n c l o s e d , o r semie n c l o s e d b a s i n s , l i k e t h e C a r i b b e a n and t h e Gulf of Mexico. They g e n e r a l l y have a normal o v e r p r i n t on t o p of t h e m a g n e t i c s t r a t i g r a p h y t h a t ' s v e r y d i f f i c u l t t o remove.
So w e d o n ' t
know t h e answer t o y o u r q u e s t i o n . I d o n ' t know t h e r e l a t i o n s h i p between N o r t h American m i c r o t e k t i t e s and t h e paleomagnetic stratigraphy a t this t i m e . Dr.
L.
Glass:
Shishkevish:
Were m i c r o t e k t i t e s found i n moon s a m p l e s ?
G l a s s b e a d s were found i n
T h a t ' a hard one t o answer.
l u n a r s a m p l e s w i t h a p p r o x i m a t e l y t h e same s h a p e and s i z e d i s t r i b u t i o n as t h e m i c r o t e k t i t e s .
Chemically t h e y a r e d i s t i n c t
from m i c r o t e k t i t e s and t e k t i t e m a t e r i a l .
M i c r o t e k t i t e s and
t e k t i t e s g e n e r a l l y have s i l i c a c o n t e n t s g r e a t e r t h a n 60%, whereas most l u n a r m a t e r i a l h a s s i l i c a c o n t e n t s l e s s t h a n 6 0 % . So c h e m i c a l l y t h e y a r e d i s t i n c t .
They may b e t h e same i n
t h a t t h e y may b o t h have been formed by t h e same p r o c e s s : t h a t i s meteorite impact.
So i f you want t o c a l l g l a s s b e a d s p r o -
duced by m e t e o r i t e i m p a c t m i c r o t e k t i t e s , t h e n y e s , m i c r o t e k t i t e s have been found on t h e moon.
I f you want a s t r i c t e r de-
f i n i t i o n t o c h e m i s t r y , t h e n , n o , t h e y ' r e n o t found on t h e moon.
567 D r . M.
C.
F i r s t of a l l I ' d l i k e t o p o i n t o u t t h a t w e d o
Keen:
know where t h e Eocene-Oligocene boundary i s , of c o u r s e .
We
may n o t know where it i s i n DSDP c o r e s , b u t w e c e r t a i n l y know where i t i s i n r e l a t i o n s h i p t o t h e s t r a t o t y p e .
J u s t before I
came h e r e I saw some a g e d a t e s t h a t had b e e n , I t h i n k f a i r l y new o n e s , by P r o f . C u r r i e .
U n f o r t u n a t e l y I c a n ' t remember
them e x a c t l y , b u t I have a f e e l i n g t h a t t h e y ' r e more i n l i n e w i t h what you w a n t .
B u t , I j u s t c a n ' t remember: I seem t o
remember s e e i n g 3 4 m . y . , Glass:
b u t i t ' s j u s t from memory.
I f you c o u l d g e t a h o l d of t h o s e and s e n d them t o m e
I ' d be v e r y happy. N o more q u e s t i o n s ? Dr.
G . Williams:
Can w e assume t h a t t h e m e t e o r i t e s d i d n ' t h i t
t h e E a r t h b e f o r e t h e Eocene. Glass:
Before?
O r i s i t a problem o f . . .
I n o t h e r words y o u ' d l i k e t o p u t them i n c o l d
s t o r a g e f o r a w h i l e b e f o r e you d e p o s i t them on t h e e a r t h i s t h a t it? Williams: Glass:
NO,
I was wondering why
...
Okay, i n o t h e r words, youmean no t e k t i t e s o r m i c r o -
t e k t i t e s would b e found o l d e r t h a n 3 4 m i l l i o n y e a r s .
I don't
know i f t h a t ' s a problem o f j u s t n o t f i n d i n g them o r w h e t h e r
i t ' s a problem of them u n d e r g o i n g s o l u t i o n o r b e i n g d e s t r o y e d o v e r t h a t t i m e p e r i o d . A s you know, o r may n o t know v o l c a n i c g l a s s e s a r e f a i r l y u n s t a b l e and a s you g e t back p r e t t y f a r i n t h e t i m e s c a l e t h e y seem t o b e l a c k i n g .
I
t h i n k t h i s i s m o s t l y a w e a t h e r i n g problem, b e i n g d e s t r o y e d . Whether o r n o t t h e y a c t u a l l y e x i s t e d a t one t i m e , o r w e j u s t h a v e n ' t found them, o r t h e y ' v e been d i s s o l v e d by s o l u t i o n , o r w h a t , I d o n ' t know. But c e r t a i n l y none have been found older than about 34 m i l l i o n years. I d o n ' t know t h e a n s w e r . Williams:
Could you t e l l m e t h e n how you c o n c e n t r a t e them
p l e a s e and how you g e t them o u t of t h e s e d i m e n t s ?
Is i t l i k e
foram p r e p a r a t i o n ? Glass:
Yes,
i t ' s very simple.
You t a k e a s e d i m e n t , d i s a g g r e -
g a t e i t and w e t s i e v e i t and l o o k a t t h e g r e a t e r t h a n 125 m i c r o n s i z e f r a c t i o n . S i n c e we're d e a l i n g w i t h p e l a g i c s e d i ments f o r t h e most p a r t t h e y a r e t h e y o n l y t h i n g i n t h e r e l a r g e r t h a n 125 m i c r o n s , u n l e s s you have f o r a m s , o r s o m e t h i n g ,
568
i n w h i c h c a s e , I h a t e t o s a y i t h e r e i n f r o n t of m i c r o p a l e o n t o l o g i s t s , b u t I u s e a l i t t l e a c i d and g e t r i d of them. The o n l y t h i n g s l e f t a r e t h e m i c r o t e k t i t e s .
569
Index t o Authors Aberdeen, E. 1 7 3 , 1 7 8 Adams, C . G . 3 9 7 , 411 Adegoke, O . S . 422, 425-27, 432 Aguirre, E. 4 0 6 Aders, W. H. 4 0 1 , 4 0 5 A l b e r i c i , A. 4 5 8 , 4 6 1 A l b e r t i , G . 3 7 1 , 3 7 3 , 3 7 8 , 392 Aldridge, R . J . 9 5 , 9 7 , 1 0 3 , 107 Alexander, C . 2 5 7 - 5 8 , 263-65 Aliev, Kh. Sh. 3 1 1 , 317 A l l e n , N. W. 250 Alves, R. J . 270 Amdurer, M. 4 3 1 , 4 5 8 , 4 6 1 Amprom, H. W. J. van 7 9 Anderson, F. W. 70, 79, 233, 241, 2 4 7 , 265 Anderson, G. C . 5 3 4 Andress, N. E. 164 Andrews, J . E. 535 Andries, R. R. 4 1 7 , 4 3 2 Antunez, M. T. 4 2 4 , 4 2 7 , 4 3 2 , 4 5 1 , 461 Apollonov, M. K. 1 7 9 Apostolescu, V. 2 8 3 , 2 8 8 - 8 9 , 291-92 2 9 4 , 2 9 8 , 4 9 6 - 9 7 , 5 0 3 , 5 0 5 , 513 Applin, E. R. 2 0 6 , 220 Araujo, S . J. 320 Aubert, J. 3 9 5 , 3 9 8 - 9 9 , 4 0 5 A u s t i n , R. L . 1 0 7 , 1 1 8 , 1 2 0 - 2 1 , 1 3 1 - 3 2
Becker, F. 7 9 Becker, G. 6 3 - 6 6 , 6 8 - 6 9 , 7 1 - 7 2 , 7 6 , 79 Beckman, J. P. 2 1 0 , 317 Behnken, F. H. 122-23, 131 Belasco, C . 1 9 9 B e l l , W. A. 7 0 , 80 Bellion, Y. 2 6 4 , 266 Belmonte, Y . C . 2 9 2 , 2 9 8 , 4 2 3 , 4 2 6 - 2 7 , 433, 448, 461 Bengtsson, R. H. 8 7 - 8 8 , 103 Benait, A. 162, 164 Benson, R. H. 263, 266, 406, 496, 5 0 8 - 1 1 , 513 Benson, W. E. 2 0 6 , 2 1 0 , 219 Berdan, J. M. 3 - 4 , 7 , 6 2 , 80 Berger, W. H. 5 2 4 , 534 Berggren, W. A. 2 0 6 , 2 1 9 , 3 9 0 - 9 1 , 393-403, 405, 408 419, 425-26, 429-30, 433, 458, 461, 504, 513, 523, 534 Bergstrom, S . M. 8 4 , 87-94, 98-99, 101-104, 1 0 7 , 1 1 3 , 1 3 2 , 169, 178 Berry, W. B. N. 5 B e r t e l s , A. 282-83, 2 9 8 , 416-17, 420-21, 433, 453-54, 461-62, 496, 498, 514 B e t t e n s t a e d t , F. 2 6 5 - 6 6 , 3 9 8 , 406 Beurlen, K. 2 8 0 , 298 Bharadwaj, D. C. 1 7 7 - 7 8 Bachman, A. 1 3 8 , 1 4 3 , 549-50 B i l l i n g s , E. 3 Baker, J. H. 513 Bjorklund, K. R. 1 7 8 , 534-35 Raker, R. A. 211, 221 Blanc, P. L . 2 6 4 , 266 Raker, R. N. 556, 559, 561 Bless, M. J . M. 6 3 , 6 5 , 6 8 - 7 2 , 75B a l l a r d , R. D. 1 8 5 , 197 7 7 , 7 9 , 8 0 , 83 Bandel, D. 6 4 , 79 Blondeau, M. A. 4 2 3 , 4 2 6 , 4 3 3 Randy, 0. 1,. 5 2 3 , 5 2 6 , 5 2 8 , 5 3 0 , 5 3 4 Blow, W. H. 3 9 4 , 3 9 7 - 9 8 , 4 1 0 , 4 0 5 5 4 8 , 550 407, 428-29, 433, 416, 450, 456, Bangerow, E. F. 8 4 4 6 0 , 462 Banner, F. 'T. 2 3 6 - 3 7 , 2 4 1 , 407 Bode, H. 11. 1.91 B a r b i e r i , F. 4 5 8 , 4 6 1 Bodrlle, J. 5 6 1 Barker, R. W, 3 9 4 , 3 9 6 - 9 8 , 4 0 3 , 4 0 5 Bold, W, A. van den 4 9 6 , 4 9 8 , 5 0 3 , Barlow, J . A. 1 9 7 5 0 5 - 0 6 , 5 0 8 - 0 9 , 514-16 Barnes, C. R. YO, 1 0 2 - 0 3 , 107 B o l l i , H. M. 4 0 1 , 4 0 6 , 4 3 1 , 4 3 3 , 4 5 2 Barnes, M. A. 5 5 4 , 559 Boltovskoy, H. M. 4 5 4 , 4 5 9 , 462 Barnes, V. E. 5 5 4 , 559 Bond, G. 4 5 5 , 4 6 2 , Barr, F. T. 2 0 9 , 219 Boogaard, M. van den 9 6 , 1 0 4 Barss, M. S . 1 9 1 - 9 2 , 1 9 7 Bouckaert, J. 1 2 0 , 1 3 1 Bassiouni, M. e l A. A. 236, 241, Boucot, A. J. 4 , 5 2 5 6 , 265 Boudda-Ahmed 198 B a s s l e r , R. S . 6 3 , 7 9 , 8 4 Bradshaw, L . E. 92, 104 Bate, R. H. 233-34, 237-38, 241, Brady, G . S . 516 2 8 0 , 298 Bramlette, M. N. 317 B a t j e s , D. A. J. 3 9 6 , 405 Brand, U. 9 1 , 1 0 4 Beaudry, F. M. 561 Braun, P. G. 0. 2 7 9 - 8 0 , 2 8 2 , 298 Becker, D. 5 0 3 , 513 Braun, W. K. 6 4 , 6 7 , 8 0 , 2 3 5 , 2 4 1
570
Brenner, G. J. 3 6 2 - 6 3 , 366 Briden, J. C. 1 3 2 , 178 Brinkmann, R. 3 9 2 , 406 B r i t o , I. M. 1 4 1 , 143 Brognon, G . 2 8 8 , 2 9 0 - 9 1 , 2 9 8 , 4 2 4 , 427, 433, 451 Bronnimann, P. 3 1 7 , 516 Brooke, M. M. 235, 241 B r o t z e n , F. 3 9 8 , 4 0 6 Brown, P. M. 239, 242, 250, 257-59, 2 6 9 , 3 9 2 , 406 Brown, R. H. 197 Brun, L. 283, 299, 424, 427, 449, 463 Bryant, W. 562 Bukry, D. 317, 346, 366, 546, 547, 5 5 0 , 5 5 7 , 559 Bulman, 0. M. B. 9 5 , 1 0 4 B u r b r i d g e , P. P. 3 7 1 , 393 B u r c k l e , L . H. 4 2 8 , 4 3 5 , 4 5 6 , 4 6 4 , 536 Burk, C. F. 8 0 Burk, C. F. Jr. 4 , 9 Burmann, G. 1 3 7 , 1 4 3 Burns, R. E . 535 Burrett, C. 101, 1 0 4 B u t l e r , E. A. 2 6 3 , 266
Clendening, J. A . 1 9 8 Gloss, D. 4 4 5 - 4 6 , 4 5 3 , 4 6 2 Coleman, B. E. 233-34, 2 3 7 , 241 C o l i n , J. P. 2 3 8 , 2 6 3 - 6 4 , 266 C o l l i n s o n , C. 1 2 0 , 131-32, 1 5 5 , 164 Colom, G. 5 4 8 , 5 5 1 Combaz, A. 1 4 4 , 1 5 6 , 1 6 4 Conkin, B. M. 5 1 , 56 Conkin, J. E . 5 1 , 56 Connin, J. 1 9 9 Cook, H. E. 101, 1 0 4 , 1 2 9 , 1 3 1 Cookson, I. C. 371-72, 392-94 Cooper, B. J. 9 5 , 1 0 4 Copeland, M. J. 3 - 5 , 7 , 6 2 , 7 0 , 8 0 C o r n e t , B. 3 2 9 , 337 Couper, R. A . 355-56 C o u r t v i l l e , J. 392 Cousminer, H. L. 1 8 6 , 1 9 1 - 9 3 , 1 9 8 , 200 Coutinho, P. N. 282, 299, 4 1 4 , 434 Couvering, J. A . van 4 0 6 , 523 Cramer, F. H. 1 3 7 - 4 1 , 1 4 3 - 4 4 , 1 5 2 , 1 5 7 , 159 C r e a t h , W. B. 70, 79, 81 C r o s s , A. T . 1 9 9 Curry, D. 3 9 6 , 406 C u r t i s , G. H. 4 0 6 Cushman, J . A . 4 0 2 , 4 0 7
Camacho, H. H. 417 Campo, M. van 356 Carnes, J . B. 9 4 , 1 0 2 , 104 Carter, G. F. E. 4 Casey, R. E. 523, 526-27, 530-32, 5 3 4 , 550 C a s t r o , J. C . 300, 413, 418, 420, 4 3 5 , 464 C a u d r i , C. M. B. 3 9 7 , 406-07 Cavaroc, V. V. 1 9 2 , 1 9 8 Cepek, P. 317 Chao, E. C. T. 5 5 5 , 559 Chapman, F. 4 2 8 , 4 3 3 C h a r r i e r , R. 4 1 8 , 4 3 3 C h e b l i , G. 2 8 1 , 299 C h o l i c h , R. de C . 519 Choubert, G. 1 9 1 , 188 C h r i s t e n s e n , 0 . B. 2 5 0 , 256 C h r i s t o p h e r , R. A. 1 4 3 Churkin, J. Jr. 535 C i e s i e l s k i , P. F. 5 4 8 - 4 9 , 551-52 C i f e l l i , R. 4 2 8 , 4 3 3 , 5 3 4 C i t a , M. B. 2 0 6 , 2 2 0 Clariond, L. 1 9 3 , 198 C l a r k , D. L . 90-91, 104, 118, 122, 1 2 3 , 1 2 7 , 131-32 C l a r k , J . M. 4 C l a r k e , R. F. A. 3 7 1 , 392 C l a r k e , W. J . 4 0 7
d a Gama, E. G. Jr. 3 0 0 Dale, B. 394 Dam, A . t e n 4 0 2 , 407 Damotte, R. 264, 266 Darrah, W. C. 199 Daubinger, J . 1 9 0 - 9 2 , 1 9 9 Davey, R. J. 3 4 6 , 3 6 6 , 3 7 1 , 3 7 4 , 392 Davidson, S . E . 3 9 3 D a v i s , T. A. 535 Dawson, J. W. 4 d e Boer, K. F. 3 3 0 , 338 D e f l a n d r e , G. 1 3 8 - 3 9 , 1 4 3 , 1 7 3 , 1 7 5 , 1 7 9 , 3 7 2 , 392 D e f r e t i n , S. 1 9 9 d e K l a s z , I. 2 8 3 - 8 4 , 2 8 9 - 9 0 , 2 9 8 - 9 9 , 423-27, 4 3 3 - 3 4 , 4 4 9 - 5 0 , 4 5 5 , 462463 Delaney, P. J. 4 5 4 , 4 7 0 Dennison, J. M. 5 Deroo, G. 2 6 3 - 6 4 , 267 Deunff, J. 1 3 7 - 3 8 , 1 4 2 , 1 4 4 Dewey, J. F. 185, 199 D i e z , M. d . C , R. 1 3 9 , 1 4 3 - 4 4 , 165 D i g r e g o r i o , J . H. 2 8 0 - 8 1 , 298 Dinesen, A. 3 9 6 , 4 0 8 D i n g l e , R. V. 288-89, 294-95, 298, 5 0 1 , 516 Dodekova, L. 3 3 2 , 337
571 Donnelly, T . W. 5 5 5 , 559 Donze, P. 2 3 5 - 3 8 , 2 4 1 , 2 5 1 - 5 3 , 2 6 2 ,
266-67 dos Reis, F. J. 300 Douglas, J. G . 393 Douglas, R. G. 409 Downie, C . 1 3 8 - 3 9 , 1 4 2 , 1 4 4 - 4 5 , 393 Doyle, J. A. 355-56, 362-67, 371,
3 7 3 , 383 Drake, R. E. 406 Drewrey, G. E. 1 3 2 , 178 D r e x l e r , E . 2 3 6 , 241 Dreyfus, R. M. 536 Drooger, C . W . 4 4 9 , 4 6 4 , 516 Druce, E. C . 8 7 , 9 1 , 104 Drugg, W. S . 3 7 1 , 383 D u m i t r i c a , P. 3 1 1 , 3 1 7 , 5 3 5 , 5 4 9 ,
551
2 0 , 4 3 4 , 4 5 3 , 463 173-74, 1 7 9 , 3 1 1 , 3 1 4 , 3 1 7 , 557 F o r s y t h , D . W. 1 8 5 , 201 F o r t e y , R. A . 1 6 9 , 179 Fourcade, E. 2 6 4 , 266 Fox, H. 175 F r a n c e s c o , F. 0 . d e 4 5 4 , 463 F u g a n t i , A. 9 4 , 105 F u n n e l l , B. M. 4 2 9 , 4 3 4 , 4 5 8 , 463 F u r n i s h , W. 8 7 , 9 5 , 1 0 5 , 1 2 4 , 131 Foreman, H. P.
423-24, 427, 433, 449, 4 5 0 , 4 5 5 , 462 Galehouse, J. S. 535 G a r l i c k , G. D. 5 5 8 , 560 G a r t n e r , S . J r . 2 0 6 , 220 G a t e s , W. L. 4 0 1 , 407 G e i g e r , M. E . 3 2 9 , 337 G e n t n e r , W. 5 5 4 , 5 5 8 , 560 G e r r y , E. 2 3 5 , 241 Gibson, L. B. 6 7 , 81 G i l l e s p i e , W. 199 Gitmez, G. u. 3 3 4 , 337 G l a s s , B. P. 5 5 4 , 556-62 G l e z e r , 2. I. 5 4 8 - 4 9 , 551 Gocht, H. 3 3 2 , 337 G o l d s t e i n , R. F. 1 6 0 , 164 G o l l , R. M. 534-35 Gonzalez, B. B . 463 Gordon, W. A. 219-20 Graham, J. J . 4 2 4 , 4 2 7 , 4 5 0 , 463 G r a n t , A. B. 535 Graves, R. W. J r . 1 7 3 , 179 Grebe, H. 7 5 , 82 G r e b e r , C . 1 9 3 , 199-200 Green, R. 7 0 , 80 G r e k o f f , N. 284, 286-87, 290-91, 2 9 8 , 5 1 6 , 517 G r o s d i d i e r , E. 2 8 4 , 2 8 7 , 2C9, 298 Gruqdel, J . 6 7 - 6 8 , 7 1 , 2 6 2 , 267 Guiraud, R. 266 G u l i n c k , M. 561 Gageonnet, R.
Dunay, R. E . 199 Dunham, J. B . 1 7 0 , 179 Durden, C . J . 199 D u t u i t , J. M. 199 E a g e r , R. M. C . 199 Eames, F. E. 3 9 4 , 3 9 7 , 407 E c h o l s , D. J. 7 0 , 7 9 , 8 1 Edgar, N. T. 5 5 5 , 5 5 7 , 559 Edmund, J. M . 534 Edwards, A. R. 535 E i s e n a c k , A. 137-39, 144, 152, 155,
1 5 7 , 1 7 5 , 3 7 1 - 7 3 , 383 E l o f s o n , 0 . 518 Emery, K. 0 . 3 9 1 , 407 E p s t e i n , A. G. 104 E p s t e i n , J. B. 104 E r i c s o n , D. B. 4 5 9 , 463 E t h i n g t o n , R. L. 8 7 , 9 0 - 9 1 , 9 5 , 104
1 0 5 , 107 Evernden, J. F. 559 Evernden, R. K. S . 559 E v e r e t t , J. C . 197 E v i t t , W. R. 389 Ewing, M. 4 2 8 , 4 3 5 , 4 5 6 , 464 F i h r a e u s , L. F. F a l c e t t a , M. M.
F i d a l g o , F. 4 5 4 , 463 F i s h e r , D. W. 144 F i s h e r , M. J. 199 F l e i s c h e r , R. L. 558-60 Fontana dos S a n t o s , J . 4 1 6 , 4 1 9 -
9 3 , 1 0 2 - 0 3 , 105 445, 453, 460, 462,
463 F a n n i n g , K. A. 534 F a u l k n e r , J. S. 4 4 9 - 5 0 , 463 Faure-Muret, A, 1 9 1 , 198 F e i s t , R. 9 6 , 105 F e l i x , C . J. 3 7 1 , 383 F e r e l l o , R. 299 F e r n a n d e s , J. M. 4 4 5 - 4 6 , 473 F e r r e i r a , C, S . 4 4 5 , 473 F e r r e i r a , J. M. 4 5 2 , 4 6 4 F e y s , R. 199
341, 343, 347, 348-49, 3 5 0 - 5 2 , 3 6 6 - 6 7 , 3 7 2 , 3 7 4 , 383-84 Hacquebard, P. A. 1 9 0 , 1 9 2 , 1 9 5 , 197 H a m a r , G. 9 4 , 105 Hamond, S. R. 5 2 6 - 2 7 , 5 2 9 , 536 Hanna, G . D. 5 4 6 , 551 Hansen, H. J. 407 Habib, D.
572 Haq, B. 401-02, 406 Harland, R. 371, 374, 383 Harper, H. E . J r . 524, 535 Hartmann, G. 496, 508-09, 516 Hartmann Schroder, G. 516 Hasenmuller, W. A . 132 Hass, W. H. 1 7 4 , 179 Hathaway, J. C . 352, 366 Havlena, V. 200 Hay, W. W. 557, 561 Hayes, D. E . 206, 220, 559 Hays, J. D. 526, 535 Hazel, J . E. 258-59, 262, 267, 401, 407 Head, J . W. 5 Hedberg, H. 392, 407 Heezen, B. C. 554, 560 Heide, S . van d e r 80 Heimdach, F. F. 236, 241 H e l f r i c h , C. T. 95, 105 Hendrix, W. E. 6 7 , 74 Hennen, G. J. 199 Hem, D o 418 Herngreen, G. F . W. 330, 338 H e r r i g , E. 259, 262, 264-65, 268 Higgins, A. C. 120, 131 Hilterman, H. 398, 409 Hinde, G. J . 173, 175, 179 H o f f m e i s t e r , W. S . 139, 144 Holdsworth, B. K. 169, 175-76, 179 H o l l i s t e r , C. 206, 219, 513 H o l l i s t e r , C . D. 352, 366, 390-91, 394, 406, 504 Hooyberghs, H. J. F. 401, 407 Hope, R. 201 Hopping, C. A. 329, 339 Horq, V. 290, 299 Huddle, J. W. 131 Hughes, C. F . 101, 108 Hulings, N . C. 513 I a c c a r i n e , S. 458, 461 Ichikawa, W. 549-50 I s r a e l s k y , M. 259, 262-64, 268 J a n s a , L. F. 200, 335-36, 338 J a n s o n i u s , J. 1 5 4 , 165 J a r d i n e , S. 140, 144 Jekhowsky, B. de 143-44, 156, 1 5 9 , 163, 166 J e n d r z e j e w s k i , J. P. 549, 551 J e n k i n s , W. A. M. 153, 155, 160-62, 165, 374, 383 Jeppsson, L . 95-97, 105 Johnson, D. A . 526, 535 Johnson, T. 305, 317, 535 J o n e s , D. E. 263, 266
J o n e s , P. J. 8 7 , 91, 104 J o n e s , T. R. 70-71, 75, 8 0 , 516 J o r d a n , H. 63-65, 68, 70, 72, 75, 77, 80-81 J o u r a , J. A. 300 K a a s s c h i e t e r , J. P. H. 396-97, 416, 516 Kanes, W. H. 125, 180 Kay, G. M. 4 , 5 , 104 Keen, M. C. 516 K e i j , A . J . 498, 516-17 K e l l e t t , B. 73, 77 Kemp, E . M. 362, 366 Kemper, E. 252-53, 268 Kennett, J. P. 394, 398, 407, 523, 535 K e s l i n g , R. V. 65, 67, 81 K i l e n y i , T. I. 235, 241, 250, 268 K i l g o r e , J . E . 65, 8 1 King, E. A . 562 Kirkby, J . W. 70-71, 75, 8 1 Klapper, G. 125-26, 131, 173 K l i e , W. 517 K l i n e , J. K. 1 5 3 , 166 Kling, S. A. 1 7 6 , 179 K l i n g e r , W. 241 Kniker, H. T. 420 K n o l l , A. H. 524, 526, 535 Knupfer, J . 94, 105 Kobayashi, T. 522, 535 Koch, R. 383 Kockel, F. 96, 105 Kohut, J. J. 93, 105 Kogbe, C. 502, 517 Kellman, K. 236, 241 KDtzian, S. B. 453, 462 Kozlova, G. 311, 317-18 Krasheninnikov, V. 317 Kremp, G. 75, 82 K r o m e l b e i n , K. 65, 67, 79, 8 2 , 84, 274, 282-84, 286, 289, 298-99, 503, 506-07 Kueler, H. G. 397, 407 Kullmann, J. 79 Kurmnerow, E. 71-72, 75, 82 Lahsen, A. Lamont, A. L a n c e l o t , Y. Landing, E. Lane, H. R. Lange, F. W. Lanzoni, E. L a r s e n , G. Latham, M.
418, 433 93-94, 105 317, 352, 366 87-88, 93, 99, 105 118, 121, 132 153, 166 140, 142, 145 396, 408 82
573
Laufeld, s.
153. 155. 159. 160. 1 6 2 , 1 6 3 , 166 Laughton, A. S. 2 0 6 , 2 0 9 , 2 1 1 , 2 2 0 , 3 9 1 , 408 Lay, C. 561 Leca, F. 1 9 3 , 198 LeCalvez, Y. 2 8 3 , 2 9 9 , 4 2 3 - 2 7 , 4 3 4 , 4 4 9 - 5 0 , 4 6 2 - 6 3 , 517 Lee, H. Y. 9 2 , 105 Legault, J. A. 9 5 , 105 Lejeune, Carpentier 383 Lentin, J. K. 3 7 2 , 3 8 3 , 385 Leschik, G. 200 Lesta, P. J. 299 Lethiers, F. 6 9 , 7 1 , 82 Leydon, R. J. 1 8 5 , 200 Liebau, A. 2 6 4 , 268 Lima, E. 2 8 1 , 2 9 9 , 4 1 3 , 434 Lindstrom, M. 9 0 - 9 1 , 93 9 4 , 1 0 5 - 0 6 , 114 Ling, H. Y. 5 4 9 , 551 Lipps, J. H. 5 4 5 , 551 Lister, T. R. 1 3 8 - 3 9 , 145 Loeblich, A. R. Jr. 3 9 4 , 3 9 8 , 551 Loeblich, A. R. I11 5 4 6 , 5 4 9 , 551 Loeblich, L. A. 551 Lohmann, G. P. 3 9 6 , 408-09 Loranger, D. M. 6 7 , 7 0 , 82 Lord, A. 2 3 6 - 3 7 , 241 Lorenz, J. 1 8 8 , 200 Lugardon, B . 356 Lund, R. 200 Lundin, R. 6 3 , 82 Luterbacher, H. 2 0 6 , 2 1 0 , 2 1 6 , 2 1 9 , 220 Luterbacher, H. P. 3 4 6 - 4 7 , 3 6 2 , 367
Mabesone, J. M. 2 8 2 , 2 9 9 , 4 1 4 , 434 Macedo, A. C. 517 Maddocks, R. F. 5 0 8 , 5 1 3 , 517 Madeira, M. L. 4 5 3 , 462 Maedler, K . 200 Magloire, L. 1 4 0 , 1 4 2 , 144-45 Maher, J. C. 2 0 6 , 2 2 0 , 3 9 2 , 408 Malloy
384
Malumian, N.
281, 416, 418-21, 434, 4 4 7 , 4 6 3 , 517 Malz, H. 2 3 4 , 2 3 6 , 241 Mandra, H. 5 4 6 , 548-49 Mandra, Y . T. 5 4 8 - 4 9 , 551 Mannil, R. 1 5 9 , 166 Manspeizer, W. 1 8 6 , 1 9 1 - 9 3 , 1 9 8 , 200 Manum, S . 3 7 1 , 384 Margerel, J. P. 4 0 2 , 408 Marie, P. 4 2 4 , 4 3 4 Marliere, R. 2 8 6 , 299 Martinez-Machiavello, J. C . 1 4 1 . 145 Martin, F. 1 3 7 - 3 9 , 145
Martini, E. 5 3 6 , 5 4 6 - 4 9 , 551-52 Masiuk, V. 2 8 1 , 4 1 7 - 1 8 , 4 2 0 - 2 1 , 4 3 4 , 473 Masoli, M. 5 0 8 , 517 Masumov, A. S . 2 3 4 , 2 3 6 - 3 7 , 241 Matthews, S . C . 8 6 , 106 Mattis, A. 1 9 1 , 200 Maurrasse, F . 5 5 4 , 5 5 6 , 561 Maxwell, A. E. 3 1 7 , 4 2 8 - 2 9 , 4 3 4 , 457-58, 463 Mazzoni, M. M. 417 McDonald, N. G. 198 McElhinny, M. W. 8 9 , 106 McGill, P. 6 4 - 6 5 , 6 7 , 82 McGuire, O . S . 82 McIntyre, A. 4 0 1 , 408 McIntyre, D. J. 3 7 1 , 384 McKenzie, K. G. 4 9 7 , 517 McLearn, F . H. 4 McMillan, N. J. 65 McMillen, K. J . 535 McMilland, G. V. 81 McNeely, B. 3 2 8 , 5 5 7 , 561 Mehas, K. 517 Meijer, M. 4 2 4 , 434 Mello, J. F. 2 0 6 , 220 Mendez Bedia, I. 7 9 Mendez, I. A. 4 1 7 , 434 Menendez 3 7 1 , 384 Meuter, F. J. C. de 4 0 1 , 407 Michel, M. Ph. 6 4 , 6 6 , 6 9 , 7 5 , 8 0 , 83 Michelsen, 0 . 2 3 3 - 3 4 , 241 Micholet, J. 2 8 3 - 8 4 , 2 8 9 - 9 0 , 2 9 8 , 4 5 0 , 462 Minard, J. P. 2 2 0 , 384 Miller, J. A. 406 Miller, J. F. 8 7 , 9 1 , 1 0 3 , 106 Millioud, M. E. 3 4 6 , 367 Milow, D. 318 Missarzhevsky, V . V . 8 7 , 106 Moguilevsky, A. 4 9 6 , 518-19 Mohler, H. P. 5 Moore, T. C . 3 1 1 , 3 1 8 , 4 0 1 , 407 Moores, E. PI. 1 3 0 , 133 Moorley, A. 2 3 6 - 3 7 , 241 Morgenroth, P. 3 3 0 , 338 Mosher, L. C. 1 2 4 - 2 5 , 132 Moullade, M. 2 5 3 , 268 Mound, M. 9 1 - 9 2 , 106 Moura, 3 . A. 2 6 1 , 281 Muller, G. W. 517 Muller, K. J. 8 7 , 106 Murphy, M. A. 1 7 0 , 179 Murray, G. E. 391-92, 408 Murray, J. 3 0 5 , 318 Hurray, J. W. 3 9 5 , 4 0 7 , 409 Musacchio, E . 2 8 0 - 8 1 , 299
574 Naeser, C. W. 560 Nazarov, B. B. 169, 172-73, 175,
179 Neale, J . W. 2 5 0 , 2 6 5 , 2 6 8 , 517 Nelson, W. G. 4 2 8 , 433 Neuweiler, F . 241 Newport, R. L . 1 5 3 , 166 N i c o l l , R. S . 9 5 , 1 0 6 , 1 2 9 , 132 N i g r i n i , C . A. 1 7 3 , 1 7 6 , 179,526,535 N i t e c k i , N. H. 1 7 3 , 1 7 6 , 179 Nogami, Y . 8 7 , 106 Noguti, I. 4 1 4 , 4 1 6 , 4 1 9 , 4 2 0 - 2 1 ,
4 3 4 , 4 4 5 , 4 5 8 , 463 N o l t i m i e r , H. C . 8 8 - 8 9 , 106 Noordermeer, E . J. 517 N o r r i s , G . 3 5 7 , 367 Odin, C. S. 5 5 8 - 5 9 , 561 O e r t l i , H. J. 2 3 5 , 2 3 7 , 2 3 9 , 2 4 1 ,
2 5 3 , 268 O'Grady, M. D. 2 0 6 , 2 1 1 , 220 Ohmert, W. 2 6 2 , 269 O'Keefe, J. A. 5 5 4 , 561 O r n e l l a s , L . P. de 518 Olsen, W. 1 8 5 , 200 Olson, E . C . 201 Olsson, R. K . 2 0 6 , 2 1 1 , 2 2 0 , 3 8 3 ,
P i n t o , I. D. 2 8 7 , 3 0 0 , 518 P i s e t t a , J . L . 4 4 7 , 464 Pitman, W. C . 1 8 5 , 1 9 9 , 2 0 1 , 2 9 1 ,
408 Ploch, R. A. 6 7 , 81 Pocok, R. 3 3 0 , 338 Pokorny, V. 6 5 , 235, 237, 241, 259,
2 6 9 , 518 P o l l a r d , J . 7 5 , 8 0 , 83 P o l l o c k , C . A. 9 5 , 106 Pomerol, C . 3 9 2 , 408 Poore, R. Z . 4 0 0 , 408 Popov, L. E. 179 P o r t h a u l t , B. 262 P o t t e r , C . A. 9 1 , 106 P o u l s e n , V. 8 7 - 8 8 , 107 P r i b y l , A. 6 3 , 6 6 , 7 6 - 7 7 , 83 P r i c e , P. B. 558-60 P r o u s t , F. 1 9 3 , 200 P u f f e r , J . 1 8 6 , 1 9 2 , 200 P u r i , H. S . 4 0 2 , 4 0 9 , 4 9 6 , 518 Ramirez, F. C. 518 Ramsbottom, W. H. C.
7 5 , 83, 176, 178-79 Rauscher, R. 1 4 1 , 145 R e i f , W. E. 176 Reinhold, T. 4 0 2 , 407 3 9 4 , 408 Omatsola, M. E. 4 9 7 , 5 0 3 , 5 0 8 , 517 Remy, W. 201 O ' N e i l , J . R. 560 Renz, 0 . 3 4 6 , 367 Opdyke, N. D. 8 9 , 1 0 6 , 5 2 6 , 535 Repetski, J . E . 9 1 , 107 O r i e l , S . S . 6 5 , 84 R e r a t , D. 4 2 3 - 2 7 , 4 3 4 , 4 4 9 - 5 0 , 462463 Orr, W. N. 131 Rexroad, C. B. 9 5 , 1 0 3 , 1 0 7 , 131 Ostenso, N. A. 1 2 8 , 132 Repent, E. 518 Oswald, D. H. 83 R e p e n t , R. A. 283, 290-95, 300, Owen, H. G . 2 3 1 - 3 2 , 241 422-23, 425-26, 4 3 5 , 448-49, 4 5 5 , Owens, J . P. 2 2 0 , 384 4 6 4 , 4 9 6 , 5 0 5 - 0 6 , 518 Rhodes, F. H. T. 9 4 , 1 0 7 , 118, 120Packham, G. H. 535 2 1 , 132 P a d g e t t , G . 198 Rickard, L. V. 131 P a k i s e r , H. M. 3 5 5 , 3 6 2 , 3 6 5 , 367 R i e d e l , W. R. 173, 177, 179, 311, Papp, A. 5 4 6 , 5 4 9 - 5 0 , 552 318, 4 0 6 , 524, 526-29, 535-36, P a s c u a l , R. 4 5 4 , 463 5 4 8 , 5 5 2 , 5 5 7 , 561-62 Paulson, 0 . L. Jr. 2 5 9 , 262 R i g a s s i , D. 3 1 7 , 516 P e n i g u e l , G. 144 Riggi, J. C . 4 1 8 , 4 2 0 - 2 1 , 4 3 4 , 4 4 7 , P e r l m u t t e r , N, M. 2 0 6 , 221 Pernjakova, M. N , 2 3 4 , 241 463 P e r r y , W . J. 3 5 5 , 3 6 2 - 6 4 , 367 Riva, 3. 104 Pessagano, E. A. J r . 2 0 6 , 2 2 1 , 3 1 1 , Robb, J. M. 201 Robbins, E . I. 3 5 5 , 3 6 2 - 6 4 , 367 318 P e t e r s o n , M. N. A. 2 0 6 , 221 Robinson, J. E. 7 1 , 8 3 , 241 6 5 , 81 P e t e r s o n , R. M. Rocha, A. T. 4 5 1 , 464 P e t r i , S. 281, 300, 443-45, 464 Rodriguez, F. B. H. 463 Petrushevskaya, M. G . 3 1 8 , 5 3 1 , 535 R o s s i . U. 458. 461 P e t t e r s , S. E. 2 0 6 , 2 0 8 - 0 9 , 2 2 1 , 384 R o s s i . d e G a r c i a , E . 241, 281, 498, P h i l l i p s , 3. D. 185, 201, 391, 394, 518-19 3 9 7 , 4 0 2 , 406 Roth, P. H. 318
5 75 Rudiman, W. F. 4 5 9 , 464 Rushton, A . W. A. 8 7 , 106 R u t t e n , M. G . 3 9 2 , 409 Ryan, W. B. F. 185, 1 9 9 , 5 1 3 , 536 Sachs, K. N. 534 S a i t o , T. 3 1 8 , 4 2 8 , 4 3 5 , 4 5 6 , 4 6 4 ,
536 Sandberg, C. A . 131-32 Sanchez de Posada, L. 6 6 , 6 8 , 7 1 , 7 6 ,
7 9 , 83 311, 318, 406, 524, 526-29, 535-36, 547-48, 552, 558, 561-62 S a n g u i n e t t i , Y . T. 2 8 7 , 3 0 0 , 518 S a r j e a n t , W. A. 3 3 4 , 3 3 7 , 3 7 4 , 38384 Saunders, J . €3. 559 S a v i n , S . M. 3 9 4 , 3 9 8 , 409 S c h a l l e r , H. 273-74, 281-82, 300, 4 1 4 , 4 1 8 - 2 0 , 4 3 5 , 4 4 3 - 4 4 , 464 Schenk, P. E. 201 S c h e u r i n g , B. W. 2 0 1 , 3 1 9 , 338 Schink, D. R. 534 Schmid, M. 1 3 8 , 143 Schmidt, R. A . M. 2 6 3 , 269 Schonlaub, H . P. 9 4 - 9 6 , 1 0 5 , 107 Schopf, T. J. M. 9 2 , 107 S c h r a d e r , H. J. 406 S c h u l t z , G . 271 Schumacher, D. 201 Schwalb, H. 1 5 5 , 164 S c o t t , Th. 7 3 , 519 Seddon, G . 131 S e i b o l d , E . 317 Sergeeva, S . P. 9 3 , 107 Serpagli, E. 8 7 , 9 3 - 9 4 , 1 0 5 , 107 S h a c k l e t o n , N. J. 3 9 4 , 3 9 8 , 407 S h a v e r , R. H. 7 3 , 83-84 S k e v i n g t o n , D. 101, 107 S l i t e r , W. V. 2 0 1 , 211 S l a n s k y , M. 4 2 2 , 4 2 5 , 4 3 5 , 4 5 5 , 4 6 4 , Smith, A. G. 1 2 8 , 1 3 2 , 178,197 Smith, S . 7 3 , 8 3 , 113 Snead, R. G . 384 S o h l , N. G . 384 Sohn, I. G . 7 2 , 8 3 , 201 S p a l l e t t i , L . A . 417 S t a e s c h e , K. 3 9 8 , 409 S t a i n f o r t h , R. M. 3 9 7 , 409 S t e h l i , F . G . 409 S t e p h e n s , D . G . 198 S t e w a r t , G . A. 6 7 , 84 Stockmans, F . 1 3 8 , 141-42, 145 S t o l k , J. 4 2 3 , 4 2 6 , 435 S t o r z e r , D. 5 5 8 , 560-62 S t e v e r , L. E. 384 Sanfilippo, A.
S t r a d n e r , H. 5 4 9 - 5 0 , 552 S t r a k a , J.J. 111 132 S t r e e l , M. 79 Stumm, E . C . 6 5 , 84 S t u r n e r , W. 1 7 3 , 179 Suarez Soruce, J . R. 4 5 4 , 464 S v e r d l o v e , M. S . 347, 367, 372, 374,
384 6 5 , 8 4 , 239, 241-42, 2 5 6 - 5 8 , 2 6 3 , 2 6 9 , 3 0 0 , 406 Sweet, W. C . 8 4 , 8 7 , 9 1 - 9 2 , 9 4 , 1 0 2 , 1 0 4 , 1 0 7 , 1 1 3 , 1 2 4 , 132 S y l v e s t e r - B r a d l e y , P. C . 513 Szaniawaki, H . 8 7 , 107 Swain, F . M.
Tabor, N. L . 6 5 , 8 1 Talwani, M. 185, 201, 391, 399,
408-09 Tappan, H. 3 9 4 , 4 0 8 , 551 Tasch, P. 2 0 1 , 1 5 3 , 166 T a t r o , J . 0 . 2 6 3 , 266 Taugourdeau, P . 1 5 4 , 1 5 6 - 5 7 , 1 5 9 ,
1 6 2 - 6 4 , 166 T a y l o r , M. E . 1 0 1 , 1 0 4 , 1 2 9 , 131 Teeter, J. W. 5 0 9 , 519 Theyer, F. 5 2 6 - 2 7 , 5 2 9 , 536 T h i e r s t e i n , H. R. 3 1 8 , 3 4 6 , 3 4 8 - 5 0 ,
3 6 0 , 367 T h i e s o n , 2. V. 4 4 5 - 4 6 , 464 Thomas, T . M. 1 2 2 , 132 Thompson, M. J. 84 Thompson, T o L . 131 Thorez, 3. 7 2 , 79 Tinoco, I . de M. 2 8 2 , 299-300, 4 1 4 ,
4 3 4 - 3 5 , 4 5 3 , 464 T j a l s m a , R. C. 3 9 6 , 4 0 1 , 4 0 6 , 408-09 Todd, R . 2 0 6 , 2 2 1 , 4 2 0 , 435 T r a v e r s e , A . 1 9 8 , 3 2 9 , 337 Tschigova (Chizhova), V . A . 71 Tucholke, B. 2 0 6 , 2 1 0 , 221 Uchupi, E . 1 8 5 , 1 9 7 , 3 9 1 , 407 U d i n t s e v , G . 3 9 9 , 409 Ulleberg, K. 3 9 4 , 409 U l r i c h , B. C . 220 U l r i c h , E . 0 . 3 , 7 , 6 3 , 84 Umnova, N . 166 Upshaw, C . E . 384 Urban, J. B . 1 5 3 , 166 Uyene, T . T. 132 Vachey, G. 144 V a l e n s i , L. 384 V a l l e n t i n e , J . W. 1 3 0 , 133 van Couvering, J . A . 534 Van de P o l l , H . W. 201, 256, 269 van der Lingen, G. J . 535
576
Vangerow, E. F. 7 5 Van Hinte, J. E. 2 0 6 , 2 1 0 , 2 1 9 , 2 2 1 , 3 6 0 - 6 1 , 3 6 6 , 3 9 8 , 407 Van Houten, F. B. 1 8 6 , 201 Van Valen, L. 518 Van Wamel, W. A . 9 3 , 108 Vasconcelos, D. N. 3 0 0 , 4 1 3 , 4 1 8 , 4 2 0 , 4 3 4 , 464 Vavdrova, M. 1 3 7 , 145 Verdenius, J. G . 4 0 1 , 409 Verdier, J. P. 3 4 6 , 3 6 6 - 6 7 , 3 7 1 , 382 Vernier, G . 433 Verrier, G . 2 8 8 , 2 9 0 - 9 1 , 2 9 8 , 4 2 4 , 4 2 7 , 4 5 1 , 462 Viana, C . G . 2 7 7 - 7 8 , 2 8 4 , 300 Viira, V. 9 3 - 9 4 , 107 Voorthuysen, J. H. van 4 0 2 , 409 Vozzhennikova, T. F. 3 7 1 - 7 4 , 384 Wade, J. A . 2 0 0 , 3 3 5 - 3 6 , 338 Wagner, C . W. 517 Wagner, G. A . 5 5 8 , 560-62 Walker, R. M . 560 Wall, D. 383-84 Walliser, 0 . H. 9 5 , 107 Walmsley, V. G . 9 5 , 107 Weaver, F. M. 5 3 2 , 5 3 6 , 5 4 8 - 4 9 , 5 5 1 , 554 Warren, J. S. 383 Webers, G . F. 9 2 , 108 Webster, G . D. 132 Weiss, M. 6 5 , 82 Wenger, R. 2 7 4 , 2 8 2 , 299 Weyant, M . 6 4 , 84 Whatley, R. C . 2 3 5 , 2 4 2 , 4 9 6 , 518 Whittington, H. B. 101, 108 Wicander, E. R. 1 4 2 , 145 Wienholz, E. 242 Wilcoxon, J . A . 3 4 6 , 367 Williams, A . 101, 108 Williams, G . L. 3 2 7 , 3 3 4 - 3 5 , 3 3 8 , 3 7 2 , 3 7 4 , 3 8 3 , 385 Willi;re, Y. 1 3 8 , 1 4 1 - 4 2 , 145 Wilson, Ch. W. 6 3 , 84 Wilson, G . J. 3 7 1 , 3 7 4 , 3 7 8 , 385 Wilson, J. T. 101, 108 Wilson, L. R. 1 4 3 , 145 Winkler, P. C. F. 79 Wold, R. J. 1 2 8 , 132 Wolfe, J. A . 3 5 5 , 3 6 2 , 3 6 5 , 367 Wollen, I. D. 198 Wollin, G . 4 7 7 , 481 Wong, T, 3 9 8 , 407 Woodland, A . W. 3 9 1 , 409 Worzel, J . L. 5 5 5 , 5 5 7 , 562 Wright, C . A . 3 9 5 , 408-09 Wright, J. D. 6 5 , 84
Wright, R. C.
5 3 4 , 550
Yochelson, E . L .
201
Zagora, K . 65 Zahringer, J . 5 5 8 , 562 Zarillo, Z. A. 5 4 9 , 551 Zhuze, A . P. 5 4 9 , 552 Ziegler, W. 1 1 6 , 1 1 8 , 132-33 Zwart, M, J. 5 5 4 , 5 5 7 - 5 8 , 562
577
Index to Genera and Species Abathomphalus 2 0 5 , 2 9 4 mayaroensis 2 0 8 , 3 1 6 , 3 9 1 Abyssocypris pykna 505 sp. 505 Abyssocythere 4 9 6 , 509 trinidadensis 505 Acaenictyle umbilicata 312 Acanthaulax paliuros 325 Acanthocythereis 263 hapsida 5 0 4 ret iculospinosa 5 0 4 sp. 2 6 4 sp. aff. A. teiskotensis 4 7 2 spiniferrima 502 Acanthodiacrodium sp. 1 4 6 Acanthodus 90 Acanthoscapha 6 2 - 6 8 Acarinina triplex 4 2 0 Achomosphaera ramulifera 3 4 4 , 352 Acidnomelos proapteron 3 1 3 - 1 5 , 318 Acontiodus 93 Acratia 6 9 , 72 centronotus 7 3 rothi 7 3 Acrocythere 2 3 7 , 2 5 1 , 2 5 4 Actinochilina spp. 24-25 suecica 26 Actinocythereis biposterospinata 5 0 3 davidwhitei 472 exanthemata 4 8 9 indigena 5 0 3 rex 503 stenzeli 4 7 2 Actinomma sp. 5 3 8 Acuticytheretta 2 6 4 Adetognathus 1 2 2 - 2 3 , 127 Adnatosphaeridium aemulum 3 2 4 caulleryi 3 2 4 , 333 Aequitriradites spinulosus 326 Afens liroides 3 1 3 - 1 5 Afrocytheridea 2 3 8 , 240 Agathammina 5 0 , 5 3 , 55 Aga thamminoides 5 1 Agrenocythere 4 7 9 , 4 9 6 antiquata 4 7 9 , 4 9 1 , 5 0 5 , 509 gosnoldi 4 7 9 , 4 9 1 hazelae 4 7 9 , 4 9 1 , 5 0 5 , 5 0 9 Akidobairdia 237 Akidograptus acuminatus 100 Alabamina 3 9 5 , 399 midwayensis 3 9 8 , 435 Alatacythere 2 5 4 , 2 6 3 - 6 4 cirrusa 4 6 8 , 4 8 0 maerkyi 505 Albaillella 1 6 9 , 1 7 5 cf. pennata 1 7 6 pennata 1 7 6 - 7 7
Albaillella spp. 1 7 6 - 7 7 Alievium gallowayi 312-15 praegallowayi 312 superbum 3 1 1 - 1 2 , 315 Alisporites 3 5 1 aff. parvus 197 bilateralis 358 minuto saccus 1 97 Allomorphina 417 aIlomorphinoides 212 Aluta 20 Ambocythere caudata 5 1 1 elongata 505 exilis 505 subreticulata 505 Amicytheridea 2 1 8 , 2 3 8 , 2 4 0 , 250 Armnobaculites 5 3 , 5 5 - 5 6 , 216-17 beveridgei 57 chappelensis 57 gutschicki 57 leptos 57 Ammodiscella 5 1 Armnodiscoides 5 3 - 5 5 Ammodiscus 5 2 , 5 5 , 2 1 4 , 2 1 6 - 1 8 , 399 longexsertus 57 semiconstrictus 57 Ammolagena 55 Ammolegena 5 3 Ammonia beccarii 4 4 7 , 4 5 2 , 4 5 4 - 5 5 beccarii parkinsoniana 4 5 2 - 5 4 Ammovertella 51, 5 3 , 5 5 - 5 6 , 218 lisae 57 pikei 57 Amorphognathus 9 1 - 9 4 ordovicicus 9 9 superbus 9 1 , 99 tvaerensis 9 1 , 99 Amphicervicis 9 1 Amphicythere 2 5 1 Amphicytherura 2 5 8 , 2 6 2 - 6 3 bartensteini 253 chelodon 252 sp. 2 9 4 Amphipyndax enesseffi 2 9 5 , 312-15 stocki 313-15 tylotus 3 0 5 , 3 1 2 - 1 6 , 318 Amphithopalum ypsilon 5 3 8 - 3 9 Amphistegina 4 5 0 , 4 5 3 lessoni 4 5 2 - 5 3 Amphitoxis curvata 34-35 Amphitremoida 5 2 , 55 Amphorula metaelliptica 3 2 5 , 334-35 3 4 4 , 348
Anchignathodus 1 2 2 - 2 4 , 127 sp. 1 1 5 typicalis 1 2 3 , 125 Ancyrochitina aequoris 1 6 3
578 Ancyrodella 118 lobata 115 ploeckensis 97, 100 Ancyrognathus 118 Angoch i1ina ca11awayens is 163 capillata 161 milanensis 161 Angulogavelinella gracilis 212 Angulogerina gracilis 399 Anisochilina 29 Annosacythere 251, 258 Anomalinoides 395, 399 acutus 395 midwayensis 395 nelsoni 211-12 pinquis 211-12 Anthocyrtidium angulare 637 Anticostibolbina jupiterensis 12,14 Anticostiella 5 elisensis 11, 13 pustulosa 12, 14 Anticythereis 283, 286, 495 inconnexa 503 schilleri 503 Antulisporites varigranulatus 326 Aparchites 5 maccoyii 30-31 Ap a tobo 1b ina p 1atyg as ter 33- 35 whiteavesi 12, 14 Apatocythere 252. 257 Apatognathus 120 scalenus 121 Aphelocythere 234 Aphelognathus 92 Apiculatisporis 189 Append ic isporites po tomacensis 357 problematicus 357 Apsidognathus 96 Apteodinium granulatum 344 Apterrinella 51 Aracajuia benderi 274 Aragonia velascoensis 213, 216 Aratrisporites sp. 193, 197 Archaeospongoprunum triplum 282 Archaias angulatus 445, 452-53 Archeoglobigerina 195-96, 199 blowi 196 Archocyrtium 73-76 Arcuaria sineclivula 33 Arenobulimina 214, 216-219 frankei 213 presslii 218 subsphaerica 213 Argilloecia 254, 264 decussata 262 sp. 506 Arikloedenia newbrunswickensis 12,15 Arkhangelskiella cymbiformis 316
Artostrobium urna 312-15 Asanoina 395 Aschemonella 52, 55 Asciocythere 238, 240, 256, 259 perforata 256 rotunda 256 sp. A. 259 Asterocyclina 395 Astrammina 51 Astrononion stelligerum 452 Astrorhiza 51 Asturiella 75-76 limburgensis 76 Atlantopollis 363 sp. 344, 349 Aulacopsis bifissurata 22 monofissurata 22 spp. 21, 24 Aureotesta clathrata 138, 146 Aurila 478, 481 amygdala 504 conradi conradi 489 galerita 504 punctata 506 Australicythere 507, 510 Bactrognathus 120-21 Bairdia 71, 240, 251-52, 254, 257, 263-64, 479, 497 aff. hondurasensis 502 alexandrina 259 amygdaloides 506 anachoretta 503 aselfingensis 237 attenuata 506 cassida 505 cespedeselis is 505 comanchensis 258 denticulata 262 dolicha 502 farinacciae 237 hahni 237 magna 502 montensis 47@ oarion 505 snldadensis 502 sp. 502 sp. aff. ceramensis 506 Bairdiacypris 236 Bairdoppilata 293: 4.73 rotunda 259 sp. 3 511 Balowella 235 Balticella 26, 29 deckeri 26 sp. 11, 13 spp. 25 Baltisphaeridium 142
579
140, 146 simplex 141 B a l t o n i o d u s 94 B a s s l e r a t i a 63 t y p a 11, 1 3 , 25-26 t y p i c a 24 B a s s l e r i t e s 5 0 7 , 510 a b a l i d i e g w u e n s i s 502 s p . a f f . B. b e r c h o n i 506 B a t a v o c y t h e r e 253 g a u l t i n a 254 h i l t e r m a n n i 253 Bathysiphon 5 2 , 5 4 - 5 5 , 2 1 3 - 1 4 , 21618, 398-99 B e e c h e r e l l a 6 3 - 6 5 , 67 Bekoma b i d a r f e n s i s 529 t e t r a d i c a 529 Belodina 92 Bensonia 507 Bergstroemognathus 92 Bernix 72 "Bernix" v e n u l o s a 7 2 B e r t h e l i n e l l a 214 B e r o u n e l l a 6 3 , 6 5 - 6 8 , 76 B e y r i c h i a b a r r a n d i a n a 2 2 , 24 ( E o b e y r i c h i a ) s p p . 34 k l o e d e n i 3 3 , 35 ( N o d i b e y r i c h i a ) p u s t u l o s a 1 2 , 15 p a u c i t u b e r c u l a t a 3 3 , 35 r i n g e r i k e n s i s 3 3 , 35 B e y r i c h i o n a 20 t r i c e p s 24 B e y r i c h i o p s i s 71-72 g l y p t o p l e u r o i d e s 72 Bigenerina 5 3 , 55-56 a r c u a t a 211 j u r a s s i c a 211 B i o r b i f e r a johnewingii 344-45, 349, 3 5 4 , 360-61 Biorbulina b i l o b a t a 451 B i s u l c o c y p r i s 2 3 5 , 2 6 4 , 279 sp. 280 B l a s t a m i n a 5 2 , 54-55 e i s e n a c k i 57 Bohemina 69 B o l b i b o l l i a l a b r o s a 1 2 , 14 p a p i l l o s a 1 2 , 14 Bolbina spp. 25 Bolb i n e o s s i a ( B o l b i n e o s s i a ) p i n e a u l t i 1 2 , 14 ( B r e v i b o l b i n e o s s i a ) berdanae 1 2 , 14 Boldia 383 B o l l i a s u b a e q u a t a 24-26 B o l i v i n a 387 compacta 452 c r a s s i c o s t a t a 426 Baltisphaeridium denticulatum
B o l i v i n a f i n l a y i 446 p l i c a t e l l a 443 p s e u d o p l i c a t a 4 5 2 , 455 s t r i a t e l l a t a 427 s t r i a t u l a 4 5 2 , 454 t o r t u o s a 452 v a r i a b i l i s 4 5 2 , 454 B o l i v i n o i d e s 2 0 5 , 2 0 9 , 2 1 2 - 1 3 , 216 c u l v e r e n s i s 208 d e c o r a t u s 208 d r a c o 208 m i l a r i s 208 s t r i g i l l a t u s 208-09 B o l i v i n o p s i s 387 h e l v e t o j u r a s s i c u s 211 Bonnernaia r u d i s 3 1 , 3 3 , 35 Bosquetina s p . a f f . B. m a r g i n a t o s t r i a t a 506 B r a c h y c y t h e r e 2 5 9 , 263-64 a g u l h a s e n s i s 294 armata 294 durhami 263 ekpo 293 i n t e r r a s i l i s 472 ledaforma 264 ogboni 502 o g u n i 294 p l e n a 472 r h o m b o i d a l i s 265 s a p u c a r i e n s i s 282 s p . g r . ekpo 2 5 9 , 264 s p h e n o i d e s 2 5 9 , 264 t a y l o r e n s i s 262 t r i a n g u l a r i s 472 Bradleya 2 6 2 , 4 7 9 , 4 9 6 , 509 d i c t y o n 505 f o r b e s i 477 h a z a r d i 2 6 0 , 265 v e s i c u l o s a 294 B r a d o r i a 20 B r a d y c y p r i s r o t u n d a 287 Bradyina 5 3 , 55-56 B u c c e l l a f r i g i d a 4 4 7 , 4 5 2 , 454 p e r u v i a n a campsi 4 5 2 , 454 Buccinosphaera i n v a g i n a t a 5 2 6 , 531 B u d n i a n e l l a shenandoense 25-26 Bulimina 399 m a r g i n a t a 455 o r p h a n e n s i s 399 p a t a g o n i c a 454 B u l i m i n e l l a e l e g a n t i s s i m a 454 a c u l e a t a 410 f a b i l i s 2 1 5 , 217 g r a t a 217 B u l l a p o r a 51 B u l l a t e l l a kauffmanensis 2 3 , 25-26 Buntonia 2 6 3 , 4 9 5 . 5 0 7 , 510 alabarnensis 502
580
Buntonia attitogoensis 502 bopaensis 502 fortunana 502 giesbrechti robusta 506 livida 502 pulbinata 502 rocanortensis 503 sp. 502 spp. 293 sublatissima dertonensis 506 vanmorkhoveni 293 Buryella clinata 5 2 9 Bythoceratina 2 6 4 scabra 5 0 5 - 0 6 , 5 1 1 Bythocypris 2 5 7 , 2 6 3 , 4 7 9 , 4 9 7 guppyi 5 0 5 sp. 505 sp. aff. B. bosquetiana 505 Bythocythere costata 503 punctulata 5 0 3 rocana 503 triebeli 503 Cadargasporites verrucosus 326 Calamospora 1 8 9 Calcitornella 5 1 , 5 3 , 55-56 Calcivertella 5 1 Callialasporites dampieri 3 2 6 , 3 3 0 , 335
segmerties 326 Callistocythere 4 9 5 - 9 6 , 5 0 9 canaliculata 4 7 8 , 4 8 9 Calocyclas turris 5 7 2 , 5 8 0 - 8 1 Calocycletta costata 529 Camaros;cTites secatus 326 Camarozonosporites rudis 326 Campylocythere ? perieri 5 0 4 Canartiscus marylandicus 537 Cancris cf. cocoaensis 4 2 6 sagra 4 4 5 Candona 282 daleyi 4 9 1 gregaria 277 huantriacoensis 282 Candorbulina universa 4 5 5 Cannartus petterssoni 529 Cannopilus 546 Carbonita 7 0 , 7 5 , 7 7 Cardobairdia 2 5 1 asynnnetrica 505 glabra 505 ovata 5 0 5 sp. aff. C. minuta 505 Carinobolbina 5 Carinocythereis 5 0 9 Carinokloedenia 63 Carniodus 96 Carpocanium eracatum 538
Carpocanistrum szyz 5 2 9 Cassidella tegulata 215 Cassideus riedeli 312 Cassigerinella chipolensis 4 1 5 , 447
Cassigerinelloita amekiensis 4 2 6 Cassidulina laevigata 4 5 2 - 5 3 laevigata carinata 4 5 0 sublobosa 4 5 2 Catapsydrax dissimilis 4 1 5 stainforthi 4 1 5 Cativella 5 0 7 , 510 ixemojai 508 novis 5 0 4 Caudites 507 Cavellina sp. cf. cumingsi 7 6 Cavusgnathus 1 2 0 , 1 2 7 altus 1 2 1 charactus 1 2 1 convexa 115 naviculus 1 2 1 Centrocythere bordeti 253 Centrocytheridea 264 Cerarchocyrtium 1 7 5 Ceratamina 5 2 , 55 Ceratacratia 6 9 Ceratobulimina parva 215 Ceratocypris longispina 2 2 , 2 4 Ceratokiscum 1 6 8 , 1 7 0 , 1 7 2 , 1 7 4 - 7 5 acatangulatum 172 avimexpectans 1 7 5 bicancellatum 1 7 6 bujugum 1 7 5 planistellare 1 7 5 spinosiarcuatum 1 7 5 tricancellatum 1 7 0 Ceratopsis 5 chambersi 30 quadrifida 11, 1 3 Cerebropollenites mesozoicus 3 2 6 , 329
Chapmanicythereis 256-57 Chatangiella maunumii 3 6 9 , 3 7 4 , 3 7 6 - 7 8 , 3 8 0 , 385
n. sp. 3 6 9 , 3 7 4 , 3 8 0 - 8 1 , 386-88 victoriensis 3 6 9 , 3 7 4 - 7 5 , 3777 9 , 385
vnigrii 3 6 9 , 3 7 2 , 3 7 4 - 7 8 , 3808 1 , 385
Chiloguembelina martini cubensis 426
morsei 4 1 4 Chilostomella 216 trinidadensis 213 Chirognathus 92 Chlamydophorella nyei 3 4 4 , 3 5 1 Choenicosphaera murrayana 538
581
Choffatella decipiens 2 8 8 Chordasporites sp. 1 9 3 , 1 9 6 Chosonodina 9 1 Chrysalogonium 399 Chrysocythere 5 0 7 , 5 0 9 - 1 0 cataphracta 5 0 6 foveostriata 506 hexastriata 5 0 6 Chytroeisphaeridia sp. (cristate) 3 4 4 Cibicides 3 9 5 , 4 0 3 asknerianus 4 5 2 bertheloti 4 4 7 , 4 5 2 - 5 3 dispars 4 5 5 harperi 212 pseudoungerianus 4 4 7 , 4 5 2 - 5 3 spp. 4 1 0 , 4 5 5 tenellus 399 variabilis 4 5 4 Cibicidoides 3 9 9 alleni 3 9 5 hercegovinensis 3 9 9 howelli 3 9 5 mediocris 4 0 2 proprius 3 9 5 succedens 3 9 5 trincherasensis 398 Cicatricosisporites autralensis 3 2 6 , 333
brevilaesuratus 357 hughesii 3 4 4 , 357 potomacensis 3 4 4 , 357 Cinclopyramis sanjoaquinenesis 3 1 3 , 315
Circularaesporites cerebroides 3 2 6 , 332
Citharina 2 1 6 kochi 2 1 0 , 2 1 5 simondsi 2 0 9 , 2 1 4 texana 2 0 9 , 2 1 4 wadei 2 0 9 , 2 1 4 Cladarocythere apostolescui 4 9 9 Classopollis torosus 3 5 7 , 359 Clathrocanium diadema 538 Clathrocorys sp. 538 Clathropyrgus bumastus 3 1 3 , 3 1 5 titthium 3 1 3 - 1 5 Clavatipollenites 3 4 4 - 4 5 , 3 5 5 - 3 7 ,
Clithrocytheridea 2 5 1 , 2 5 4 , 256-57 garretti 4 7 2 magna 4 7 0 sp. 502 sp. aff. C. tombigbeensis 502 Cnestocythere truncata 9 9 , 4 7 8 Cobanocythere 5 0 9 Coleodus 92 CollOsPhaera tuberosa 5 3 7 , 5 3 9 Colonammina 5 2 , 5 5 Complexiopollis 3 4 4 - 5 6 , 3 5 8 - 5 9 , 3 6 1 , 363-64
sp. 3 4 4 , 3 5 9 Conbathella biporata 1 2 , 14 equilateralis 1 2 , 14 inornata 1 2 , 14 Conchoecia 2 5 1 Conchoides minuta 2 1 , 2 2 , 2 4 Conchoprimitia eos 2 4 leperditioidea 26 Concnoprimites 2 6 spp. 2 5 Conochitina 1 5 1 , 1 5 5 , 1 6 3 cactaceous 1 6 1 dactylus 1 5 5 elegans 161 hirsuta 1 6 2 micracantha 1 5 5 , 1 6 2 minnesotensis 1 5 5 robusta 1 5 5 , 1 6 2 wesenbergensi 162 Conradidium firmamentum 1 4 2 , 1 4 7 Contignisporites cooksonii 3 3 5 problematicus 3 3 0 sp. 3 2 6 , 3 3 0 , 338 Convolutispora 1 8 9 klukiforma 3 2 6 , 329 Coquimba congestocostate 5 0 4 Corbisema 546 Cordaitina 1 8 9 Cordosphaeridium costatum 3 2 4 , 33234
Cordylodus 9 1 , 9 3 , 99 angulatus 9 1 Coriac ina coriacea 2 7 8- 7 9 Cornicythereis 2 5 6 Cornigella 7 5 363-64 Cornikloedenia 7 Cornuspira 5 3 , 5 5 - 5 6 hughesii 3 5 5 - 5 7 , 3 6 1 Cornutella profunda 532 SP. 3 4 4 , 3 5 6 - 5 7 Corollina meyeriana 3 2 4 - 2 9 , 336 tenellis 357 torosus 3 2 6 , 3 2 8 - 2 9 , 335 Clavigerinella 4 2 5 Clavulina 216 Coronatispora valdensis 326 Coronifera oceanica 3 4 4 trilaterus 2 1 1 - 1 4 Cleistosphaeridium tribuliferum 3 2 4 , Coryphidium elegans 1 3 8 , 1 4 6 Corythoecia 1 7 4 , 1 7 6 351 Climacamina 5 1 , 5 3 , 5 5 - 5 6 Coryphostoma 2 0 5 incrassata 2 1 3 - 1 4
582
Coryphostoma plaitum 211-12 Costa 502 barracoloradensis 504 bellipulex 504 cubana 504 dohmi 504 dahomeyi 502 hasenbankae 503 kugleri 504 maquayensis 504 meka 504 praedohmi 504 robinsoni 504 ruggierii 506 ._ santacrucensis 504 stokesae 504 tricostata 478 variabilacostata recticostata 504 variabilocostata variabilocosta ta 504 walpolei 504 Costaveenia 262 Couperisporites jurassicus 3 2 6 , 335 Craspedobolbina armata 3 3 , 35 (Mitrobeyrichia) boltoni 1 2 , 14 percurrens 3 3 , 35 variolata 3 3 , 35 Criboconcha 70 Cribrohantkenina inflata 429 Cribroperidinium muderongensis 351 Cribrorotalia sp. 436 Cribrostomoides 3 9 9 , 410 Cribrostomum 51 Crithionina 51 Croneisella 51 Crycella espartoensis 312 Cryptophyllus 7 1 Ctenentoma plana 2 2 , 24 Ctenidodinium 334 aff. tenellum 3 2 4 , 330 continuum 324 culmulum 3 2 5 , 334-35 ornaturn 3 2 4 , 3 3 2 , 334 pachydemum 3 2 4 , 330 panneum 3 2 4 - 2 7 , 334-35 schizoblatum 325 sp. A. 324 Ctenobolbina 29 ciliata 30 minor kuckersiana 26 ornata latimarginata 26 spp. 25 Cubotholus cf. rhombicus 538 Cucullograptus scanicus 100 Curfsina 2 5 8 , 2 6 2 , 265 communis 2 6 0 , 265
344,
Curfsina nuda 262 Curtognathus 92 Gushmanidea 497 mayeri 472 Cyamocytheridea 497 felix 503 punctatella 477 Cyathochitina 151, 158 alata 158 calix 158 campanulaeformis 1 5 5 , conica 157-58 cylindrica 156 dispar 158 dispar yerrucata 158 d jadoensis 158 157-58
158
_I
fistulosa 158 fusiformis 158 hymenophora 158 hymenophora nigarica 158 infundibulifonnis 1 5 6 , 158 koumeidaensis 158 kuckersiana 158 novempopulanica 158 obtusa 158 regnelli 158 stentor 157-58 striata 1 5 6 , 158 Cycadopites 189 cf. jansonii 326 deterius 326 sp. A. 3 2 6 , 3 2 9 , 338 sp. B. 3 2 6 , 3 2 9 , 338 subgranulosus 3 2 4 - 2 7 , 329 Cyclamina 41C, 417 Cyclogranisporites 189 Cycolgyra involvens 454 Cyclonephelium distinctum 3 4 4 , 349 Cyclotron 20 Cymatiosphaera velicarina 1 4 2 , 147 Cypridea 2 3 6 , 2 3 8 , 2 4 7 , 2 5 0 - 5 1 , 270 aculeata 248 ambigu: 279 bapounon 287 cf. C. primaria 279 clavata 248 (Cypridea) hilariensis 290 dequeenensis 256 diminuta 2 5 6 , 286 dorsispinata 248 dromedarius 277 granulata fasciculata 248 hollensis 287 kegeli 277 loango 287
583
Cypridea (Morinina) bibulluta 2 7 3 , 278-79 (Morininoides) candeiensis 2 7 3 , 2 7 7 , 279 nanorostrata 284 opifera 279 paulsgrovensis 248 (Pseudocypridina) fabeolata 279 riojoanensis 279 (Sebastianites) fida 2 7 3 , 279 (Sebastianites) fida fida 278 (Sebastianites) fida minor 279 (Sebastianites) sostensis sostensis 278 setina 2 4 8 spinigira 248 tenuis 248 wyomingensis 256 Cyprideis 2 6 4 , 4 7 8 , 4 9 5 - 9 7 locketti 5 0 4 ovata 504 pascagoulaensis 5 0 4 salebrosa 5 0 4 similis 508 sp. aff. C. ovata 5 0 4 sp. aff. C. torosa 506 Cypridelliforma 7 5 Cypridina 7 5 , 2 5 1 radiata 7 3 CypridoLJs i p 264 Cyptopholobus 3 4 Cyrtentactinia 168 primotica 1 6 8 , 1 7 4 Cyrtisphaeractenium 1 7 5 Cyrt isphaeronemium 1 7 5 Cyrtocapsella tetrapera 5 2 9 , 537 Cyrtograptus ellesae 100 linnarssoni 100 lundgreni 100 murchisoni 100 rigidus 100, 1 7 0 Cyrtopera laguncula 5 3 2 , 537 Cystograptus vesiculosus 100 Cythere cornuta var. gulfensis 262 foersteriana 2 6 2 Cythereis 2 5 1 , 2 5 3 - 5 4 , 2 5 6 - 5 9 , 2626 4 , 282 acuticostata 254 austinensis 2 6 2 - 6 3 bartensteini 254 bekumensis 253 bicornis 259, 2 6 2 - 6 3 buchlerae 254 burlesonensis 258 cf. bicornis 259 dallasensis 2 6 2 - 6 3 deltaensis 5 0 2 eaglefordensis 258-59
Cythereis geometrica 253 hannai 259 indocilis 5 0 3 ivii 2 6 5 marssoni 2 5 9 ornatissima 259 paraustinensis 259 pidgeoni 265 praeornata 256 roanokensis 258 rug0sissima 6 3 sp. A. 2 8 9 teiskotensis 502 worthensis 258 Cytherella 2 3 6 , 2 4 0 , 2 5 1 - 5 2 , 2565 9 , 2 6 2 , 2 8 2 , 4 7 9 , 497 austinensis 2 5 8 , 2 6 2 , 293 australamericana 503 consueta 4 7 3 excavata 4 7 2 guasarensis 5 0 2 kellettae 5 0 2 lagenalis 4 7 0 montensis 4 7 0 'munsteri' 259 navarroensis 263 navetensis 5 0 5 obliquirugata 262 sp. 5 0 2 sp. aff. C. bullata 5 0 3 sp. aff. C. postdenticulata 506 s p . aff. C. pulchea 506 sp. aff. C. sordida 506 sp. aff. C. utilis 5 0 3 s p . aff. C. vulgata 506 tubulifera 260 Cytherelloidea 2 3 6 , 2 5 1 - 5 2 , 2 5 4 , 257-59, 262-3, 482 dalbyensis 253 damariacensis 4 7 3 , 487 montgomeryensis 4 7 2 , 487 nanafaliensis 4 7 2 ovata 252 patagonica 5 0 3 sp. 502 spirocostata 503 Cytheretta 4 7 8 , 4 8 2 alexander 4 8 7 alexanderi 4 7 2 , 477 arrugia 502 bambruggensis 4 7 4 burnsi 4 8 9 carita 4 7 4 cellulosa 4 6 8 , 4 7 4 - 7 5 , 4 8 1 , 4 8 5 costella costellata 4 7 4 costellata antecalva 4 7 4 costellata costellata 4 8 4
584 C y t h e r e t t a c o s t e l l a t a c r a t i s 485 c o s t e l l a t a g r a n d i p o r a 474 e o c a e n i c a 4 6 8 , 4 7 3 , 4 8 1 , 485 f o r t i c o s t a 4 7 3 , 485 g e o u r s e n s i s 489 haimeana 4 7 4 , 485 howei 477 i n a e q u i v a l v i s 489 l a t i c o s t a 468, 474-75, 477, 481,
485 n. s p . 468 n. sp. a f f . eocaenica 481 n . s p . c f . l a t i c o s t a 480 n e r v a m o n t e n s i s 470 n e r v a n e r v a 4 6 8 , 4 7 0 , 480 o r t h e z e n s i s 489 p o r o s a c o s t a 4 7 7 , 485 " p r e - e o c a e n i c a " 474 r i g i d a r i g i d a 468 s a h n i 489 s c r o b i c u l o p l i c a t a 4 7 3 , 4 8 0 , 485 t e n u i p u n c t a t a 477 t e n u i p u n c t a t a a b s o l u t a 489 C y t h e r i d e a 2 5 7 , 2 7 4 , 482 a f f . r i g i d a r i g i d a 474 bronnimanni 504 i n t e r m e d i a 4 7 5 , 485 k o l l m a n n i 504 m e d i a t e n a 504 newburyensis 473 p e r n o t a 4 7 7 , 485 p r i m i t i a 4 7 3 , 485 r i g i d a r i g i d a 4 7 4 , 4 8 1 , 485 Cytheromorpha 264 a i l l y e n s i s 470 a f f . a i l l y e n s i s 471 b u l l a 491 C y t h e r o p t e r i n a 257 t r i e b e l i 250 C y t h e r o p t e r o n 2 4 0 , 2 5 7 , 2 6 2 - 6 3 , 282 c o r y e l l i 265 l i o g l u m a 503 mexicanum 505 oba 502 rocanum 503 s p . 259 s p . A . 2 8 9 , 506 s p . a f f . C . d i v i u s 502 s p . a f f . C . hamatum 506 s p . B. 289 t h i l i e n s i s 470 t r i n i d a d e n s i s 505 C y t h e r u r a 264 C y t r o c a p s e l l a t e t r a p e r a 538 D a c t y l o f u s c a maranhensis 1 4 1 , 146 Dahomeyella 5 0 7 , 509 a l a t a 502 Dalmanitina 31
Damonella ? t i n k o u s s o u e n s i s 287 Dapsilodus 94 Darwinula 280 kwangoensis 290 l e g u m i n e l l a 287 oblonga 287 Daucina ermaniana 448 Daucinoides c i r c u m t e g e n s 448 D a v i e s i n a 395 D e f l a n d r e a andromedensis 373 b a k e r i i 3 6 9 , 3 7 5 - 7 7 , 389-91 d i e b e l i 373-75, 378, 389, 391,
395 344-45, 347, 353, 354, 3 6 0 , 3 7 2 , 3 7 4 - 7 5 , 3 7 7 - 7 8 , 3 9 1 , 395 minor 3 6 9 , 3 7 5 , 3 7 7 - 7 8 , 3 9 1 , 395 oebisfeldensis 369, 373, 376, 378, 3 8 9 - 9 1 , 395 p h o s p h o r i t i c a 373 v e s t i t a 344-45, 347, 350, 352,354 3 5 6 , 360 D e n s o i s p o r i t e s v e l a t u s 3 3 3 , 3 2 6 , 335 Deroocythere 262 Desmochitina l a t a 162 minor 162 s c a b r o s a 161 Desmospyris s p o n g i o s a 537 D e u n f f i a 139 e i s e n a c k i 139-40 eoramusculosa 140 f u r c a t a 1 4 0 , 146 monospinosa 140 Diacanthum h o l l i s t e r i i 3 4 4 , 350 D i c e l l o g r a p t u s anceps 99 complanatus 9 9 D i c h o d e l l a 94 Diconodinium firmum 3 6 9 , 3 7 4 , 3767 8 , 3 8 9 - 9 1 , 395 D i c o t e t r a d i t e s s p . 360 Dicranella 5 m a r g i n a t a 11, 13 D i c r a n o g r a p t u s c l i n g a n i 99 Dictyocha 5 4 5 - 4 6 , 548 Dictyomitra duedecimcostata 313, 315 k o s l o v a e 3 1 3 , 315 l a m e l l i c o s t a t a 3 1 3 , 315 pseudomacrocephala 3 1 3 , 315 somphedia 312 v e n a t a 312 Dictyonema f l a b e l l i f o r m s 99 Dictyophimus c r i s i a e 538 Dictyophyllidites equiexinus 326, 335 Didymograptus " b i f i d u s " 9 9 e x t e n s u s 99 hirundo 99 m u r c h i s o n i 99 echinoidea
585
Dielymella 2 0 Diexallophasis absona 1 4 2 , 1 4 7 cuspidus 1 4 2 Digmocythere russelli 4 7 2 , 505 Dingodinium cerviculum 3 4 4 , 3 4 9 , 3 5 1 jurassicum 3 2 5 , 3 3 4 Dinogymnium acuminatum 3 4 4 , 353 Diplofusa gearlensis 3 4 4 , 3 5 3 Diplograptus multidens 9 9 Disaccites indet. 1 8 9 Discoaster barbadiensis 557 saipanensis 557 Discocyclina 4 2 3 , 4 2 5 gr. aguerreversi 4 2 3 pratti 4 2 8 sp. 4 2 6 varians 4 2 8 Discoidella 7 1 Discopulvinulina hertheloti 4 5 5 Discorbis floridanus 4 5 6 minutissima 2 1 0 , 2 1 8 mira 4 5 2 obtusa 4 5 2 peruvianus 4 5 5 Discorbinopsis aguayoi 4 5 5 Distephanus fibula 5 4 5 - 4 8 speculum 548 Distobolbina 5 Distobolbina teicherti 11, 1 3 Distomodus 96 Dizygopleura 67 acuminata 3 2 , 3 3 , 35 sp. 1 2 , 1 5 Dolerocypris kinkoensis 2 9 1 Dolocythere rara 253 Dolocytheridea 2 5 2 , 2 5 4 , 2 5 6 - 5 8 atlasica 2 5 8 bosquetiana 2 5 4 , 256 crassa 258 intermed ia 2 53 - 5 4 intermedia intermedia 2 5 4 spp. 252 wolburgi 252 Domasia 1 3 9 elongata 1 4 0 , 1 4 7 trispinosa 1 4 0 Dorcadospyris alata 5 2 2 , 5 2 4 , 5 2 9 ateuchus 5 2 9 dentata 529 Dordoniella 2 5 8 Dorothia 2 0 5 , 2 1 4 , 2 1 6 , 217 bulletta 2 1 3 ellisorae 2 1 3 .oxycona 2 1 3 praehauteriviana 2 1 0 , 218 retusa 2 1 3 Drepanellina clarki 7 , 3 2 - 3 3
Drepanodus 3 5 , 93 Drepanoistodus 92 9 3 Druggidium apicopaucicum 3 4 4 - 4 5 ,
-
348, 351, 354, 361 344-45. 349-51. 354. 361 rhabdoreticulatum 3 4 4 - 4 5 , 3 4 8 - 4 9 , 350, 354, 360-61 Druppatractona 3 1 5 , 318 sp. A . 313 Dumontina 2 5 8 , 2 6 2 , 2 6 4 stellata 4 7 0 Duvernaysphaera aranaides 1 4 0 , 1 4 6 radiata 1 4 2 , 1 4 7 Dyocibicides biserialis 4 5 2
deflandrei
Earlandia 5 3 , 5 5 , 56 consternatio 57 Echinitosporites cf. iliacoides 3 2 4 2 7 , 3 2 9 , 3 3 6 , 338
sp. A . 3 2 6 , 3 3 8 Echinocythereis 4 7 3 isabenana 4 7 3 jacksonensis 4 7 2 , 487 reticulatissum 4 7 3 , 4 8 0 , 4 8 7 scabra 4 8 9 scabropapulosa 4 7 4 Ektyphocythere 2 3 4 - 3 5 , 2 4 0 El 1ipsod imorphina 3 9 9 Ellipsoglandulina 2 1 4 Ellipsoidictyum cinctum 3 2 4 Ellipsoxiphus sp. A . 3 1 3 , 3 1 5 , 318 sp. B. 3 1 3 , 3 1 5 , 3 1 8 Elliptocyprites longula 26 parallela 2 5 Ellisonia 1 2 3 - 2 4 sp. 114 Elofsonella amberii 4 8 2 Elphidium 3 9 5 , 4 0 3 , 4 4 5 advenum depressulum 4 5 2 articulatum 4 5 4 cf. crispum 4 5 0 discoidale 4 5 3 - 5 4 excavatum 4 5 4 - 5 4 galvestonense 4 5 2 - 5 3 gunteri 4 5 2 - 5 3 incertum 4 1 0 , 4 5 2 margaritaceoum 4 5 4 poeyanum 4 4 3 , 4 4 5 sagrum 452 selseyense 4 5 3 Endoscrinium eisenacki 3 2 4 , 3 3 4 luridum 3 2 5 , 3 3 3 - 3 4 Endosporites 1 8 9 jurassicus 3 2 6 , 335 Entactinia 1 7 0 - 7 1 , 1 7 4 , 1 7 6 - 7 7 Entactinosphaera 1 7 1 , 1 7 4 , 1 7 6 , 1 7 7
586
Entomidella 20 marrii 22 Entomozoe marstoniana 33 Enzonalasporites vigens 1 9 3 , 196 Eoasianites 1 7 6 , 203 Eocytheropteron 256-57 aff. thiliensis 4 7 1 - 7 2 s tchepinskyi 253 trinitiensis 256 tumidum 250 wetherelli 4 7 7 , 487 Eographiodactylus eos 2 5 - 2 6 Eoleperditia 4 'Eoleperditia' bivia 4 Eoplacognathus 9 4 foliaceus 9 1 , 99 lindstroemi 9 1 , 99 reclinatus 9 1 , 99 robustus 9 1 , 99 suecicus 9 1 , 99 variabilis 9 1 , 99 Eopoikilofusa striatifera stericula 146
Eoschulerid ea 234 Epelidosphaeridia spinosa 3 4 4 , 352 Ephedra 3 5 5 Ephedripites 3 5 5 , 3 6 3 multicostatus 3 4 4 - 4 5 , 3 5 5 , 3 5 7 , 3 6 0 - 6 1 , 363-64
virginiaensis 3 4 4 Epigondolella 1 2 6 - 2 7 abneptis 1 2 5 bidentata 1 2 5 multidentata 1 2 5 mungoensis 114, 1 2 5 - 2 6 Epiplosphaera reticulospinosa 3 2 5 , 334
Epistomina lacunosa 217 Ep istom inel1a 3 9 5 exiqua 4 0 2 Eponides moremani 2 1 0 , 2 1 5 procera 4 5 5 repandus 4 4 5 Eponidopsis eshira 4 4 8 , 4 5 1 Eriella 67 Erismodus 92 Estastra 1 3 9 - 4 0 Euchitonia elegans 538 Eucomiidites minor 357 Euconochitina 1 5 1 , 1 5 5 Eucraterellina sp. cf. E. oblonga 15
Eucytheroides 256 Eukloedenella umbonata 3 2 , 3 3 , 35 Eupoikilofusa striatifera 1 4 0 Euprimitia gamachei 1 1 , 1 3 labiosa 11, 1 3 Euprimites 26 bursellus 25-27 effusus 2 4 , 26 locknensis 2 5 - 2 7 suecicus 2 5 , 27 Eurychilina 5 dorsotuberculata 2 2 , 2 4 strasburgensis 2 5 , 27 subradiata 11, 1 3 Eurycythere 2 2 1 , 2 2 7 , 2 2 9 , 234 Euryitycythere 252 Eustephanella ? jupiterensis 1 2 , 14 Eusyringium langena lagena 5 2 9 Evisceratocythere 5 0 9 Evittia sommeri 141, 1 4 6 Excentropyloma cenomana 3 1 1 - 1 2 Exesipollenites tumulus 3 2 6 , 357 Exilisphaeridium simplex 1 4 2 , 1 4 7 Exopthalamocythere 2 5 1 Fabanella 2 4 3 , 2 5 1 ansata 2 5 0 boloniensis 250 Flirliella 5 1 Falcisporites 1 8 9 Falites 2 0 Fasciolites 3 9 6 - 9 7 Flexus gutzwilleri 4 6 8 , 4 7 6 , 4 8 1 Florinites 1 8 9 Foramenella 5 phippsi 11, 1 3 Formaniella 1 7 5 cibdelosphaera 1 7 4 Fossocytheridea 2 5 6 - 5 7 Framella 7 7 Frostiella 8 groenvalliana 3 4 - 3 5 Fuhrbergiella 2 3 5 , 238 Furnishina 8 8 Furnishius 1 1 5 , 1 2 4 - 2 5 triserratus 1 1 5 Fursenkoina croneisi 215 Fusulina 7 3 Fusulinella 7 3
1 2 , Gabonella gigantea
424
Galliaecytheridea 2 3 5 , 2 3 8 , 2 5 1 Eucyrtidium calvertense 5 3 2 , 5 3 7 , 5 3 9 teres 2 5 0 , 254 Eucyrtis tenuis 312 Garniella concentrica 1 2 , 1 5 Eucytherura 2 4 0 , 2 5 1 - 5 3 , 2 6 2 , 2 6 4 Gastroammina 5 1 decorata 502 Gaudryina 2 0 5 , 2 1 2 , 2 1 5 , 216 nuda 253 ellisorae 209
587 Gaudryina l a e v i g a t a 213 G a v e l i n e l l a 2 0 5 , 216-19 ammonoides 218 b a l t i c a 2 1 0 , 215 b e c c a r i f o r m i s 396 beninens i s 448 c f . barremiana 210 d a k o t e n s i s 2 1 0 , 215-16 d a n i c a 3 9 5 , 399 h e n b e s t i 211-12 minima 215-16 m o n t e r e l e n s i s 215-17 n e e l y i 395 n a c a t o c h e n s i s 212 plummerae 210, 215-17 G a v e l i n o p s i s 219 cenomanica 210, 2 1 5 , 218 G e i n i t z i n a 51 G e i s i n a 7 5 , 77 Ginkgocycadophytus n i t i d u s 357 G l a d i o g o n d o l e l l a t e t h y d i s 115 Globanomalina a f f . p s e u d o i o t a 419-20 G l o b i g e r a p s i s 425 i n d e x 4 2 8 , 430 s e m i i n v o l u t a 4 1 4 , 4 1 9 , 558 ' G l o b i g e r i n a ' 424 G l o b i g e r i n a 415 a m p l i a p e r t u r a 4 1 5 , 420-21 a n g u l i o f f i c i n a l i s 436 angulisuturales 421 aquiensis 4 2 0 , 430 b u l l o i d e s 4 5 1 - 5 2 , 4 5 5 , 459 c f . t r i a n g u l a r i s 443 c i p e r o e n s i s 4 1 5 , 4 2 1 , 436 c i p e r o e n s i s c i p e r o e n s i s 420-21,447 c r e t a c e a 443 d e c o r a p e r t a 465 d i g i t a t a 459 d i s s i m i l i s 451 d r u r y i 465 eocaena 420 eogubina 414-15 e u a p e r t u r a 447 g o r t a n i i 436 h e l v e t i c o j u r a s s i c a 209 hexagona 459 n e p e n t h e s 4 4 6 , 465 o f f i c i a n a l i s 4 2 0 - 2 1 , 436 pachyderrna 4 5 2 , 459 quinqueloba 459 r u b e s c e n s 459 s o l d a d o e n s i s 425 t r i l o b u s c f . i r r e g u l a r i s 451 t r i l o c u l i n o i d e s 420 t r i p a r t i t a 4 4 5 , 458 woodi woodi 447 Globigerinatella insueta 451
G l o b i g e r i n e l l a 209 Globigerinelloides eaglefordensis
209 u l t r a m i c u s 209 G l o b i g e r i n i t a a e q u i l a t e r a l i s 459 d i s s i m i l i s 3 9 4 , 458 unicava 3 9 4 , 451 G l o b i g e r i n o i d e s b i s p h a e r i c u s 446 c o n g l o b a t u s 459 f i s t u l o s u s 456 glomerosus 446 h i g g i n s i 419 r u b e r 455 s a c c u l i f e r 455 t e n e l l u s 459 t r a n s i t o r i u s 446 t r i l o b u s 4 5 3 , 455 G l o b i v u l v u l i n a 5 3 , 55-56 Globoconusa d a u b j e r g e n s i s 4 1 4 , 4 1 6 ,
435 G l o b o l e b e r i s 264 Globoquadrina d e h i s c e n s 455-56 G l o b o r o t a l i a 415 a c o s t a e n s i s 4 0 3 , 4 1 2 , 4 3 4 , 446 a c u t a 415 aequa 425 a f f . w i l c o x 419 a l t i s p i r a 415 a n g u l a t a 415 a r a g o n e n s i s 415 c e n t r a l i s 4 2 8 , 430 c e r r o a z u l e n s i s 4 1 5 , 4 1 9 , 557 c f . c r a s s a t a 443 compressa 4 1 4 , 4 1 6 - 1 7 , 4 2 0 , 435
443 conomiozea 465 c r a s s a f o r m i s 400 c r a s s u l a 465 d u t e r t r e i 415 f o h s i 4 1 5 , 4 4 6 , 4 4 8 , 4 5 1 , 458 f o h s i b a r i s a n e n s i s 450 f o h s i f o h s i 450 f o h s i lobata 450 f o h s i p e r i p h e r o a c u t a 4 4 4 - 4 5 , 458 f o h s i p e r i p h e r o r o n d a 445 f o h s i r o b u s t a 458 formosa 426 formosa formosa 4 1 5 , 428 h i r s u t a 459 i n f l a t a 4 0 1 , 4 5 5 , 459 k u g l e r i 4 1 5 , 4 2 1 , 458 m a r g a r i t a e 455 mayeri 4 1 5 , 4 4 4 - 4 6 , 458 membranacea 420 m e n a r d i i 4 1 5 , 459 m e n a r d i i archaeomenardii 446 m e n a r d i i m e n a r d i i 459
588 G l o b o r o t a l i a m e n a r d i i miocenica 450 m e n a r d i i tumida 459 merotumida 458 moizea 400 nympha 465 o b e s a 446 opima 436 opima opima 41.5, 419, 4 2 0 , 447 p a c h y d e m a forma s i n i s t r o r s a 447 palmerae 419 p e r c l a r a 414 p s e u d o b u l l o i d e s 414-17, 435 pseudomenardii 41.5-16, 4 2 8 , 4'30 pseudo t o p i l e n s i s 425 p u n c t i c u l a t a 400 p u n c t u l a t a 459 p u s i l l a p u s i l l a 415-16 q u e t r a 419 r e x 415 r u b e r 415 s c i t u l a 459 t o s a e n s i s 459 t r i n i d ad ens i s 4 1 5 t r u n c a t u l i n o i d e s 41.5, 459 tumida 451 ( T u r b o r o t a l i a ) c o l l c c t e a 428 u n c i n a t a 415 ve1ascoensi.s 41.5-1.6, 428, 436 w i l c o x e n s i s 419 G l o b o r o t a l o i d e s s u t e r i 426 Globotruncana 205 c a l c a r a t a 208, 391 c o n c a v a t a 310, 316 e l e v a t a 208, 316, 391 f o r n i c a t a 208, 316 g a n s s e r i 208, 316, 391 m a r g i n a t a 294 spp. 294 s t u a r t i f o r m i s 296 G l o b u l i n a 205 l a c r i m a 212 Gloecocapsamorpha p r i s c n 139 Glomospira 5 2 , 54-55, 216-3.8 g o r d i a l i s 213, 216 G l o m o s p i r e l l a 2 3 , 25 Glossomorphites s p . 24 t e n u i l i m b a t a 21, 24 G l o s s o p s i s d e p r e s s o l i m b a t a 2 2 , 24 Glymmatobolbina ? s p i n o s a 11, 1 3 Glyphostomella 5 3 , 55-56 G l y p t o b a i r d i a c o r o n a t a 504 G l y p t o c y t h e r e 234-35, 237 G l y p t o g r a p t u s p e r s c u l p t u s 100 t e r e t i u s c u l u s 99 G l y p t o p l e u r a e l a p a 71 Gnathodus 1 2 0 , 122-23, 127 a n t e t e x a n u s 121
Gnathodus b a s s l e r i b a s s l e r i 123 b a s s l e r i symmetricus 1 2 3 bilineatus 121 g i r t y i c o l l i n s o n i 121 mononodosus 1 2 1 p u a c t a t u s 110 semiglaber 1 2 1 texanus 1 2 1 G o l o c o c y t h e r e 252 Gonyaulacysta a l d o r f e n s i s 324, 329 ambigua 325 a c u l e a t a 325 c a s s i d a t a 351 c l a d o p h o r a 324-27, 329, 334 z h r e n g e r g i 325 e x i l i c r i s t a t a 344, 352 f i l a p i c a t a 324-27, 330-31 g r a n u l a t a 325, 333 g r a n u l i g e r a 325, 333-34 h e l i c o i d e a 344, 348 j u r a s s i c a 324-27, 329, 333-34 j u r a s s i c a s s p . l o n g i c o r n i s 325 G o t l a n d o c h i t i n a 163 Guembelina g l o b u l o s a 443 G u t t u l i n a 205, 2 1 2 Cypsina 395 Gyroidina 395 G y r o i d i n o i d e s 205, 216, 218 a f f . n i t i d u s 217 d e p r e s s u s 212-14 g i r a r d a n u s 213, 398-99 n i t i d u s 213-14 octacarneratus 217 u m b j l i c a t a 214 H a b r o c y t h e r e 254, 257 Hadrognathus s t a u r o g n a t h o i d e s 97 Hagiastrin c f . Staurolonchidium tuberosum 313, 315 Halesium spxangulum 312 Halkyandia 395 IIdmarodus 94 H a m i a p o l l e n i t e s 189 Xammatocythere h e b e r t i a n a 4 6 8 , 4 7 7 , 4 8 1 , 489 Hantkenina 425, 432 a l a b a m e n s i s 4 2 8 , 431) a r a g o n e n s i s 429 p r i m i t i v a 427-28, 430 H a p l e n t a c t i n i a 1 7 2 , 1 7 4 , 176 a r r h i n a 183 H a p l o c y t h e r i d e a 256-57, 259, 263, 471, 481, 497 amygdaloides b r e v i s 265 d e b i l i s 468, 477, 481 f a b a f o r m i s 265 f a b a f o r m i s m u l t i l i r a 265 g l o b o s a 260
589
Haplocytheridea grangerensis 263 Hermanites setocpnsis 502 kummi 252 sp. 502 macropora 265 sp. aff. H. haidingeri 506 montgomeryensis 4 7 2 , 4 7 7 , 487 tschoppi 504 moodyi 472 ulrichi 502 parvasulcata 259 Hertzina 88 plummeri 2 6 0 , 2 6 3 , 265 Hesperidella 26 ulrichi 265 esthonica 27 Haplophragmoides 5 3 , 5 5 - 5 6 , 2 1 5 , 218 spp. 25 Hastigerinella 427 Hesslandona 20 Hazelina 469 Heterocypris 279 aranea 473 sp. 280 cf. aranea 479 Heterolepa mexicana 398 indigena 477 Heterostegina 397 sangalkamensis 502 Hexagonifera chlamydata 3 4 4 , 352 sp. aff. H. sangalkamensis 502 cylindrica 344 Healdia 7 5 jurassica 3 2 5 , 3 3 2 , 334 Healdiopsis 7 1 Hexastylus i 7 3 Hechtina 219 Hibbardia 65-66 Hedbergella 2 0 5 , 209 Hindeodella 9 4 , 96-97 globigerinelloides 209 crassa 9 7 , 100 hauterivica 209 crispa 9 7 , 100 infracretacea 209 sagitta 9 7 , 100 trochoidea 206 steinhornensis eosteinhornenHelotholus vema 537 9 7 , 100 sis Hemicyprideis cacigalensis 504 Hipponicharion 20 cubensis chicoyensis 504 Histiodella 92 cubensis cubensis 504 Hlubocepina 65 curta 504 Hoeglundina charlottae 2 1 0 , 215-17 helvetica 491 supracretacea 2 1 2 , 214 karlana 504 Hoioeciscus 174 larosaensis 504 Ho 1I. inella 7 5 - 77 montosa 4 7 6 - 7 7 , 4 9 1 Holocryptocanium barbui 311-12 subovata 504 nanum 311-12 Hemicythere deformis 4 7 8 , 481 tuberculatum 3 1 1 - 1 3 , 315 deformis minor 489 Huantraiconella prina 5 0 3 , 509 gordoni 504 Hutsonia 2 3 8 , 2 4 3 , 2 5 0 , 257 Hemicytherura howei 2 5 1 , 2 5 4 , 264 capelensis 250 Hemigordius 5 3 , 55 Hyperammina 5 2 , 5 4 - 5 5 , 218 Hemisphaerammina 5 1 - 5 2 , 54-55 casteri 57 geometrica 57 compacta 57 osgoodensis 57 constricta 57 Hemsiella 34 gracilenta 57 aff. maccoyiana 34 kentuckyensis 5 7 , 59 anterovelata 33 nitida 57 maccoyiana 3 3 , 35 rockfordensis 57 maccoyiana mclearni 1 2 , 15 sappingtonensis 57 maccoyiana sulcata 1 2 , 15 Hystrichodinium pulchrum 3 2 5 , 3 4 4 , Henryhowella 4 7 9 , 497-8 349 cf. evax 498 voigtii 3 4 4 , 349 evax 4 7 7 , 489 Hystrichogonyaulax cornigera 324 ex. gr. asperrima 4 9 8 , 505 nealei 324 Herend eenia p isciformis 3 2 5 Hystrichokolpoma feros 3 4 4 , 351 Hermanites 4 7 3 , 479 Hystrichosphaeridium arundum 3 4 4 , 352 hornibrooki 504 cooksonii 3 4 4 , 352 hourcqi 506 Hystrichosphaeropsis ovum 3 4 4 , 353 huantraicoensis 503 hutchisoni 504
590
Icriodella 9 2 - 9 4 , 96 deflecta 9 7 , 100 discreta 9 7 , 100 inconstans 9 7 , 100 irregularis 9 7 , 100 Icriodina 96 Icriodus 1 1 5 - 1 6 , 126-27 bilatericrescens 1 1 6 - 1 7 corniger 1 1 6 - 1 7 huddlei 1 1 6 - 1 7 huddlei curvicauda 1 1 7 huddlei huddlei 117 latialatus 9 7 , 100 latericrescens 1 1 5 latericrescens n. subsp. 1 1 7 obliquimarginatus 1 1 7 pesavis 1 1 5 - 1 7 post-woschmidti 115-17 spp. 117 woschmidti 1 1 5 , 1 1 7 Idiocythere 259 bartoniana 477 definita 262 replicata 262 Idiognathodus 1 2 0 , 1 2 2 - 2 3 , 127 acutus 1 1 5 ellisoni 1 2 3 humerus 1 2 3 n. sp. A . 1 2 3 noduliferus 1 2 3 Idiognathoides convexus 1 2 3 Illinites 1 8 9 Ilyocypris compressa 290 luzubiensis 290 triebeli 282 Imbatodinium antennatum 325 kondratjevi 3 2 5 , 3 3 4 Inhotepia similis 262 Indiana 20 Islandiella norcrossi 4 2 3 spp. 4 1 0 Isobuntonia 509 Isochilina arctica 2 4 cristata 2 0 , 2 2 , 2 4 gregaria 2 0 , 2 2 , 2 4 seelyi 2 0 , 2 2 , 2 4 Isocythereis 2 5 4 fissicostatus 2 5 4 fortinodis 2 5 4 Isohabrocythere 509 Isthomolithus recurvus 557 Janusella biceratina 1 2 , 15 Jenningsina 6 5 Jonesites semilunatus 11, 1 3 Jordanites 7 5 - 7 6 Jugosocythereis sp. 502
Jugosocythereis grimsdalei 502 Juviella 3 4 Kalyptea diceras 324 Karsteneis 2 5 9 , 262 Kegelites 7 1 Kellettina 7 5 Kentrodictyocythere 251-52 Kerionamina 5 1 Kikliocythere 2 6 2 , 2 6 4 Kikliopterygion 262 Kingmaina 2 6 4 cf. opima 4 7 1 Kinkelinella 2 3 4 - 3 5 , 240 Kirkbya 7 2 Kirkbyella (Berdanella) obliqua
12,
14
Kladognathus 1 2 0 - 2 1 mehli 1 2 1 primus 1 2 1 Klieana 235 Kloedenella scapha 3 2 - 3 3 , 35 sp. 12, 1 5 Kloedenia 8 , 32 crassipunctata 3 4 - 3 5 'Kloedenia, f a l s e ' 7 wilkensiana 1 2 , 15 Kloedeniopsis 7 retifera 1 2 , 1 5 sp. 1 2 , 1 5 Klukisporites pseudoreticulatus 3 2 6 , 3 3 0 , 335
Knoxisporites 189 Kockelella 9 7 patula 97 Kozlowskiella 6 3 , 66 Kraeuselisporites reissingeri 3 2 6 , 3 2 9 , 3 3 6 , 338
Krausella 2 2 4 rawsoni 11, 1 3 variata 2 5 , 27 Krithe 2 5 7 , 2 6 3 - 6 4 , 4 7 3 , 4 7 9 bartonensis 477 cancuenensis 505 crassicaudata 505 cubensis 505 cushmani 252 dolichodeira 505-06 guatemalensis 5 0 5 hiwanneensis 505 lambi 505 langhiana 506 londinensis 4 7 3 montensis 4 6 8 , 4 7 0 , 4 8 0 morkhoveni 505 prolixa 505 reversa 505
591 K r i t h e rocana 503 s a u n d e r s i 505 s p . a f f . K. p e r a t t i c a 502 t r i n i d a d e n s i s 505 Kyphopyxa c h r i s t n e r i 212 L a b i i s p o r i t e s g r a n u l a t u s 196 L a c c o c h i l i n a b u l b a t a 2 4 - 2 5 , 27 d o r s o p l i c a t a 2 2 , 24 p a u c i g r a n o s a 25-27 L a e v i g a t o s p o r i t e s 189 L a f f e t e i n a 395 Lagena 213 s u l c a t a 217 L a g e n a m i n a 5 2 , 54-55 Lamprocyclas h e t e r o p o r o s 5 3 2 , 5 3 7 ,
539 L a - p r o c y r t i s h a y s i 529 n e o h e t e r o p o r o s 529 L a n t e r n a s p o r t u l a 325 Lapidopiscum 175-76 Leguminocythereis 4 8 2 , 495 b o p a e n s i s 502 b u l l a t a 474 d e l i r a t a 4 7 7 , 487 l a g a g h i r o b o e n s i s 502 n. s p . 474 o e r t l i i 487 s e n e g a l e n s i s 502 s p . a f f . L. l a g a g h i r o b o e n s i s 502 s t r a i t o p u n c t a t a 4 7 4 , 487 Leiocyamus s p . 1 2 , 15 L e i o f u s a 141-42 b e r n e s g a 141 b r a z i l e n s i s 1 4 1 , 146 j u r a s s i c a 147 m u l l e r i 141 L e n t i c u l i n a 214 ex. g r . m u e n s t e r i 210 g a u l t i n a 2 1 0 , 215 klagshamenensis 435 o u a c h e n s i s 210 o u a c h e n s i s m u l t i c e l l a 210 p o l o n i c a 211 r o t u l a t a 447 s a x o c r e t a c e a 2 1 0 , 218 L e o n i e l l a 141 L e p e r d i t i a c y l i n d r i c a 3 1 , 33 s c a l a r i s 34 s c a l a r i s praecedens 35 L e p i d o c y c l i n a f a v o s a 398 L e p t o c h i r o g n a t h u s 92 Lep tod inium "arcuatum" 325 egemenii 3 2 5 , 3 3 3 , 334 r e g a l e 3 2 4 , 3 3 0 , 332 s u b t i l e 3 2 4 , 332 s u b t i l e s s p . p e c t i n i g e r u m 325
L e p t o l e p i d i t e s e p a c r o r n a t u s 357 major 326 p s a r o s u s 326 Levisulculus 5 m i c h i g a n e n s i s 11, 13 Ligonodina 97 L i l i a c i d i t e s d i v i d u u s 357-58 p e r o r e t i c u l a t u s 357-58 t r i c h o t o m o s c u l c a t u s 357-58 Limbergina 2 5 8 , 2 6 2 , 264 b i l a m e l l o s a b i l a m e l l o s a 470 l o n g i p o r a c e a 470 o r n a t o i d e l l a f i s s u r a t a 4 7 0 , 480 L i m i t i s p o r i t e s 189 Limnocythere 2 3 5 , 251 L i n g u l i n a 2 17 n o d o s a r i a 2 1 0 , 218 L i t h e l i u s minor 538 "Lithocampe" 1 7 6 , L i t h o c y c l i a a r i s t o t e l i s 529 L i t h o d i n i a j u r a s s i c a 3 2 4 , 332 v a l e n s i i 324 L i t h o m e l i s s a h e r o s 315 h o p l i t e s 3 1 3 - 1 5 , 318 p e t i l l a 3 1 3 , 315 Lithostrobus punctulatus 313-15,
318 s p . A . 3 1 3 , 3 1 5 , 318 L i t h r aph i d i t e s quad ra t u s 3 16 L i t o s p h a e r i d i u m siphoniphorum
344, 352 L i t u d a subgoodlandensis 2 1 0 , 218 L i t u o t u b a 5 2 , 5 4 - 5 5 , 214 semiplana 57 Llandoveryznathus 96 L o c k h a r t i a 395 Londinia 8 a f f . a r i s a i g e n s i s 34 a r i s a i g e n s i s 1 2 , 1 5 , 34 k i e s o w i 3 4 , 36 s p . 34 L o p h o c t e n e l l a spp. 34 Lophocythere 235 Lophokloedenia 32 m a n l i e n s i s 3 4 , 36 Lophophaena p o l y c y r t i s 3 1 3 - 1 5 , 318 L o p h o t r i l e t e s 189 Loxoconcha 263 a n t i l l e a 504 c r e t a c e a 265 l a g o s e n s i s 502 l i e n e n k l a u s i 504 l i n e a r i s l i n e a r i s 481 p r a v a 479 s i m i l i s 503 s p . a f f . L. a u s t r a l i s 506 t u r b i d g 506
592
Lox~~dus91 Loxostomoides applinae 435 Loxostomum limbatum 452 Lucina inornata 476 Ludibundus 27 Luehndea spinosa 3 2 4 , 330 Lundbladispora 189 Lunucamina 5 1 , 5 3 , 55-56 Lychnocanium grande 5 3 2 , 5 3 7 , 539 Lychnocanoma elongata 529 Lycospora 189 Lyramula 546 Macrocypris 254 Macrodentina 2 3 5 - 3 6 , 238 aspera 234 Macronotella cf. M. praelonga 30 spp. 3 0 , 31 Macrypsilon aff. salterianum 34 salterianum 1 2 , 1 5 , 3 3 , 36 Majungaella 2 3 4 , 2 3 8 , 240 kimeridgiana 238 mundula 238 oxfordiana 238 perforata 238 praeperforata 238 Manocodinium semitabulatum 2 3 4 - 2 7 , 3 3 0 , 3 3 2 , 336 Mandelstamia 251 sexti 2 5 0 , 254 Mand j ina excavata 427 Mantelliana purbeckensis 250 Maranhites mosesi 147 sp. 142 Marginalia minuta 211 Marginotruncana 205-06 concavata 208 helvetica 208 renzi 208 signali 208 Marginulina siliquina 2 1 0 , 215 Marsipella 5 2 , 55 Marssonella doneziana 211 Marsupipollenites 189 Martinotiella communis 410 Mas iukcy t here 498 Mastigobolbina lata 3 1 , 3 3 , 36 typus 3 1 , 3 3 , 36 Matronella 254 matronae 257 Ma turod inium 3 30 inornatus 324 Mauritsina 2 5 8 - 5 9 , 264 sp. 264 Megastomella africa 449 compressa 446 Mehesella 507 biafrensis 502
Meiourogonyaulax deflandrei 324 stoveri 3 4 4 , 350-51 Melonis soldanii 402 Mendicodinium reticulatum 3 2 4 , 330 Mesocena 545-46 Mesocythere 5 0 7 , 509 Mesomphalus magnificus 1 2 , 15 Messinella guanajayensis 505 jamaicensis 505 Mestognathus 120 beckmanni 1 1 5 , 121 Metacypris 2 5 8 , 290 consobrina 287 sp. 2 8 7 , 402 Metacytheropteron 2 5 3 , 258 paganum 293 Metamorphina 5 1 Micrhystridium fragile 141 Microcheilinella 7 1 variolaris 33 Microdinium veligerum 3 3 4 , 353 Micropneumatocythere 234 Microzarkodina 93 psrva 9 1 , 99 Micrystridium 1 3 9 , 1 4 2 , 360 stellatum 147 Micula mura 316 Miliolinella subrotunda 4 5 4 Minutosaccus potoniei 197 Miogypsina 457 complanata 449 cushmani 398 gunteri 398 (Miogypsinoides) bantamensis 449 (Miolepidocyclina) burdigalensis 449 negrii 449 nigeriana 449 Miogyps ino ides cornplanatus 398 Mixoneura neuropteroides 193 Monoceratella decorata 11, 13 teres 2 3 , 2 5 , 27 Monoceratina 7 1 , 79 Monogenerina 51 Monograptus angustidens 100 bouceki 100 convolutus 100 crispus 100 cyphus 100 gregarius 100 griestoniensis 100 perneri 100 riccastonensis 100 sedgwicki 100 turriculatus 100 Monosaccites 189 Monosulcites glottus 3 3 4 , 357 Mooreinella 5 3 , 55-56
593
Moorites 7 5 Neogloboquadrina pachydema 401 Morrocanium simplex 1 3 8 Neogondolella 1 2 3 - 2 4 , 1 2 6 - 2 7 Mosaeleberis 2 6 2 , 2 6 4 bisselli 1 2 3 canaliculata 4 7 0 constricta 1 2 5 - 2 6 Muderongia simplex 3 2 5 , 349 jubata 1 2 5 Muellerina 88 (conodont) milleri 1 2 5 latimarginata 4 8 9 (ostracode) mombergensis 1 2 5 - 2 6 Multioistodus 92 orientalis 1 2 3 Multiplicisphaeridiumgranulatispinosum regale 1 2 5 139 rosenkrantzi 1 2 3 hoffrnanensis 1 3 8 , 1 4 6 serrata 1 1 5 moroquense 1 3 8 , 1 4 6 serrata postserrata 1 2 3 rayii 1 4 6 serrata serrata 1 2 3 spp. 1 3 8 sp. 1 1 5 verrucarum 1 4 2 , .47 Neomonoceratina 4 9 5 - 9 6 , 507 Munseyella 4 9 5 , 507-08 helvetica 4 8 1 bermudezi 5 0 4 mediterranea 506 bollii 5 0 4 Neospathodus 1 2 3 - 2 4 , 1 2 6 - 2 7 huantraicoensis 5 0 3 arcucristatus 1 2 3 hyalokystis 502 conservaticus 1 2 5 laurea 5 0 3 cristagalli 1 1 4 , 1 2 5 muhlemanni 5 0 4 dieneri 1 2 5 punctata 5 0 4 kununeli 1 2 5 reticulata 5 0 4 newpassensis 1 2 5 Murrayina 2 6 4 pakistanensis 1 2 5 Mutilus 5 0 7 , 5 0 9 timorensis 1 2 5 Neostreptognathodus 1 2 3 , 1 2 7 Nanacythere 2 3 4 clinei 1 2 3 Nannoceratopsis gracilis 3 2 4 - 2 7 , 3 3 0 , pequopensis 1 2 3 336 prayi 1 2 3 Nanopsis nanella 2 2 , 2 4 sulcoplicatus 1 1 5 , 1 2 3 Naviculopsis 546 Neothlipsura sp. 1 2 , 1 5 Navifusa drosera 1 4 2 , 1 4 7 Neoveenia 507 Neobeyrichia 3 4 argentinensis 5 0 3 nutans 3 3 , 36 Neoveryhachium carminae 1 3 8 - 4 0 , regnans 33 146 ringerikensis 3 4 newcarminae 1 4 6 Neobulimina 217 Netrocytheridea 2 6 4 albertensis 2 1 5 , 217 Nigeria 2 6 3 , 2 8 3 , 293 minima 217 Nigeroloxoconcha oniscgvni 5 0 2 , spinosa 212 509 Neocaudites macertus 5 0 4 Nodibeyrichia 8, 3 4 triplistriatus 5 0 4 scissa 3 3 , 3 6 Neoconorbina orbicularis 4 5 4 tuberculata 3 4 Neocyprideis 2 6 4 , 4 8 2 Nodosinella 5 1 , 5 3 , 55-56 apostolescui 4 9 1 Nonion 395 colwellensis 4 7 7 , 4 9 1 affine 4 4 7 , 4 5 2 durocostoriensis 4 7 0 - 7 1 , 4 9 1 boueanum 4 5 0 grandinatus 4 7 1 centrosulcata 4 4 8 murciensis 2 6 4 , 4 7 1 commune 4 5 5 williamsoniana 4 7 7 , 4 9 1 granosum 4 5 5 Neocythere 2 5 4 scaphum 4 5 5 mertensi 2 5 4 zaandamae 4 1 0 vanveeni 2 5 4 , 257 Nonionella atlantica 4 4 7 , 4 5 2 - 5 3 Neodiversograptus nilssoni 100 cretacea 212 Neoflabellina reticulata 212 magnalinqua 4 3 6
594
Nonionella robusta 214 Norodiodus 9 4 Norochilina nora 11, 13 Notorotalia sp. 4 3 6 Novocypris 4 7 3 eocaenica 473 whitecliffensis 4 6 8 , 4 7 3 - 7 4 , 4 8 9 91 Novocythere 238 santacruceana 2 8 1 Nummulites 3 9 7 - 9 8 , 4 2 3 , 4 2 5 - 2 6 , 4 3 2 ,
0pimoLy:here martina 472 niarylandica 472 nanfaliana 4 7 2 Orbiculiforma renillaeformis 312 vacaensis 312 Orbitolites 396-97 Orbulina suturalis 4 4 6 , 4 5 1 universa 4 4 4 Orbul inoides beckmanni 4 1 9 Ordovicina 5 1 - 5 2 , 55 Oridorsalis eucadorensis 3 9 8 - 9 9 444 umbonatus 402 operculinoides furoni 426 Orionina 2 6 4 , 4 9 7 , 507-08 rockallensis 398 butlerae 504 Nuttallides 399 serrulata 5 0 4 crassaformis 396 vaughani 504 truempyi 3 9 6 , 398-99 Oroscena carolae 5 3 7 , 539 sp. (digitate) 537 Occisucysta sp. A . 325 Orthonotacythere 2 5 1 , 256-57 Occultocythereis bituberculata 506 mvili 287 Octonaria 6 4 sp. A. 2 8 9 Odontochitina operculata 3 4 4 - 4 5 , 3 4 8 , sp. B. 289 3 5 0 - 5 2 , 3 5 4 , 356 Orthovertella 5 3 , 5 5 Oepikella 5 Oryctoderma 5 1 - 5 2 , 5 5 labrosa 11, 1 3 Osangularia 2 0 5 , 2 1 3 , 216-17 Oepikium 5 navarroana 213 Oepikodus 9 1 plummerae 395 evae 91 Ostracode sp. 502 Oertliella 2 5 8 - 5 9 , 262 Oulodus 92-93 aculeata 4 7 9 , 485 Ovalipollis sp. 1 9 3 , 1 9 6 ducassae 4 7 9 Ovocytheridea 2 5 9 , 2 9 3 , 4 9 7 , 509 Ogmoconcha 236 brevis 259 Ogmoconchella 236 nuda 2 9 4 Ogmoopsis nodulifera 2 2 , 2 4 producta 259 Oistodus 9 2 - 9 3 pulchra 502 Oligocythereis gauthieri 237 Ovoidinium scabrosum 3 4 4 Oligosphaeridium complex 3 4 4 , 3 4 8 - 4 9 , Oxinoxis 5 2 , 5 4 - 5 5 352 ligula 27 dividuum 325 ligula A. 57 Ommartartus antepenultimus 5 2 9 ligula B. 57 hughesi 5 2 9 , 537-38 swallowi 57 penultimus 5 2 9 , 537-38 Ozarkodina 9 4 , 96-97 tetrathalamus 537 Oneotodus 90 Paijenborchella brevicosta 4 6 6 Oodium mordidum 1 3 8 , 1 4 6 eocaenica 4 7 7 , 4 8 1 , 487 Oolina melo 4 5 4 galerita 506 hexagona 4 5 4 s p . aff. P. malaiensis 506 Operculina conalifera 423 Paijenborchellina ijinensis 502 Operculinoides 4 5 0 Palaeohystrichophora infusorioides bermudezi 395 3 7 2 - 7 3 , 3 7 5 - 7 8 , 3 8 1 , 385 Operculodinium sp. 3 4 4 paucisetosa 3 7 2 - 7 3 , 3 7 5 - 7 8 , 381 Opimocythere 2 5 8 , 4 7 1 , 482 Paleomonsmirabilia 264 browni 472 genlensis 4 7 0 elonga 4 7 2 Paleoscenidium 1 6 8 , 1 7 2 gigantea 472 aff. cladophorum 1 7 4 incisa 4 7 0 cladophorum 1 7 4
595
Palaeoscenidium quadriramosum 1 7 4 Palaeotextularia 5 1 , 5 3 , 5 5 - 5 6 Palmatolepis 1 1 8 , 1 2 6 - 2 7 crepida 1 1 9 gigus 1 1 9 marginifera 1 1 5 rhomboidea 1 1 9 schindewolfi 115 triangularis 1 1 9 Paltodus 9 1 , 93 deltifer 9 1 Panderodus 9 2 - 9 3 Paraarchocyrtium 1 7 5 Parachirognathus 1 2 4 - 2 5 Paracirculina quadruplicus 1 9 6 scurrilis 1 9 6 tenebrosa 1 9 6 Paracypria longaensis liloensis 286 makawensis 2 9 1 Paracypridea brazilensis 2 7 3 , 2 7 7 - 7 9 obovata 236 obovata obovata 2 7 3 , 2 7 8 - 7 9 quadrirugosa 279 Paracyprideis graysonensis 258 Paracypris 2 5 1 , 2 5 7 , 2 6 3 - 6 4 alta 258 angusta 263 caerulea 250 communis 502 contracta 4 8 7 monmouthensis 265 siliqua 262 tenuicula 262 weatherfordensis 256 Paracytheretta maracensis 502 reticosa 4 6 8 - 7 0 , 4 8 0 , 4 8 5 Paradictyocha 546 Paraechmina spinosa 3 1 , 3 3 , 36 Paraexopthalmocythere 252 Paragnathodus 1 2 0 Paragondolella 1 2 6 - 2 7 polygnathiformis 125 Parakozlowskiella 6 6 - 6 7 Parakrithe 2 6 4 dactylomorpha 506 elongata 505 robusta 506 vermunti 505-06 Paranesidea elegantissima 5 0 4 Paranotacythere 2 3 6 , 256-57 blanda 253 damottae damottae 253 diglypta 2 5 2 , 2 5 4 fordensis 2 5 6 inversa inversa 253 inversa tuberculata 253
Paraparchites 7 1 carbonarius 7 0 Parapyxion 26 SPP. 25 subovatum 27 Pararotalia 3 9 5 , 398 Paraschuleridea 2 4 0 , 2 5 7 , 279 spp. 280 Paratextularia 2 8 0 Parenthatia 2 9 Pareodinia ceratophora 3 2 4 , 330 Pariceratina tricuspidata 262 Paroidtodus 9 3 originalis 91, 9 9 proteus 9 1 , 99 Parvisaccites radiatus 3 4 4 , 357 Parvocavatus tuberosus 325 Patagonocythere 4 9 5 - 9 6 , 5 0 7 , 509 Patinasporites densis 1 9 6 Patrognathus variabilis 1 2 1 Patellina 5 3 , 5 5 - 5 6 corrugata 4 5 2 subcretacea 218 Patulibrachium dickensoni 312 lawsoni 312 Pelekysgnathus 97 Pelosina complanata 2 1 1 , 216 Peneroplis 4 5 3 pertusus 4 5 2 Pericythere 5 0 7 , 509 Periodon 9 2 - 9 4 Peripyramis circumtexta 532 Perissocytheridea 2 5 6 - 5 7 , 4 9 5 - 9 6 , 508
alata 5 0 4 matsoni 5 0 4 Petrobrasia 2 8 4 diversicostata 279 marfinensis 2 7 3 , 278 Phacorhabdotus 2 5 9 , 2 6 2 , 2 6 4 , 497 pokornyi 259 sangalkamensis 502 simplex 5 0 3 sculptilis 505 semiplicatus 262 s p . 5c5 Philomedes donzei 2 5 4 Phoberocysta neocomica 3 4 9 , 3 5 1 , 354, '361
Pholadomya ludensis 4 7 5 - 7 6 , 4 8 7 , 4 8 9 Phormocyrtis striata striata 529 Phratocytheridea ruginosa 4 7 2 Phragmodus 9 2 Physocythere 262 Picostella perforata 1 4 6
596
Pilosisporites sp. A. 3 2 6 , 3 3 3 , 3 3 5 , 338 trichopapillosus 3 2 6 , 334 Pinnatulites procera 2 2 , 24 Pintopsis 7 , 32 tricornis 1 2 , 15 Pinuspollenites spp. 358 Pirea dubia 1 3 8 , 146 Piretia geniculata 2 4 - 2 5 , 27 Placopsilina 5 3 , 55-56 Plagionephrodes 67-68 Planileberis 262 Planomalina buxtorfi 2 0 9 , 362 Planorbulina mediterranea 452 Planularia pseudoparallela 211 Planulina 209 marialana 398 renzi 398 sp. 426 texana 2 0 9 , 214 wellerstorfi 4 0 2 , 410 Platybolbina bulbosa 3 3 , 36 cf. P. elongata 30-31 cf. P. plana 30-31 shaleri 11, 13 spp. 2 5 , 34 Platycosta hazeli 503 inornata 503 Platycythereis 2 5 3 - 5 4 , 256-57 gaultina 254 rectangularis 254 Platyvillosus 124-25 Plectodina 92 Pleurograptus linearis 9 9 Pleurostomella 399 torta 213 Pliolepidina tobleri 397 Plummerinella 51 Pneumatocythere 234 Podocyrtis ampla 529 chalara 529 goetheana 5 2 9 , 557 mitra 529 phyxis 529 sinuosa 529 Pokornyella 473 citrea 473 Poloniella 6 2 - 6 5 , 67-68 Polycope 2 5 1 , 2 5 4 , 264 Polyedrixum sp. 142 Polygnathoides 97 siluricus 9 7 , 100 Polygnathus 1 1 6 , 118, 1 2 0 , 126-27 asynmetricus 119 bischoffi 121 conununis 121 cristatus 119 dehiscens 116
Polygnathus dehiscens lenzi 117 faveolatus 117 inornatus 121 lacinatus121 pennata 115 spp. 117 styriacus 119 varcus 116-17 Polygonium gracile 146 Polyplacognathus 92 Polysphaeridium warrenii 3 4 4 , 3 4 8 , 351 Polystephanophorus paracalathus 3 2 4 , 334 sarjeantii 325 Polytaxis 5 3 , 55-56 Polyzygia 6 3 - 6 6 , 68-69 Pontocyprella 251-53 alexanderi 258 aturica 473 Pontocypris felix 250 Popofskyellum 1 6 8 , 176 Porcellispora longdonensis 1 9 3 , 196 Porcellisporites longdonensis 326 Porodiscus 176 Porticulasphaera 425 Poseidonamicus 4 7 9 , 4 9 6 , 509 Potonieisporites 188-89 Praebulimina 2 0 5 , 216 arkadelphiana 213 aspera 212 carseyae 211-12 cushmani 214 exiqua 215-16 fabilis 214 kikapooensis 213 Praeglobobulimina 395 Praeglobotruncana delrioensis 208 Praeorbulina glomerosa 4 1 5 , 4 4 5 , 458 Praepilatina 6 3 - 6 4 , 66 Praeschuleridea 2 3 4 , 238 batei 234 caudata 234 pseudokinkelinella 234 Primitia bicornis 2 5 , 27 sanctipatrici 30-31 simplex 2 5 , 27 sp. 24 Primitiella 34 brevisulcata 2 4 , 51-52 champlainensis 2 5 , 27 constricta 2 4 - 2 5 , 27 parallela 3 3 , 36 spp. 25 Primitiopsis p1anifror.s 33 spp. 34
597
Prioniodus 94 alobatus 91, 99 elegans 91, 99 gerdae 91, 99 navis 91, 99 triangularis 91, 99 variabilis 91, 99 Priscotheca siempreplicata 138 Pristograptus fecundus 100 ludensis 100 transgrediens 100 tumescens 100 ultimus 1OC Processobairdia 68-69 Proconodontus 88 Procythereis deformis 504 howei 504 laresensis 504 Procytheridea 235, 237, 2$4 martini 237, 240 reticulata 234 Procytherura 235, 240 Profusulinella 73 Progonocythere 234-35, 238 Prolixosphaeridium deirense 325 granulosum 325 Prooneotodus 88 Propontocypris 497 s p . 502, 506 Prosagitrodontus 87 Proscandodus 88 Protallinella grewingki 24, 51-52 Protelphidium tuberculatum 447 Proteoniana 51, 218 Protobuntonia 259, 507, 510 keiji 502 numidica 264 punctata 502 Protocosta 507, 509 struvae 503 Protocythere 235, 251-54, 256-57 bedoulensis 253 cancellata 253 cf. P. revili 251 croutensensis 253 derooi 253 divisa 252 frandei 252 hannoverana 252, 254 hechti 252 helvetica 252 mazenoti 251 mertensi 254 nodigera 254 oertlii 253 paquieri 251 praetriplicata 252
Protocythere pnmila 253 reicheli 252 speetonensis 254 strigosa 253 triplicata 252 villierensis 253 Protocytheretta karlana 489 P r o t o d i p l o x y p i n u s g r a c i l i s 196 Protoellipsodinium sp. 344, 353 Protognathodus 118-19 Pro tohaploxypinus 189 Protojonesia 264 Protopanderodus 92-94 Protoxiphotractus perplexus 312 Prunopyle titan 532, 537, 539 Psaligonyaulax apatela 325, 334 Psammosphaera 52, 54-55, 410 Pseudastrorhiza 52, 54-55 baccula 57 digitata 57 lanceola 57 Pseudoaulophacus lenticulatus 313, 315 pargueraensis 313, 315 Pseudobythocypris 76 Pseudobythocythere 253 Pseudoceratium pilliferum 348, 351 Pseudohastigerina eocenica 428 wilcoxensis 420 Pseudohutsonia 235, 238 Pseudoleperditia 72 poolei 72 tuberculifera 72 Pseudolunulidea imperatrizensis 141, 147 Pseudonodosaria 217 Pseudopalmula 52, 55 Pseudoparaparchites 75 Pseudopolygnathus multistriatus 121 Psilatricolporites 344-45, 355-56, 358-59, 361, 363-64 psilatus 358 sp. 344 Pterocanium praetextum praetextum 526 prismatium 529, 537-39 537 prismatium S.S. Pterocorys bicornis 538 splendens 537 zancleus 538 Pterospermopsis brazilensis 142, 147 Pterygocythere 258 saratogana 259 Pterygocythereis 258-59, 262-63 bartonensis 475 cornuta 474
598 P t e r y g o c y t h e r e i s p u s t u l o s a 4 7 7 , 487 t o k i a n a 259 Pterygognathus amorphognathoides 9 7 , 1 0 0 c e l l o n i 9 7 , 100 P t y c h o b a i r d i a s c h a u b e r g e r i 236 sp. 237 P t y c h o m i l i o l a s e p a r a n s 4 5 0 , 455 P u l l e n i a 2 0 5 , 3 9 5 , 399 b u l l o i d e s 410 c r e t a c e a 213 quinqueloba 410 s u b c a r i n a t a 447 P u l l e n i a t i n a 443 o b l i q u i l o c u l a t a 450 P u l s i p h o n i n a prima 2 1 1 - 1 2 , 435 Pulvinosphaeridium 1 3 9 Punctatisporites 179 P u n c t o b e e c h e r e l l a p u n c t a t a 1 2 , 14 Puriana 478 i n t e r r a s i l i s 506 r e t i c o s a 469 s c u l p t a 469 Pygodus 9 4 Pylentonema 1 7 3 , 175 Pyramidina s z a j n o c h a e 213 Pyrgo 4 5 4 nasuta 454 peruviana 454 r i n g e n s 454 s u b s p h a e r i c a 445 P y x i d i e l l a s p . 335 Pyxidinopsis challengerensis 344, 348, 350 Pyxion c a r i n a t u m 27 s p . 25 Quadracythere 473 a n t i l l e a 504 b i c h e n s i s 504 brachypygaia 504 diversinodosa 4 6 8 , 476-77, 481 g o l d e n e n s i s 502 l a g a g h i r o b o e n s i s 502 macropora 477 p r o d u c t a 504 s p . 502 s p a r s a 504 stadnichenkoae 502 Quadraditium f a n t a s t i c u m 1 4 0 , 146 Q u a s i b u n t o n i a sp. a f f . Q. r a d i a t o p o r a 506 Q u a s i h e m a n i t e s 251 Q u a s i l l i t e s 67 Q u i n q u i l o c u l i n a 215 lamarckiana 442 s a b e l l a 218 seminulum 4 5 3 - 5 4
Radiicypridina 73 Radimella 507 c o n f r a g o s a 504 s p . 504 w a n t l a n d i 504 Raibosammina 5 1 R a n i k o t a l i a 4 2 3 , 432 Raymoorea 251 Rayosoana q u i l i m a l e n s i s 2 8 0 - 8 1 "Reconcavona" b a t i k e 287 Rectocornuspira 5 3 , 55-56 Rectoplacera 6 9 , 7 9 Recurvoides 399 Rehacythereis 252 f r e d r i c k s b u r g e n s i s 260 senckenbergi 252 Reophax 5 2 , 5 5 - 5 6 , 2 1 4 , 217 buccina 57 c a l a t h u s 57 h e l v e t i c u s 2 1 0 , 218 lachrymosus 57 mcdonaldi 57 n o r t h v i e w e n s i s 57 sp. A. 57 R e t i c u l a t i s p o r i t e s 189 R e t i t r i c o l p i t e s 3 4 5 , 356 georgensis 344-45, 355, 358-59, 3 6 1 , 363-64 magnif i c u s 358 s p h a e r o i d e s 358 Retusotriletes 189 R e u s e l l a s p i n u l o s a 452 R e u s s i c y t h e r e r e u s s i 504 Reymentia 5 0 7 , 509 Rhabdammina 5 2 , 5 4 - 5 5 , 399 Rhabdochitina magna 1 6 2 t u r g i d a 162 u s i t a t a 162 Rhinocypris kroemmelbeini 287 Rhizammina 218 Rhodanicites t r i p a r t i t a 4 8 1 Rhodesognathus 92 Rhopalosyringium colpodes 3 1 3 , 3 1 5 sparnon 3 1 3 , 315 R i c h t e r i a ? sp. 34 R i c h t e r i n a (R.) a f f . l a t i o r 6 8 , 7 1 R i g i d e l l a e r r a t i c a 24 R i s a l t i n a 258 Robulus r o t u l a t u s 452 R o c a l e b e r i s 4 9 7 , 507 nacena 503 Rocella 546 R o t a l i a bensoni 395 t r o c h i d i f o r m i s 395 P s t a l i p o r a 205-06 appenninica 209 a p p e n n i n i c a ap ennin i c a 3 6 2
599 Schuleridea thoerenensis 252 travisensis 2 5 9 , 263 tumescens 258 virginis 253 washitaensis 258 Scofieldia bilateralis 11, 1 3 Scolopodus 9 0 , 92 Scriniodinium attadalense 3 4 9 campanula 3 4 4 , 349 crystallinum 3 2 4 , 3 3 2 , 342 dictyotum 3 2 5 , 3 3 3 , 3 4 4 Scyphiodus 92 Semitextularia 5 2 , 55-56 spp. 57 Sergipella transatlantica 274 Sethocapsa trachyostraca 3 0 8 , 312 Shidelerella 5 1 Sigmobolbina sigmoidea 25-27 Signetopsis semicircularis 3 3 , 36 spp. 3 4 Silenites 7 1 Similis pulchra 7 5 Saccamina 5 1 - 5 2 , 5 4 - 5 5 Siphocampe erucosa 5 3 1 howei 57 S iphogenerinoides elegantus 435 Saccorhiza ramosa 396 plummerae 212 Saetograptus fritschi linearis 100 Siphonina advena 398 leintwardinensis 100 tenuicarinata 398-99 Sagrina sp. 426 S iphonochit ina pel luc ida 162 Saida 2 5 9 , 262 Siphonodella 1 2 0 , 126-27 Sakesaria 395 cooperi 1 2 1 Sansabella 7 1 crenulata 1 2 1 Saracenaria bononiensis 215 duplicata 1 2 1 duckcreekensis 2 1 0 , 215 isosticha 1 2 1 Saxocythere dividera 254 praesulcata 1 1 9 notera 254 quadruplicata 1 1 5 , 1 2 1 tricostata tricostata 253 sulcata 1 2 1 Scabriculocypris 2 5 1 Sleia equestris 3 4 , 36 Scaliognathus 120 Sorosphaera 5 2 , 54-55 Scandodus 93 cooperensis 57 Scaphignathus velifer 1 1 9 Schizamina 5 4 papilla 57 Soudanella 4 9 5 , 5 0 7 , 510 Schizocythere appendiculata 4 7 4 arca 506 edominnini 502 laciniosa 502 Schmidtognathus hermani 1 1 9 nebulosa 502 Schopfipollenites 1 8 9 triangulata 502 SchulaDacvthere 253 Schuleridea 2 3 4 , 2 3 8 , 2 4 0 , 2 4 3 , 251- Spathognathodus 1 1 5 - 1 6 , 1 1 8 , 1 2 0 2 5 2 , 2 5 6 , 2 5 8 , 2 6 3 , 288 bidentatus 117 costatus 1 1 9 bernouilensis 253 exiguus exiguus 117 caudata 237 muricatus 1 2 3 deroii 253 n. sp. C. 117 dorsoventrus 255 (Eoschuleridea) wealdensis 250 n. sp, Q. 117 plumulus 1 2 1 extranea 253 robustus 1 2 1 jonesiana 2 5 4 , 256 steinhornensis 115-17 juddi 250 perforata perforata 4 7 4 sulcatus 1 1 6 - 1 7 sulcatus (late forms) 1 1 7 praethoerenensis 252 Rotalipora cushmani 208 greenhornensis 208 Roundyella 1 6 , 7 5 - 7 6 , 7 9 simplicissima 7 6 Ruggieria 5 0 7 , 509 sp. aff. R. rotundata 5 0 6 tattumi 502 triangulata 506 Rugoglobigerina 209 Rugotruncana 2 0 5 - 0 6 , 209 subcircumnodifer 2 0 8 , 3 8 1 Rugubivesiculites reductus 358 rugosus 3 5 8 , 363 Rugulidium microrugulatum 1 3 8 rugulatum 1 3 8 , 1 4 6 scabratum 1 3 8 triangulatum 1 3 8 varirugulatum 1 3 8 Ruttenella 5 0 7 , 509 Rzehakina epigona 2 1 3 , 216
-~
600
Spathognathodus t r i d e n t a t u s 121 Sphaero i d i n a bul loi d e s 39 9 Sphaeroleberis 264 pseudoconcentrica 260 Sphaerostylus l a n c e o l a 312 Sphenolithus pseudoradians 557 Spinicythereis 259 S p i n i f e r i t e s c i n g u l a t u s 344, 351 ramosus 352 S p i n o l e b e r i s 256, 258-59, 262 S p i r i l l i n a 53, 55-56, 216-18 Spirolocammina 410 Spiroplectanunina 53, 55, 213, 215, 2 1 7 , 399 a l e x a n d e r i 210, 218 mexiaensis 213 s p e c t a b i l i s 399 S p i r o s i g m o i l e n e l l a 410 Spongas t e r 528 berminghami 527-29 c r u c i f e r u s 530 p e n t a s 443, 526-31, 537-38 t e t r a s 531 t e t r a s t e t r a s 537-38 Spongopyle o s c u l o s a 531, 538 Spongosaturnalis h u e y i 313, 315 p r e c l a r u s 313, 315 Spongotrochus g l a c i a l i s 538 S t a c h e i a 51 Stacheoides 51 S t a p l i n i s p o r i t e s caminus 326 Staurosphaera septemporata 312 Stegnamnina 5 2 , 54-55 S t e l l i n i u m c r i s t a t u m 142, 147 S t e n s i o e i n a 213-14, 216 Stephanelytron caytonense 325, 332 r e d c l i f f e n s e 325 scarburghense 325, 332 S t e u s l o f f i a a c u t a 22, 24 c o s t a t a 26 linnarsoni 26 p o l y n o d u l i f e r a 22, 24 Steusloffina 5 c o s t a t a 25, 27 l i n n a r s s o n i 25, 27 u l r i c h i 11, 13 "Stichocapsa" 176 S t i c h o c o r y s delrnontensis 529, 539 p e r e g r i n a 529, 537, 539 w o l f f i i 529 S t i c h o m i t r a asymmetra 313, 315, 318 313, 315, 318 sp. A. Stichopilidiurn t e s l a e n s e 313 Stigmosphaera 170 S t i l o s t o m e l l a 213-14, 399 a c u l e a t a 399 c u r v a t u r a 399
Stolodus 93 Stomasphaera 52, 55 S t r a v i a c r o s s a t a 252 Streptognathodus 1 2 7 e c c e n t r i c u s 115 u n i c o r n i s 121 S t r e p u l i t e s d a l h o u s i e n s i s 1 2 , 15 S t r i a t i t e s 189 S t r i a t o a b i e t i t e s 189 S t r i a t o p o l l i s paraneus 358 Striomonosaccites 189 Stylochlamidium a s t e r i s c u s 538 S t y l o s p h a e r a 173 Subbotina l i n a p e r t a 394 patagonica 394 t r i l o c u l i n o i d e s 416 S u b t i l i s p h a e r a p e r l u c i d a 344, 351, 354 pontis-mariae 344, 353, 356, 361, 369, 374-75, 377, 381, 385 S u l c a t i s p o r i t e s 189 S u l c o s t o c y t h e r e 495-96, 508-09 S v e a l u t a 20 Sweetognathus 123, 1 2 7 w h i t e i 123 Systematophora a r e o l a t a 324, 335 f a s c i c u l i g e r a 325, 344 o r b i f e r a 324, 332 schindewolfi 325, 334 t u r o n i c a 324, 334 Taeniophora i u n c t i s p i n a 325 Ta 11i n e 11a d imo rpha 2 5 -2 7 t r i d e n t 25, 29 Taphrognathus 120 v a r i a n s 121 Tasrnanites spp. a f f . T. mourai Tenua h y s t r i x 325 r i o u l t i 324 sp. 344 s p . A. 324 sp. B. 324 v i l l e r s e n s e 325 T e t h y s i a 251 Tetradella 5 a f f . q u a d r i l i r a t a 29-30 a n t i c o s t i e n s i s 11, 13 buckensis 11, 13 complicata 25, 29 s p . 24 thomasi 11, 13 u l r i c h i 11, 13 T e t r a g r a p t u s approximatus 99 Tetralithus gothicus t r i f i d u s n i t i d u s t r i f idus 316 t r i f i d u s 316
141
316
601 Tetrasacculus 7 1 Tetrataxis 5 3 , 55-56 Tetrentactinia 1 7 4 , 1 7 6 Textularia 5 3 , 5 5 - 5 6 , 2 1 4 , 217 gramen 4 5 2 - 5 3 rioensis 110, 1 1 8 ripleyensis 1 1 2 washitensis 110 Thalmannia ? fusa 506 Thalmannita 395 Thekanunina 5 1 Theocampe altamontensis 3 1 3 , 3 1 6 apicata 3 1 3 - 1 4 , 316 ascalia 3 1 3 , 316 bassilis 3 1 3 - 1 4 , 3 1 6 daseia 3 1 3 - 1 4 , 316 lispa 3 1 3 , 316 mongolfieri 529 salillum 3 1 3 , 316 Theocapsomma comys (group) 3 1 1 - 1 3 ,
Tolypammina bulbosa 57 cf. prestwichiana 4 7 9 cyclops 57 frizzelli 57 gersterensis 57 jacobschapelensis 57 laocoon 57 rotula 57 Trachycythere 2 3 4 , 262 Trachyleberidea 2 5 8 , 262 cubensis 505 goochi 479 mammidentata 505 prestwichiana 4 7 3 , 485 Trachyleberis bermudezi aff. crebripustulosa 5 1 1 bermudezi bermudezi 505
bermudezi crebripustulosa 505 bollii 505 floriensis 4 7 2 , 4 7 7 , 4 8 7 316 teren 3 1 3 - 1 4 , 316 huantraicoensis 5 0 3 Theocotyle cryptocephala cryptocephaspiniferrima 4 7 5 la 5 2 9 spinosissima 475 nigrinae 5 2 9 teiskotensis 502 Theocyrtis redondoensis 5 3 2 , 5 3 7 , weiperti 503 539 Trepeilopsis 5 3 , 55-56 tuberosa 5 2 9 , 566 glomospiroides 57 prodigalis 57 Theriosynoecum 2 3 5 , 258 recurvidens 57 varietuberatum 277-78 Treposella 6 6 Thlipsura cf. T. v-scripta 1 2 , 1 5 Tretaspis 3 1 Thlipsuropsis inaequalis 1 2 , 15 Triactinosphaera sp. 3 1 6 , 318 Tholosina 5 1 - 5 2 , 54-55 Triadispora obscura 1 9 6 elliptica 57 sp. 1 9 6 Thomasatia falcicosta 11. 1 3 . 2 4 - 2 5 . Triangulina alargada 141 29 Triceraspyris 537 Thurammina 5 2 , 54-55 Tricladiodus 92 arenacorna 57 Tricolpites 356 congesta 57 auritus 3 4 4 , 358 triradiata 57 micromunus 3 4 4 , 358 Thuramminoides 5 4 minutus 3 4 4 - 4 5 , 3 5 5 , 3 5 8 - 5 9 , Thyrsocyrtis bromia 5 2 9 , 5 5 6 - 5 7 , ,
I
564-65
finalis 5 5 7 , 564-65 tetracantha 5 5 6 , 5 6 4 - 6 5 triacantha 5 2 9 , 5 5 6 , 564-65 Timirasevia 235 Togoina 2 9 6 , 4 9 5 , 510 attitogoensis 502 australis 5 0 3 kugleri 502 kugleri soldadoensis 502 obesa 502 Tolypammina 5 2 , 5 4 - 5 5 , 218 aff. semiplana 4 7 1 botonucus 57 bransoni 57
3 6 1 , 363-64
Tricolporites sp. 3 4 4 , 359 "Tricolporopollenites" triangulus 358
Tricorni.na 6 3 - 6 6 , 6 8 - 6 9 , 7 1 (Bohemina) 7 6 (Tricornina) 7 6 Triebelina crumena 5 0 4 howei 5 0 4 Trigonopyxidia ginella 3 4 4 , 353 Trilobosporites jurassicus 3 2 6 , 3 3 2 , 335
Triplacera 62 Triporopollenites 359 Tritaxia capitosa 2 0 9 , 2 1 4 Trithyrodinium suspectum 3 4 4 - 4 5 , 347 353-54, 361
602
Trochamnina 5 3 , 5 5 - 5 6 , 217-18 mehli 57 Trochamninoides irregularis 213 Trocholina transversarii 211 Truncorotaloides collactea 420 rohri 4 1 9 Truyolsina truyolsi 7 6 Tuberitina 5 1 Tumidoleberis 264 Turritellella 5 2 , 55 Tuscaritellum 68 Tvarenella 26 carinata 29 sp. 25
Vernuil inoides 2 1 8 Verrucosisporites cf. cheyneyi 3 2 6 , 329
sp. A . 3 2 8 , 338 Veryhachium 1 4 2 , 360 lairdi 1 3 8 Vesicaspora 1 8 9 bulbiferum 1 3 9 carminae 141 exasperatum 1 4 1 irregulare 1 4 2 , 147 legrandi 141 reductum 141 trisulcum 141 Vestrogothia 20 Vestustocytheridea 4 7 1 , 482 Uhakiella coelodesma 29 fornicata anteronoda 472 spp. 25 fornicata fornicata 4 7 1 Ullerella triplicata 2 5 , 29 guitrancourtensis 470 Ulrichia (Ulrichia) bipunctata 2 5 , 2 9 lignitarum 470 (Ulrichia) verticalis 1 2 , 1 4 Ulrichodina 90 macrolaccus munseyi 472 Uroleberis sp. aff. U. ranikotiana sp. 4 7 1 502 Virgatocypris edwardwi 4 9 1 Uvigerina 3 9 5 , 3 9 9 , 446 grisyensis 4 9 1 abbreviata 436 tenuistriata 4 9 1 germanica 436 Virgiana 6 tenuistriata 4 3 6 Vitreisporites pallidus 3 2 6 , 357 Vittatina 1 8 8 - 9 0 Vaginulina debilis 2 1 0 , 215 Vulvulina 214 Valensiella ampulla 324 Walchia linearifolia 193 ovulum 3 2 4 , 332 piniformis 1 9 3 vermiculata 3 2 4 - 2 7 , 332-33 Wallodinium krutzschii 3 4 4 , 3 4 9 , Vallasporites ignacii 1 9 6 351 Valvulinella 5 3 , 55-56 Wanaea spectabilis 325 Valvulineria 2 1 6 , 218-19 "Webbinella" 5 1 allomorphinoides 212 Webbinelloidea 5 2 , 54-55 infrequens 2 1 0 , 217 lotterlie 2 1 0 , 215-17 similis 57 plummerae 214 Wiekkoella 5 4 Welleria 7 , 32 Veenia 2 5 4 , 2 5 7 , 2 5 9 , 262-64 acuticostata 502 obliqua 3 3 , 36 Welleriopsis 32 arachoides 265 jerseyensis 3 4 , 36 florentinensis 253 Wichmannella 5 0 7 , 509 harrisiana 254 meridionales 503 (Nigeria) 2 6 3 , 2 8 3 , 296 ozanana 262-63 Wolburgia 282 neocretacea 288 paratriplicata 263 paleocenica 503 reticulata 259 spoori 262 Xestoleberis 253-54 ughelli 294 Itelere 502 varriensis 294 warriensis 502 mauryae 502 sp. A. 506 Veenidea 263-64 Veliferites tenuimarginatus 1 4 1 , 146 XiPhOstYlus 1 7 3 Ventilabrella 205 Zabythocypris heterodoxa 511 Venzavella sp. 1 2 , 14 Zygobeyrichia 7 , 6 3 Verneuilina 213 Zygocosta 6
603
Zygobursa 6, 7 praecursor 1 2 , 14 Zygobolba anticostiensis 1 2 , 14, 31, 33, 36 decora 7, 12, 14, 31, 33, 36 erecta 31, 33, 36 robusta 12, 14 twenhofeli 1 2 , 14 Zygobolbina emaciata 31, 33, 36 Zygosella postica 31, 33, 36
This Page Intentionally Left Blank
