Professional Documents
Culture Documents
I. Jarvis
Sc~hool of Geological Sciences, Kingston Pol>*technic Rood, Kingston
, Pc~nrh~~n upon Thntne~,
Surre_v KTl 2EE, U.K.
G. A. Carson
Department of Earth Sciences, lJniz~ersit_v of Liewpool, Brozonlow Street. PO Box 147.
T,ieterpool. L69 3BX, U.K.
M. K. E. Cooper
Strntigraphic Services International Ltd., Chancellor Court, 20 Priestly Road, Guildford.
.Surre>,, GU2 5 YL, U.h’.
D. Home
Department of Geology, City of London Pol>stechnic, H’nlhurgh IIozrse, Bigland Street. London
El 2,VG, l.:.h-.
and A. Rosenfeld
Geological Surr,e\l of Israel, 30 ;%falkhe Yisrael Street. Jerusalem 95 501, Israel.
I. Jar\G, G. A. Carson, M. K. I-. Cooper, RI. B. Hart, P. ,X. Lear?, B. .4. ‘rocher, D. Horne
and X. Kosenfeld. Microfossil .4ssemhlages and the Cenomanian-Turonian (late Cretaceous)
Oceanic Anoxic Event. Cretaceous Research (1988) 9, 3-103. The effects of the Cenomanian-
‘I’uronian Oceanic Anoxic Event (OAE) in the Chalk Sea of NW Europe have been
investigated using published macrofossil records combined with new detailed sedimentolog-
ical. foraminiferal, ostracod, calcareous nannofossil, dinoflagellate cyst and stable-isotope
data from Dover, England. The ranges of individual fossil species are displayed against
lithostratigraphic logs, and their relation to the Cenomanian-Turonian houndarv (defined
using macrofaunal data) is discussed. .4 positive carbon stable-isotope excursion, indicating
the stratigraphic extent of the OAE, spans the stage boundary. Correlation with successions
elsewhere in NW Europe suggests that the OAE was isochronous, and that major hiostrati-
graphic marker horizons are characterised by distinctive 6°C values. All microfossil groups
display uppermost Cenomanian abundance and diversity minima which correspond closely to
the peak of the carbon stable-isotope excursion. We propose that the OAE was a phase of
increased upwelling which led to a widespread espansion and intensification of the oxygen-
minimum zone in the oceans. As a result, increasingly dysaerohic bottom waters developed
Lvithin the Chalk Sea, and were responsible for progressive disappearances in the benthonic
microfaun?,including the extinctions of many typical Cenomanian taxa. At the same time the
oxygen-mlmmum zone rose in the water column, causing the extinction of deeper water
planktonic foraminifera and then the gradual loss of intermediate-\vater groups. A temporar!
disappearance of dinoflagellate cysts and a proliferation of calcispheres were associated v ith
these events. As the OAE waned, new species eraduallv evolved to fill niches left vacant
following the extinctions of Cenomanian &a. The appearance of these new species defines the
base of the Turonian. the stage division beinn a direct conseauence
I, .,
of the OAE. It is concluded
the 0.41:s provide a major mechanism for controlling rates of evolution and extinction
throughout the Phanerozoic.
KIX \~ORDS: Cretaceous; Cenomanian; Turonian: Chalk; Oceanic Anoxic Event; Foramini-
fera; Ostracoda; Xannofossils; Dinoflagellates; Correlation: Stratigraph!; Palaeocean-
ography: Oxygen-Minium Zone: Extinction; Evolution.
Contents
1. Introduction (I.J.)
11. Effects of the OAE on the Chalk Sea biota (I. J. ef al.)
11.1 Microfossil abundance and diversity
11.2 Oxygenation levels and the biota
11.3 Expansion of the oxygen-minimum zone into the Chalk Sea
11.4 Extinction. evolution and the OAE
1. Introduction
DOVER
West
Houghton
-I
2km
..
&FOLKESTONE
Microfossil Assemblages and the Oceanic Anoxic Event 7
(Sled)
~ivw kmoa
I I
(wed)
WY3 JaM0-l
blicrofossil Assemblages and the Oceanic Anosic Event 9
Figure 3. The Cenomanian-Turonian boundary sequence on Abbots Cliff path, Dover. The bases of the
Plenus Marl Formation and Melbourn Rock Beds (Shakespeare Cliff Member, Dover Chalk) are
indicated by the lower and upper discontinuous white lines, respectively. Numbered subdivisions of
the Plenus Marl indicated by the white numerals are the bed numbers of Jeff&es (1962, 1963q h).
Locations of samples collected for microfossil and stable isotope analysis are marked by the black-
filled circles and black numerals (samples ABCS-21). The base of the Turonian. defined by the
appearance of &,rytiloides spp. inoceramid bivalves coincides with sample ABC19.
-
Microfossil .&xemblages and the Oceanic .-\nosic Event 11
2.4 Discztssiort
The Plenus Marl is differentiated from the Abbots Cliff Chalk not only by its
higher clay-mineral content, but also by containing an increased proportion
of silt-sized and coarser grained bioclasts. These observations, together with
the prominence of omission surfaces in the sequence, suggest that removal of
micritic (clay-sized) carbonate by current winnowing was probably a major
process during the deposition of the Plenus Marl. VVinnowing would not
necessarily lead to the removal of clay minerals to the same extent as clay-
grade carbonate. The former possess charged surfaces which cause them to
form silt-sized floes that behave as coarser grained aggregates rather than
individual particles. Winnowing could, therefore, concentrate a range ofsilt-
sized components, both clay minerals and bioclasts.
An alternative hypothesis is that both the proportions of clay minerals and
the coarser-grained carbonate fraction have been increased by the disso-
lution of clay-grade carbonate. Dissolution is certainly not a significant
process in the lower Plenus Marl (Beds l-3), where calcareous microfossils
(including nannofossils) are very well preserved. Above this, however, in
Beds 4-8 and in the basal metre of the Melbourn Rock Beds, microfossils
display evidence of etching. We interpret this dissolution as having been
dominantly a seafloor process (Section 11.3), but later diagenetic dissolution
processes may have had some effect on the composition of the lithofacies.
A major increase in the proportion of calcispheres is seen towards the top
of the Plenus Marl, and calcispheres remain a major component throughout
the Melbourn Rock. The increase in the proportion of calcispheres is a result
of several factors. Firstly, winnowing and dissolution have led to the removal
of clay-sized carbonate. Secondly, there is a significant decline in the
abundance of foraminifera, which form a major component of the coarse-
grained fraction of the lower beds. Finally, calcispheres must have been more
abundant at that time, since the proportion of other bioclasts (e.g. bivalve and
echinoderm debris) is also low.
There are three principal features which differentiate the Melbourn Rock
from the summit of the Plenus Marl, below. Firstly, there is a major decline
in the percentage of clay minerals. Secondly, there is the addition of micritic
cements, both within hardgrounds and isolated nodules. Thirdly, there is an
increase in the mean size of the bioclastic fraction due to the addition of
significant proportions of coarse-grained inoceramid bivalve shell debris.
Fauna1 records for the Dover successions are available from several sources,
particularly Jefferies (1962, 1963a, b) and Kennedy (1969) for the upper
Abbots Cliff Chalk, Jefferies (op. cit.) and Wright and Kennedy (1981) for the
Plenus Marl, and Jukes-Browne and Hill (1903) and Woodroof (1981) for the
basal Dover Chalk. These records are used to define the position of the
Cenomanian-Turonian boundary in the succession, and provide a biostrati-
graphic framework with which our new microfossil and stable-isotope data
can be compared.
Macrofossils are uncommon in the upper beds of the Abbots Cliff Form-
ation, although abundant inoceramid bivalve debris and fragments of thin-
shelled echinoids (Holaster sp.) occur at some levels. Fauna1 records
(Jefferies, 1962, 1963a, b; Kennedy, 1969; A. S. Gale, pers. comm., 1987)
indicate that Holaster trecensis Leymerie and the bivalves, Inoceramus pictus
J. de C. Sowerby, Pycnodonte ztesicularis (Lamarck) and Neithea quin-
quecostata (J. Sowerby) are the commonest fossils. Other records include the
bivalves, Plagiostoma globosa (J. de C. Sowerby), Plicatula inflata J. de C.
Sowerby, Chlamys beaveri (J. Sowerby), Entolium orbiculare (J. Sowerby),
the brachiopods Terebratulina (dominantly T. striata Wahlenberg), Lingula,
Concinnithyris, and the worm tube ‘Serpula’ umbonata Mantell. No am-
monites are recorded from these beds in the coastal successions, although by
comparison with inland exposures (Kennedy, 1969), the sequence is regar-
ded as being Late Cenomanian Calycoceras guerangeri Zone (Figure 2). The
Late Cenomanian age is confirmed by the occurrence of Inoceramuspictus (cf.
Troger, 1981).
Microfossil Assemblages and the Oceanic Anoxic Event 15
_
The hardground sequence at the base of the Melbourn Rock Beds yields
Inocevamus pictus, Sciponocevas hohemicum ante&s Wright and Kennedy,
Tylocidaris hirudo spines, pycnodonteine oysters and hexactinellid sponges
(Woodroof, 198 1; Wright and Kennedy, 198 1). Inoceramus pictus ranges up
to just over 1 m above the top of the Plenus Marl. Immediately above this,
inoceramids \vith the shape of MJltiloides but possessing an external orna-
ment similar to that of I. pictus occur in association with true Mytiloides spp.
(C. J. Wood, pew. comm., 19X7), particularly AZ. mytifoides (Mantell).
A4Jltiloides labiatus (Schltitheim) appears in the upper part of the Shake-
speare Cliff Member, immediately above the Melbourn Rock (Woodroof,
1981).
‘I’he Melbourn Rock at Dover is best known for its abundant echinoid
fauna (Price, 1877; Rowe, 1900; Jukes-Browne and Hill, 1903), particularly
Discoidea dixoni Forbes and C’avdiaster pygmaeus Forbes. Salenia granulosa
Forbes, Tylocidaris hirudo and rare Glzrphocyphus radiatus Hoenig also occur,
and ~‘o~~ulussubvotwzdus Forbes and 6. ‘castanea’ (Brogniart) make their first
appearances towards the summit of these beds. This echinoid fauna occurs
throughout the Shakespeare Cliff Member, but is uncommon above the
Melbourn Rock.
‘I’he calcarenitic component of the Melbourn Rock primarily consists of
comminuted inoceramid shells with subordinate echinoid debris. However,
the brachiopods Orbirhynchia cuzievi (d’orbigny), Concinnithvris sp.,
Terebratuli?la sp. and hexactinellid sponges are common throughout the
Shakespeare Cliff Member. The kival\res PIicatula sp., Plagiostoma sp. and
Lima sp. are also recorded.
Ammonites are uncommon in SE England above the Plenus Marl,
although Sciporzoceras bohemicum antevius and S. gvacile occur in the basal
part (see above). Watinorevas sp. is recorded from Merstham (Wright and
Kennedy, 1981), while Mammites twdosoides (Schliiter) and Lezoesiceras
pevamplunz (Mantell) occur in the higher beds of the Melbourn Rock at
Do\,er, and rarely in the upper part of the Shakespeare Cliff Member.
Table 1. AlphabetIcal list of foraminiferal species and their stratigraphic distribution AC, Abbots Cliff
Chalk [upper Capel-le.Ferne Member]; PM, Plenus Marl Formation; DC, Dover Chalk Formation
[Shakespeare Cliff Member]).
Species Distribution
4.2 BiostratigraphJr
4.2.1 Abbots Cliff Chalk Formation. Samples from the Abbots Cliff Chalk
(Figures 6, 7) consistently yielded abundant foraminiferal assemblages, with
over 50°, of the fauna being represented by planktonic taxa (Figure 9). The
planktonic fauna is dominated by Hedbergella spp., with rotaliporids [Rotali-
pora cushmarzi (Morrow), R. greenhornensis (Morrow)] and praeglobo-
truncanids/discarinellids each averaging only 25O, of the planktonic total.
It is important to note the appearance of the Dicarinella group, within
the uppermost 3 m of the formation (Figure 6), since this genus is norm-
ally regarded as a Turonian-Santonian indicator. The benthonic fauna
(Figures 6, 7) is of high diversity and is typical of the late Cenomanian (Zone
13 of Carter and Hart, 1977), being dominated by species of Arenobulimina
[e.g. Figure 10(a)], Gavelinella [Figures 1 O(l)-(n)], Lenticulina [Figure 10(j),
(k)], Plectina [Figure 10(g)] and Tritaxia [Figure 10(b), (c)l.
4.2.2 Plenus Marl Formation. The eight beds of the Plenus Marl (Jefferies,
1962, 1963a, 6) each contain a distinctive foraminiferal assemblage. Bed 1 is
characterised by a fauna which is very similar to that in the underlying
Abbots Cliff Chalk (Figures 6,7). The diversity of the benthonic assemblage
is also comparable, but there is an increased percentage of planktonic taxa
(Figure 9). Rotalipora cushmani [Figures 12(b), (c)] is considerably more
abundant (the genus constitutes > 50°10 o f the planktonic assemblage), and
t
cll
0”
cl
*n
0
00
0
Microfossil Assemblages and the Oceanic Anoxic Event 21
4.2.3 Dover Chalk Formation. The cemented nodular chalks of the Mel-
bourn Rock are extremely difficult to process for foraminifera, but even in
more marly intervals (where the samples are less indurated, and larger
numbers of individuals were recovered; Figure 9), the fauna is characterised
by low numbers and low di\:ersities. The planktonic fauna is generally poorly
Planktonics
Stage
Formation
90
60
70
60
i
I I iI iI
0 0
I
I I I 0 0
Lithology
o-o-. ,::~~~~~~~
:i’:g.:yi::;I: .:: : .:::;, :i:::::
g
0
,,::, : ::::i .,.,:.,_
::. ::::,,:::....>..y.,
a
-
.- -m- - --y$&l!y.-
,.a_
,‘,.
:.. ::;
e:
.,. : ” /
.-
o-o- l -.-~
o-o-o-o-o
o-o-o-o-a-o:
l - l- . - . -
0-o-a’ -----a-
o-o-*
Cenomanian Turonian
ABBOTS CLIFF CHALK DOVER CHALK
,.::fy;
‘:‘:~.,:i?J,~:,::~ .,.
D+--- o-o-o rritaxia mactedyenr
l- .---.-o-o r. pyramldata
0. ceno*an,ce
o-o-o-o
0 GyrOidlnOideS paw.3
0 n i c s
preserved in the basal beds, but above this diversities gradually increase,
caused by the’ appearances of ‘Hedbergella’ archaeocretacea (Pessagno)
[Figure 12(h)] and Dicarinella spp. [including D. imbricata (Mornod)]. Also
appearing in the MeIbourn Rock Beds are low-spired hedbergellids that
develop faint marginal keels [Helvetoglobotruncana ‘praehelvetica’ (Truj-
illo)]. There is then a complete gradation between such early forms and true
(Boreal) Heln~etoglobotrzuzcanan helvetica (Bolli). The transition may not be
synchronous with such changes in ‘l’ethyan areas, as this evolution is
probably at least in part a function of available water depth (Hart and Bailey,
1979; Hart, 1982). Marginotvuncana marginata (Reuss) [Figures 12(i), (j)]
Turonian
DOVER CHALK
r .yr:>.‘)
,;_.+ ~;,‘:;:’
0 ,
2 \’ ..J_’ . . ..
@ 8
E
l
9 . ;
-0-o
I
I
. - - - - ..- - - -
--m-v ----
-o-0
-0 0 ; o-o-.
,
I .-•-
I
XIicrofossil .%ssrmblages and the Oceanic Xnosic Event 27
Benthonics
I
I
I
I
0
I
0
i
I
I
0
I
I
I
1 0
::,
I
I I
I
3
‘.
7
I
.: I “.j I
I _,
: :
:
:..
’ I
3
1
_- I _. .- .- ; .,__
-I -
I
0 1 I
I
I
I
I
I
I
.I
I
I
I I
i 0
I I
I I I I
-
U~!LlOJfl~ ueluelUoua~
I I
,
: I
Microfossil Assemblages and the Oceanic Anosic Event 29
appears above the Melbourn Rock, in the higher beds of the Shakespeare
Cliff Member (Figure 8). The first occurrence of true H. helvetica (Figure
12e-g) was noted immediately above the top of the Shakespeare Cliff
Member, in the Warren Marls (lithostratigraphic terminology of Robinson,
1986), approximately 7 m above the top of our detailed sampling (Figure 8).
Planktonic assemblages in the upper beds of the Shakespeare Cliff Member
are characterised by increasing proportions of praeglobotruncanids and
dicarinellids, and declining percentages of hedbergellids (Figure 9). This
changing population structure is considered to be a response to increasingly
favourable, probably deeper water, conditions during the early Turonian.
The benthonic fauna is of low abundance and diversity throughout the
Shakespeare Cliff Member, although there is a gradual increase in diversity
towards the top of the unit (Figure 9). Particularly noteworthy are the
appearances of Arenobulimina preslii (Reuss) [Figure 12(l)], which occurs in
the Melbourn Rock, and Tritaxia tricarinata var. jongmansi Schijfsma
[Figure 12(k)] and LinguZogaveZin~ZZa aumalensis (Sigal) [Figure 12(m)],
which are first recorded in the higher beds of the member (Figure 8).
Figure 10. Cenomanian-Turonian foraminifera from Dover I. SEM photomicrographs of typical late
Cenomanian benthonic species. 411 of these taxa are common throughout the upper Cenomanian
except Ataxophragnium depressurn, which is a rarity that occurs in flood abundance in Plenus Marl
Bed 1. (a) Arenobuliminn advena (Cushman), ABC7. x 75; (b) Tritaxia macfadyeni Cushman, ABC7,
x 10: (c) Tritczxia pyramid&a Reuss, ABC6, x 115; (d) Dorothiagradata (Berthelin), ABCl, x 115;
(e) Marssonella trochus var. turris (d’orbigny). ABCl, x 150; (f) Eggerellina breuis Marie, ABCl.
x 115; (g) Plertina ma&e (Franke), ABCl, x 150; (h) Nodosaria sp. A, ABC4, x 55; (i) Dentalina sp.
A. ilBC8, x 55; (j) Lenticulina rotulata var. A, .4BC2 x 115; (k) Lenticulina rotulata var. B, ABC 1,
x 115; (1) Gawlinella baltica Brotzen, ABCl, x 75; (m) Gawlinella cenomanica (Brotzen), ABCl,
x 115; (n) Gavelinelln intermedia (Berthelin), ABC3. x 1 SO: (o) Atu.xophragmium depressurn (Perner).
ABc’7, x 115.
Microfossil .&emblages and the Oceanic anoxic Event 33
SpWieS IIistributron
Bairdoppilata sp. A
H. pseudoseptentrionalis Mertens, 1956
B. southerhamerlsis Weaver, 1982
B~thoceratirra herrigi \Veaver, 1982
B. umbonata (Williamson, 1817)
B. umbonatoides (I&ye. 1964)
C’urfsina senior Porkom$. 1967
(‘ythereis sp. A sense Weaver, lY82
c ‘. sp. cf. C’. lo,~~ur~~~z I’orkwny. 1063
C~\‘t/rr/d/us,,. cf. (‘. clzclt/lrl?namls \veaver, 19x2
C. . U,I,C~~1‘ULVVezi\t‘r. I OX?
C‘.sp. cf. (‘. ~ofrtrrzl‘tn van Yeen. 1932
C’. wata Koemer. 1 X41
(‘~therrlloidea bonnrmai 1Veaver. 1982
r. hindei &ye, 1964
(‘. kayei Weavrr, 1982
C’. obliquirugata (Jones and Hinde, 1890)
C‘. stricta (Jones and Hinde, 1890)
Herrigocvtherr donzei (W”eaver, 1982)
Imhotepia eugl_vpheo Weaver. 1982
Isoc~thereis elongatn \Veaver, 1982
l,:~~~x~~who~ blrtrhel1crrsi.s\VC;I\er. 1OX?
Mosarleberis sp. X
Owtliella alata Weaver, 1982
0. donzei Weaver, 1982
Pawac~thrreis subpawa (Porkomq, 1967)
Phodrucythere cuneifortnis Weaver, 1982
Pontoc~prella robusta Weaver, 1982
Psrudob~thoc_vthere colini \Veaver, 1982
Ptervgocvthere sp. cf. P. dinrinuta Weaver, 1982
/‘. S,‘. ct. I’. lutlcr-r.stata (hsquet, 1854)
I’. sp. cf. I’. rohrrsto (Jones aml Hinde. 1 XYO)
Residues were dried at 6O”C, stored in glass phials, and picked for
ostracods using standard micropalaeontological techniques.
5.2. Biostratigvnph>f
5.2.1 Abbots Clzff C‘halk Formation. ‘The Abbots Cliff Chalk (Figure 13)
yielded species typical of the English Upper Cenomanian (Weaver, 1981,
1982). ‘f’he lowest sample (ABCl) contained more than 200 specimens
comprising 16 species, the most diverse fauna in the succession (Figure 13).
Platycopids dominate the assemblage, being represented by four species of
Cytherella and tvvo species of Cytherelloidea. Cytherella ouata (Roemer)
[Figure 1 S(a)] constitutes > 50”,, of the total population. ‘l‘he dominant
podocopids, each comprising - 4’,, of the total fauna are Pontocyprella
klicrofossil Assemblages and the Oceanic Anoslc Event 35
5.2.2 Plenus Marl Formation. Ostracod assemblages from the basal Plenus
Marl (Bed la of Jefferies, 1962, 1963a, b) are similar to those in the
underlying Abbots Cliff Chalk. The lowest sample, however, yielded more
than 300 individuals, with C. oejata again dominating (- 709b) the fauna.
Isocytheresis elongata Weaver [Figure 19(d)] first appears at this level (Figure
13), comprising - 5 Y!bof the assemblage. This species is common through-
out the Plenus Marl at Dover but was not recovered from any other level,
although elsewhere in southern England it is known to be common through-
out the Middle and Upper Cenomanian (Weaver, 1982). Imhotepia euglyphae
Weaver [Figure 19(c)] and Pseudobythocythere colini Weaver [Figure 19(h)]
were only identified in the lower part of Bed 1, although these species are also
known to have more extensive ranges in the Cenomanian (Weaver, 1982).
Phodeucythere cuneiformis Weaver [Figure 16(d)] occurs throughout Bed I,
and is apparently entirely restricted to it (Weaver, 1982). A single adult
carapace recorded from the top of the Abbots Cliff Chalk (Figure 13) was
probably introduced from the basal Plenus Marl as part of a burrow fill. The
upper part of Bed 1 (lb of Jefferies, 1962, 19630, 6) yielded a similar
assemblage to the basal portion except for the absence of I. euglyphae and P.
colini and the occurrences of Cytherelloidea bonnemai Weaver [Figure 1 S(e)]
and Cythereis sp. A [Figure 18(a)-(d)], both of which are first recorded in the
underlying Abbots Cliff Chalk (Figure 13). A significant fauna1 change
occurs at the top of Bed 1, with the disappearances of four species,
C_vtherelloidea bonnemai, Oertliella alata Weaver, L.? bluebellensis and P.
cuneiformis.
Assemblages from Beds 2-3 contain fewer individuals (- 100) than those
from Bed 1 (Figure 13), but are of comparable diversity. Bythoceratina
umbonata (Williamson) [Figure 16(f)] is first recorded in Bed 3 and ranges
into the basal Dover Chalk, although it is known to occur throughout the
Cenomanian in southern England (Weaver, 1982). A major decline in
diversity begins in Bed 4 which yielded the last occurrences of P. robusta, B.
pseudoseptentrionalis and Herrigocythere donzei Weaver [Figure 19(a), (b)],
and the first appearances of Cytherelloidea hindei Kaye [Figure 1 S(f)] which
occurs sporadically throughout the remainder of the sampled interval, and
Pterygocythere sp. cf. P. robusta (Jones and Hinde) [Figure 16(j)]. The latter
species was recorded only from Beds 4-7 (Figure 13), although elsewhere it
occurs as a rarity lower in the Cenomanian (Weaver, 1982). A high
proportion of Cytherella specimens in Bed 4 and the higher parts of the
Plenus Marl (Beds 5-S) show presumed dissolution effects, with valves
displaying rough, etched surfaces, in contrast to their normal smooth
appearance.
Diversity continues to fall in Beds 6 and 7 (Figure 13), which yielded
assemblages containing only eight species. These assemblages are dominated
by platycopids, the only podocopids being Pterygocythere sp. cf. P. robusta,
I. elongata and Cythereis sp. A (the last species being represented only by
Cenomanian Turonian
ABBOTS CLIFF CHALK PLENUS MARL DOVER CHALK
-
0-O -0-o-o-a B -0-o
*
. .
--
-a -_
0 o-o-
7 5
39 Turonian
2 u, E; DOVER CHALK 0”
E? .-
c”.g
3 c- Shakespeare Cliff Member .s
iVicrofossi1 .ksemblages and thv Oceanic ;\nosic Event 39
‘igurr 15. Cenomanian-Turonian ostracods from Dover I. SEM photomicrographs showing represen-
tative specimens of Cytherella spp., Cvtherelloidea spp. and Rairdoppilata pseudoseptentrionn1i.s; all
external lateral views. Th e f o 11owmg .. abbreviations are used in the description of Figures 15 IY:
>I = male; F = female; RV = right valve; I>\: = left valve; car. = carapace; ext. lat. =external lateral
view; int. lat. =internal lateral view; dors.=dorsal xieu. All specimens illustrated have hren
deposited in the British Museum (Natural History); catalogue numbers (OS 1234 etc.) refer to thl
collections of the BM(NH). (a) (‘y~herrllrz ovnt~r Rocmer, AHC‘I, F, RV, oSl312Y, > cr.<;
(1~) C’Jtherrlla sp. cf. c’. chathumrnsis Weaver. XBCI, ~u\enilr, K\‘, oSl 3132 x 0.i; (c) ~‘~thrll~i
corrrar’a Weaver, ABCI, F. R\‘, oSi310. x h5; (d) ~~rlrerrlla sp. cf. ( ‘. contracta \ an Vren, AHC’I
1;.RV, oS1 3 131, x 65; (e) Cythrrelloidea bot~nrmai IYra\ er. AIK‘l, F, I,\‘. oS13092, x X0; (f) CI\,tlwr
tlloidea hindei Kaye, ABC1 3, F, RI’, oS13 142 x 75;(g) (‘_~~thrrr//oidertr ohliqzrirugata (Jones 8r Iiinde).
.AKSll, F:, RV, oS13144, x 00; (h) ~‘~hwllmkn krrwi M’WVCT-, ARCl. F, R\‘, oS1 i 13-I.
x 90; (i) C’ytherelloidea strrcto (Jones & Hlnde) XH(‘Z. I:. 1.1.. 051 WW. xX0: (j) Raivdopp~lcrtcr
ptCl~dOS~~t(‘ntrlOIlNllS \\lel-tens. -\RC0. I,\‘, &13Olft j -!.5.
ure IO. Cenomanian-‘l’urr,nidn ostracods from Dover II. SEM photomicrographs shox+Ing re
sentatl\e spccimcns of Auirdoppilntrr spp., Porrtocyprello robustu, Phodeucytherr cuneijormis, ‘BZ’-
thocrratinu spp. and Pterygorythe “pp.; all extcmal lateral viexks. Abbreviations arc describe zd ‘in
Figure 15.(a) Bairdopilnta southeuhanwrsis Weavrr, ABCI, l.‘i:, oS13139. x 50; (b) Bairdopilat asp.
A, AKSI3, LV. oS13154, x 60; (c) Pmtoc~prella robusta Weaver. XBC3. I>\‘, 0513141, x 7.0;.
(d) Phodeuc_vtkerr cme~jo~mzs W’ea\w ABC7, RV, oS13 193, x 125; (c) Rythoceratina h<v-n@
Weavrr, ARCl, F, I,)‘, r&13089. x 75; (f) f?ythocwstitz~umbortataWilliamson, ABC19, LV.
oS13 117. x 35; (g) Bythorrrntinu umbonutoides &ye. ABCl, l,V, 0513088, x 75; (h) Pfel ‘yggo-
cythrre sp. cf. P. dintinuta Weaver, ABCl9. LV, us131 18. x 70; (i) Pterygocythere sp. cf. P. lati-
rristata (Bosquet), .4X9, RV, oS13 101, x 60; (j) Ptrrygrxythere sp. cf. P. rohustn(Jones and
Hinde), ABC13, I,V. oS131 17, x 60.
Microfossil Assemblages and the Oceanic Anoxic Event 41
Fig1 are 17. Cenomanian-Turonian ostracods from Dover III. SEM photomicrographs show ,ing
representative specimens of Mosaeleberis sp. A. Abbreviations are described in Figure 15. (a) AK S3,
M, LV, ext. lat., oS13124, x 75; (b) Detail of dorsomedian area of (a), x 200; (c) AKS2, F, LV, (ext.
lat.,oS13123, x75;(d)AKS12,F,RV, ext. lat.,oS13148, x 75;(e)AKS12, F, LV,ext. lat.,oS131 49,
x 75; (f) Detail of dorsomedian area of (e), x 200; (g) AKS13, F, LV, ext. lat., oS13126, x 75;
(h) AKS13, F, LV, int. lat., oS13127, x 75; (i) AKS13, F, car., dors., oS13152, x 75; (j) AKS13 . F.
RV, int. lat.. oS13150, x 75; (k) Detail of anterior hinge element of (jj. x 350.
42 1. ~;11.\.is rl a/.
Figure 18. Cenclmanian-‘l‘ur(,nlan ostracods from Dover IV. SE41 photomicrographa shwwng repre-
sentative specimens of C’vthereis spp. and C’urfsina senior.Abbreviations are described in Figure 15.
(a) Cythereissp. A., ARCS, F, R\‘. ext. lat., oS13119, x 65; (b) (‘ythereis sp. A., ARCI, M, RV, ext.
lat.. oSl3085, x 65; (c) C’yther-ris sp, A-\..AR~‘X,~u~enile, RY. est. lat.. oS13120, x 65; (d) C~derris
sp. A.. ABCl, 141. RV, int. lat., oS13086. x 65; (e) C’ythereis sp. cf. C’. longaeua Porkom$, AKSl, LV,
ext. lat., oS13146, x 60; (f) Detail of posteroventral part of(e), x 150; (g) Curjsina senior PorkornG,
AKS, ?F. LV, ext. lat., oS13147. x 75; (h) Detail ofposterodorsal areaof( x 350; (i) Curfsinasenior
Porkom~, AKSIO, ?X’l. car., right wt. lat.. oSl3128. x 75; (j) &tail of dorsomedian area of(i),
x 3.50.
Xlicrofossil Assemblages and the Oceanic Xnosic Event
F;IyuI.c IO. (‘cnomanian-‘l’urcjnian ostracods from Dover 1.. SEM photomicrographs showing rrl >rc-
sentative specimens species. all external lateral views except (j). Abbreviatiom
of several arr
described in Figure 15. (a) Hevrigoc\jthere ciorzzcG (\!‘eaver), ABC9, R\‘, 0S13107. x 90:
(b) Herrigocythere donzri (Weaver), ARCI, LV, oS13091, x 90; (c) Imhotepia euglyphea W’ra ver-.
ARC7, F, car., left side, oS13098, x 100; (d) Isocytherrrs elongate Weaver. ABC7, car., left s ide.
nSl3102, x95; (e) Parewcythereis suhpawa (Porkom+), AKSll, R\‘, oS13153, x 100; (f) Oe, vtli-
r//n domei Weaver, .4BCl. LV, oS13135, x 100; (g) Oertliefla akatcz Wezaver, ABC%, LV, oS131 121,
x 90; (h) Pseudobvthoc~lthera colini Weaver, .4BC7, F, RV. oS13100. x 150; (i) Lo.tocont ./lU?
hlrtebel/en.sis Weaver. ARCI, F. I,V. oS13138, x 105: (j) I,oxoconcha? bluebellmsis Weaver, ABC’1 , 1:.
KY, int. lat., oS13137, x 10.5,
in sample ABC1 5, and fragments of Bythoceratina hevrigi and B. umbonata
in sample ABC1 7.
Lowest Turonian sediments taken from immediately above the basal
hardground sequence (samples ABC1 9, 20), however, yielded more abund-
ant ( - 150 specimens) and diverse (6-9 species) assemblages. These are again
dominated by five platycopid species (Figure 13), with C. ozlata forming
> 700, of the total fauna. Podocopids are limited to four species, which
include the highest occurrences of B. herrigi (sample ABC19) and
B. umbonata (sample ABC20). Pterygocythere sp. cf. P. diminuta Weaver
[Figure 16(h)] was recorded only from this level. Sample ABC21 yielded
only a few Cytherella spp.
Higher material from the lower ‘l’uronian (samples AKSl-13) contained
assemblages which continue to be dominated by platycopids, particularly
ci. ozlata (Figures 13, 14). Diversity gradually increases through the
Shakespeare Cliff Member, however, as species which are important in
the higher Turonian at Dover (on the evidence of our own limited studies)
appear for the first time. First appearances in the Melbourn Rock are
Cytheveis sp. cf. C. Zongaeva Porkorny [sample ASKl; Figures 18(e), (f)]
and Curfsina senior Porkorny [Figures 18(g)-(j)] with Mosaeleberis sp. A
(sample AKS2; Figure 17). The new species Parvacythereis subparva
(Porkorny) [Figure 19(e)] and Cytherelloidea obliquirugata (Jones and Hinde)
[Figure 15(g)] first appear in the upper beds of the Shakespeare Cliff
Member, at the level of the Round Down Marl (Figure 14). Complete adult
valves of Bairdoppilata sp. A [Figure 16(b)], also common higher in the
Turonian, were only found in our top sample (AKS13), although juvenile
or fragmentary specimens from several lower horizons are tentatively
assigned to this species (Figures 13, 14).
The diversity of the ostracod assemblages at the top of the sampled
interval, however, is still only half that of assemblages recovered from the
upper Abbots Cliff Chalk and basal Plenus Marl. Preliminary studies of more
widely-spaced samples taken throughout the low and mid-Turonian at Akers
Steps, however, indicate that the trend to higher diversity is continued in the
overlying beds of the Dover Chalk.
Pterygocythere sp. cf. P. robusta (Jones and Hinde, 1890) [Figure 16(j)]
Our specimens, like those of Weaver (1982), p assess a short longitudinal rib
just below the posterior half of the dorsal margin; in this respect they differ
from those illustrated by Neale (1978) ( as Alatacythere robusta) which
resemble P. diminuta Weavrer.
5.4 Discussion
Assemblages throughout the section are dominated by platycopids, parti-
cularly Cytherella ozrata. One species, C_vtherelloidea bonnemai disappears at
the top of Bed 1 of the Plenus Marl, to be replaced by C. hindei which first
appears in Bed 4 (Figure 13). Podocopids, on the other hand, show a dramatic
decrease in diversity through Beds 1 -8, particularly at the tops of Beds 1 and
4. lsocythereis elongata is the only podocopid to occur in every Plenus Marl
sample and only three podocopids appear to have struggled through into the
overlying Dover Chalk. One of these, Cythereis sp. L4 is represented only by a
single fragment at the base of the Melbourn Rock, while the others, both
species of BJtthoceratina, disappear in the basal few decimetres of the
‘I’uronian.
The refilling of the niches left vacant by these disappearances was ver)
gradual. l‘he lower numbers of ostracods obtained from samples above the
Plenus Marl can undoubtedly be attributed, at least partly, to difficulties in
processing more indurated chalks. ‘l‘he evidence from the easily disag-
gregated samples from the Plenus Marl suggests, however, that they do, to
some extent, reflect a real reduction in abundance as well as diversity.
Work on Cenomanian ostracods in southern England by Weaver (1982)
included bed-by-bed sampling of the Plenus Marl at Shillingstone (Dorset)
and Pitstone (Hertfordshire), which demonstrated that major changes in the
ostracod fauna occur through these sequences. The changes recorded by
Weaver (1982) are generally similar to those demonstrated by us at Dover,
but they differ in detail. At Dover the major disappearances occur at the
tops of Beds 1 and 4, but at Pitstone the tops of Beds 1 and 2 are the sig-
nificant levels. At Shillingstone only the top of Bed 2 shows a major effect,
although several species disappear at the top of the underlying ‘Grey Chalk’
(= Abbots Cliff Chalk).
The sequence of changes which occur in the ostracod assemblages appears
to be virtually the same at all localities. Losoconcha? bluebeltensis and
Imhotepia eugl_vphae (the latter \vas not recorded at Shillingstone; Weaver,
1982) are, for instance, the first species to disappear. HervigocJltheve donzei
and Pontocypvella robusta go next, at the top of Bed 2 at Shillmgstone and
Pitstone, but at the top of Bed 4 at Dover. Isocytheveis elorzgata survives as far
as Bed 8 at all localities. Cythevelloidea hindei first appears in Bed 4 at Dover,
in Bed 3 at Pitstone, but at Shillingstone is occurs throughout the Plenus
Marl as well as in the underlying ‘Grey Chalk’, and at Southerham (Sussex)
Weaver (1982) recorded a few specimens in the Middle Cenomanian. It
would be premature to speculate on the significance of these geographical
differences in the lithostratigraphic distribution of the different species. It
does seem, however, that appearances and disappearances are stratigraphi-
tally lower to the west and south of Dover. ‘l’he extension of our current
investigations to other localities, both in England and on the Continent,
should be informative in this respect.
Microfossil Assemblages and the Oceanic Anosic Event 47
Weaver (1982) also examined a few samples from above the Plenus Marl,
noting difficulty in processing the harder chalks, but those species which he
did record are essentially the same as those found at Dover in the Melbourn
Rock: Bythoceratina herrigi, B. umbonata, Cytherelloidea hindei and numer-
ous Cytherella spp.
Detailed ostracod records are not available for other areas of southern
England or northern France. Studies of ostracods in the Cenomanian-
Turonian boundary sequences of the Saumur (Maine-et-Loire) and Givray-
de-Touraine (Indre-et-Loire) areas (Damotte in Robaszynski et al., 1982),
SW Anglo-Paris Basin, have revealed only low diversity faunas. These have
few species in common with those in southern England and a direct
comparison is not possible.
Samples were 5 g splits taken in the field from 2 kg samples collected for
microfossil and stable-isotope analysis. Thirty-two samples were analysed,
sample numbers AKS9 and 13 not being taken for calcareous nannofossil or
stable-isotope analysis.
Calcareous nannofossils were extracted after first soaking the samples in
Calgon overnight. Disaggregated fines were suspended by shaking, an
aliquot being pipetted-off and used to produce a smear slide. The remaining
suspension was poured-off and centrifuged until a clear solution was
obtained. A second concentrated smear-slide was produced from the cen-
trifuged residue. Nannofossils were identified using light microscopy on
both the original and preconcentrated smear slides.
6.2 Biostratigraph)
6.2.1 Abbots Cliff Chalk Formation. The Abbots Cliff Chalk yielded 58
nannofossil species (Table 3), with diversities ranging from 34-45 species in
individual samples. The ranges of 21 species are shown in Figure 20. The
‘l’ahlc 3. Alphabetical list of nannofossil species and their stratigraphIc distribution (.4(‘, .4bhots (‘lift
Chalk [upper Capel-lr-Frrnr 1lcmbrr]; PM, PI enus Marl Formation; DC, Dover Chalk Formation
[Shakespeare (‘liff LIemhrr]).
Species Distribution
I MRB
*-•- o-0-0
o-o-o-o-
o-o-o-.-
o-o-
& ..j:,
-
..::.
0 - w+<
;i
o-o- l -o-o--_~*,~
l-•-•-•-.-•_+
Microfossil Assemblages and the ( keanic .Anos~c Event 51
i i i
I I I
I 7 i
l 0 i
I I I I
l !, l 0
52 1. JanIs rt LZl
podorhabdus sp. cf. T. decorus (Deflandre in Deflandre and Fert) Wind and
Wise in Wise and Wind, ” Vekshinella” dibrachiata Gartnkr and Zygodiscus
elegans Gartner. Of these, only the disappearances of A. albianus, C. striatus,
L. acutum, and R. asper were believed to represent extinction levels, which all
occur towards the top (Beds 7 and 8) of the formation (Figure 20). Other
species are known to have more extensive stratigraphic ranges elsewhere.
6.3 Discussion
MRB I
1’Iicrofossil A4ssemblages and the Oceanic Anoxic Event 55
I
0 0
_ - _ .
I .
i I I
Quadrum gartneri appears (marking the base of the Q. gartneri Zone) within
the low Turonian at Cap Blanc-Nez (Manivit, op. cit.), some distance abovre
the base of the stage, and possibly, therefore, stratigraphically higher than at
Dover. The disappearances of Axopodorhabdus albianzts close to the
Cenomanian-Turonian boundary, and of Prediscosphaera columnata in the
lower Turonian, occur at similar levels at both localities. Lithraphidites
acutum, which disappears in the Upper Cenomanian at Dover, is recorded as
ranging into the lower Turonian at Cap Blanc-Nez (Manivit, op. cit.).
The first appearance of Nannoconus multicadus around the level of the first
Q. gartneri has been noted at both Dover and Cap Blanc-1Vez, although
N. truittii is recorded higher at the former locality. It can be concluded,
therefore, that there is good general agreement between nannofossil
records from Dover and Cap Blanc-Nez. Minor discrepancies are probably
due in part to the (presumably) larger number of samples analysed from the
Cenomanian-Turonian boundary sequence at Dover.
Comparative calcareous nannofossil data have also been published for the
Turonian stratotype region of Touraine in the SW Anglo-Paris Basin.
Working in the area around Saumur (Maine-et-Loire), Manivit and Zeigh-
ampour (in Robaszynski et al., 1982) recognised both the M. decoratus ‘Zone’
and the Q. gartneri Zone in the succession. Similar data were presented
(Manivit in Robaszynski et al., 1982) for the Civray-de-Touraine (Indre-et-
Loire) borehole.
Several species are common to both the Dover and Touraine sequences,
but there are some differences in detail. Quadrum gartneri appears some
metres above the base of the Turonian in Touraine (Manivit, op. cit.).
Axopodorhabdus albianus occurs only in the low to mid-Cenomaman ot
Touraine, and does not extend up into the M. decoratus ‘Zone’, while
Ahmuellerella octoradiata appears earlier than at Dover, within the low
Cenomanian. Prediscosphaera columnata disappears in the lowest Turonian
in western France (lower than at Dover), while Lithraphidites acutzdm ranges
higher, disappearing only at the summit of the lower Turonian. Manivit
(op. cit.) noted that species of Nannoconus are generally rarer in the
marginal calcarenitic and detritus-rich sediments of Touraine than they are
in coeval chalks, although at Dover we have noted that Nannororzus first
becomes abundant in the calcarenitic chalks of the Melbourn Rock.
The variations in the ranges of nannofossil species between Dover and
Touraine is probably the result of provincialism, but more data are required
from sections elsewhere in the Anglo-Paris Basin to identify the palaeo-
geographical factors which might control the stratigraphic distribution of
Cenomanian-Turonian calcareous nannofossils.
Table 4. Alphabetical list of dinotlagellate cyst species and their stratigraphic distribution (AC, Abbots
Cliff Chalk [upper Capel-le-Ferne Member]; PM, Plenus XIarl Formation; DC, Dover Chalk
Formation [Shakespeare Cliff Member]).
Species Distribution
.dldo$ia d@andrei (Clarke and Verdirr, 1967) Stover and l<\,ltt, 197X xc
~n~~~rcasplzaern euteiches (Davey. 1969) Davey. 1979 AC, P>l, DC
C’anrtingia collic~eri Cookson and Eisenack, 1960 AC, PM, DC
Cleistosphneridiun~ ? aciculare Davry, 196Y AC, PM
C. mmntum(Deflandre, 1937) Davry. 1969 AC
C. ~lnvuluw~ (Davey, 1969) Below, 19X2 AC. PM. IX’
(‘rihroperidinium ewilicristatum (Davey. lY69) Stover and I:vitt. lY78 ilc
fJ_~c/onephelium distincturn Deflandre and Cookson. 1955 .4C. PM. DC
(‘. mmbroniphortm Cookson and Eisenack, 1962 AC. PM. DC’
E.w&osphaeridium bi’dztnz (Clarke and Verdier, 1967) Clarke et ni. lY68 A<‘
E. pbragmites Davey et al.. 1966 PM
E‘romae granulosa (Cookson and Elsenack, 1974) Stover and Evitt, 1978 AC
Hete~osphoeridilrm ? heterrrranthum (Defiandre and Cookson, 1055)Eisrnack
and Kjellstrom, 1971 AC. PM, DC
H~ctrirllosphaeridizrm dijjicile Manum and Cookson, 1964 IX
H. palmatum (White. 1842 ex Bronn. 1848) D ownie and Sarjeant, 1965 AC. PM, DC
Kallnsphnrridium sp. B Tochcr and Jarvis, 19X7 DC
K. rlrrgnesiorum (.22anum and Cookson, iY64) Tocher and Jarvis, lYX7 1’11, DC
Kiokansium pol~$as (Cookson and Eisenack. 1962) B&W. 1YX2 AC‘, Phi
Lebwidocysta chlnm_ydattr (Cookson and Eisenack, lY62) Stover and Evitt,
197X AC
1, d&ccata (Davey and Verdier, 1973) Stover and Evitt, I Y78 .4C. DC
Odrmtochitina mstata Albcrti, 196 1; emend Clarke and Verdier. 1967 AC, I’M, DC’
0. nprrculata (0. Wetzel. 1933) Dellandre and Cookson, lY55 XC‘, PXI, DC‘
O/iec)sphaeriditcNI albertense (Pocock, 1962) Davey and \Villiams. 1 Y60 XC
0. rwnples (N’hite. 1812) Dave! and Williams. 19h6 A<‘, I’>l. 1x
0. p~~c~ulun~ Jain, 1977 AC‘, DC
0. prolixispinosum Davcy and $Villiams, 1966 AC’, PM. DC
Prolr.~osphaeridiiunr conulun~ Davry, lY6Y .A\(‘, PZI. DC
Ptrrodinium cingularmr vrtiruiutzcnt (Davch- and Williams. I ‘MI Lentin and
\Vllliams, 1981 AC
Rh>whodiniopsiv sp, cf. R. aptiona Drfandre, 1935 AC
SanoGzsphaera mirroreticrtlata Rrideaus and McIntyre. I’)75 DC
,S. wtundata Clarke and Verdier. 1967 rx-
SI,,,tllsldirrirrrr,~1’. .A ‘I’ocher and Jat-1-x. IW7 1’.\1.DC:
S sp. 13Tocher and Jarvis, 1987 Lx:
S. sp. C Tocher and Jarvis, 1987 P1I. DC
Slc~rrllosphueridium? longifrtrcatum (Flrtion, 1952) Dave? et (II.. 1966 .4C
?;rn?,osphaeridiunr variecalanrus Dave? and Williams, 1966 P1\1
,Gwccwr.s cerntioides (Dellandre. lY37) Lentin and Williams, lY73 XC
r
3
0
0
2
.-.-
R;licrofossil Assemblages and the Oceanic Xnosi,, Event
.. 0
I
0
Number of specimens
0 single 0 51-100
. 2-10 l 101-200
0 11-20 l 201-500
8 !I-50 501-1000
0
. . .
RDM
.
. 0 .
. .
I .
.
I
.
I .
.
I I .
I I
0 1 . ;. 0
. .
.
I
I .I .
1
I
I
.
Lithology
Figure 23. Dinofagellatr cyst distribution and lithostratigraphy in the higher beds of the lower
‘ruronian at Dover. The succession is a continuation of that shown in Figure 22. Lithologies and
symbols are explained in Figures 5. 6 and 13. The horizontal dashed line indicates the top of the
Melbourn Rock.
Microfossil Assemblages and the Oceanic Anoxic Event 61
0
.
.
7.2 Biostratigraphy
7.2.1 Abbots Cliff ChaZk Formation. Samples from the Abbots Cliff Chalk
yielded dinoflagellate cyst assemblages typically consisting of several
hundred to one thousand individuals (Figure 22). Twenty-seven species and
subspecies were recorded (Figure 22; Table 3), eleven of which are restricted
to this formation. Assemblages invariably contained Oligosphaeridium com-
plex (White) Davey and Williams as the most abundant cyst, accompanied
by common Batiacasphaera euteiches (Davey) Davey [Figure 24(c)],
Cyclonephelium distinctum Deflandre and Cookson [Figure 25(g)],
C. membraniphorum Cookson and Eisenack, Hystrichosphaeridium palmatum
(White ex Bronn) Downie and Sarjeant, Odontochitina costata Alberti;
emend. Clarke and Verdier [Figure 24(j)] and 0. operculata (0. Wetzel)
Deflandre [Figure 24(i)]. All of these species continue up into the overlying
Plenus Marl (Figure 22). Other common constituents were Cleisto-
sphaeridium armatum (Deflandre) Davey, C. clavulum (Davey) Below,
Prolixosphaeridium conulum Davey [Figure 25(h)], Pterodinium cingulatum
reticulaturn (Davey and Williams) Lentin and Williams [Figure 24(b)] and
Rhynchodiniopsis sp. cf. R. aptiana Deflandre [Figure 24(g), (h), (k)].
Eleven species were restricted to the Abbots Cliff Chalk (Figure 22) and of
these, seven [Aldorfia deflandrei (Clarke and Verdier) Stover and Evitt,
Cleistosphaeridium armatum, Exochosphaeridium bifidum (Clarke and Ver-
dier) Clarke et al., Leberidocysta chlamydata (Cookson and Eisenack) Stover
and Evitt, Pterodinium cingulatum reticulatum, Surculosphaeridium? longifur-
catum (Firtion) Davey et al. and Xenascus ceratioides (Deflandre) Lentin and
Williams] have been recorded from post-Cenomanian sediments elsewhere
in the Anglo-Paris Basin (Clarke and Verdier, 1967; Foucher, 1979, 1981;
Foucher in Robaszynski et al., 1980, 1982; Jarvis and Tocher, 1983; Tocher
and Jarvis, 1987; Jarvis et al., 1987, 1988).
Cribroperidinium exilicristatum (Davey) Stover and Evitt and Fromea
granulosa (Cookson and Eisenack) Stover and Evitt [Figure 25(k)] were also
restricted to the Abbots Cliff Chalk, and have not been recorded from
sediments younger than Cenomanian elsewhere in southern England
(Davey, 1969; Jarvis et al., 1988). Oligosphaeridium albertense (Pocock)
Davey and Williams [Figure 25(i)], a third restricted species, is recorded for
the first time from the Anglo-Paris Basin but elsewhere it ranges throughout
the mid and upper (Barremian to Maastrichtian) Cretaceous (Yun, 1981;
Below, 1982).
A cyst referred to Rhynchodiniopsis sp. cf. R. aptiana occurred only in the
lllicrofossil Assemblages and the Oceanic Anosic Event 63
(i)
* ”
1~1g:ure 25. Crnoman~an-‘l’u~~)~llan dmofla~ellate cysts from Dmw 11. Light phot~)mlc~oKraphs of wmc
representative sprcnnens. (a) Oligospharridium poculum Jain. AKS/D 19, x 300;(h) Oligosphaeridirrm
poculum Jain, XKS/DlO, x 300; (c) Kallosphaeridium ringnesiowm (Illanum and Cookson) Tocher
and Jarvis, ABC/DlS, x 300; (d) Kallosphaevidium ringnesiovum (1lanum and Cookson) Tocher and
Jar\is, ABC;D15,, x 300; (e) Oligosphaeridiumpvolixispinosum Davey and Williams. ABC/D-b, x 300;
(f) Oligosphaeridzum pwlixispinosum Dabey and Williams, ABC/D+, x 300; (g) (‘_wlone~helirrm
distincturn Deflandre and Cookson, ABC/Dl2. x 300; (h) Pvolisosphaeridium conulum Davey,
ABC/D6, x 300; (i) Oligosphaeridium albertense (Pocock) Davey and Williams, ABC;D4, x 300; (j)
Leberidocysta dej7occata (Davey and Verdier) Stover and Evitt, ABC:I>Z, x 300; (k) Fromeagmnulosu
(Cookson and Eisenack) Stover and Evitt. ABC/D6, x 300; (1) Knllospkar~idium vingnesiorum
(\\lanum and Cookson) Tocher and Jarvis, ABCiDl-l, x 300.
Microfossil Assemblages and the Oceanic Anoxic Event 65
Abbots Cliff Chalk [Figures 22, 24(g), (h), (k)]. Rhynchodiniopsis uptiana has
previously been recorded from Aptian and older sediments (Deflandre, 1935;
Below, 1981). It is unclear whether our material from Dover is referable to
that species or represents a closely related form. If the former can be
demonstrated, then our records indicate that either the range of the species
should be extended into the Late Cenomanian, or that reworking of Aptian
sediments occurred during the Late Cenomanian in the Dover area.
7.2.2 Plenus Marl Formation.. The Plenus Marl yielded nineteen species and
subspecies of dinoflagellate cysts (Figure 22; Table 3). Assemblages in the
lower beds (l-3) are dominated by the same seven species which are
commonest in the Abbots Cliff Chalk (Figure 22), although 0. complex does
not invariably remain the most abundant cyst. The numbers of cysts
recovered from the lower beds of the Plenus Marl are generally lower (several
horizons yielded < 100 individuals per sample) than in the underlying
formation, and specific diversity is also lower (Figure 22). Cleistosphae-
rid&m? aciculare Davey [Figure 24(e)] and Kiokansium polypes (Cookson
and Eisenack) Below, which occur rarely in the Abbots Cliff Chalk, both
disappear within the lower beds of the Plenus Marl (Figure 22). The former
species has not been recorded in the Anglo-Paris Basin before, although it
has been found in Albian-Cenomanian sediments elsewhere (Davey, 1969),
whereas the latter is known from post-Cenomanian sediments within the
basin (e.g. Tocher and Jarvis, 1987). Heteros@zaeridium? heteracanthunz
(Deflandre and Cookson) Eisenack and Kjellstrom, Oligosphaeridium pro-
lixispinosum Davey and Williams [Figure 25(e), (f)] and Prolixosphaeridiunz
conulum [Figure 25(h)] also disappear in Beds l-3, but all reappear in the
lower Turonian.
.4 pronounced drop in cyst abundance (samples typically contained 50 or
fewer individuals per sample), and diversity occur in Bed 4, followed by the
progressive disappearance of most cysts in Beds 5 and 6 (Figure 22). This
decline is, however, initially accompanied by increased proportions of
Kallosphaeridium ringnesiorum (Manum and Cookson) Tocher and Jarvis
[Figure 25(c), (d)]. This species is as a minor component of the assemblage in
Bed 1 (Figure 22) but constitutes 60°0 of the cysts in Bed 5 and 96”, in Bed 6.
The uppermost beds of the Plenus Marl contain few dinoflagellate cysts, Bed
7 yielding only two specimens of K. ringnesiorum and Bed 8 a single example
of Odontochitina operculata. The appearance of K. ringnesiorum in the upper
part of Bed 1 at Dover, is the earliest record of this species in the Anglo-Paris
Basin.
The distribution of dinoflagellate cysts through the Plenus Marl suggests
that the environmental changes, which in Bed 4 caused the decline of the
dominant cyst assemblage, initially favoured K. ringnesiorium, allowing it to
reach an acme immediately prior to the disappearance of all cysts at the
bottom of the Dover Chalk.
It is noteworthy that the decline in dinoflagellate cysts coincides with a
marked increase in the proportion of calcispheres (Section 2.2). At least some
calcispheres (particularly Pithonella spp., a form which is common in the
Plenus Marl) may be calcareous-walled dinoflagellate cysts (Keupp, 1979).
The inverse relationship between the two groups, therefore, suggests that the
decline in organic-walled dinofagellate cysts during the latest Cenomanian
and earliest Turonian was balanced by an increase in calcareous-walled
groups.
7.3 Discussion
at Dover are characterised by the same species which are abundant in the
underlying Abbots Cliff Chalk. The upper beds (S-7), however, contain
mainly Kallosphaeridium ringnesiorum. While forms such as C. distinctum and
0. complex are relatively common throughout the basin (e.g. Foucher in
Robaszynski et al., 1980, 1982), B. euteiches and K. ringnesiorum are rarer.
The latter two species are thick-walled proximate cysts [Figures 24(c), 25(c),
(d)], and have not been recorded from the Plenus Marl or its equivalents at
any other localities.
Dinoflagellate cyst assemblages from the Plenus Marl in the Pas-de-Calais
(Foucher, in Robaszynski et al., 1980) and Isle of Wight (Clarke and Verdier,
1967), and the Plenus Marl equivalent in SE Devon (Jarvis et al., 1988)
continue to be dominated by Palaeohystrichophora infusorioides and Spin~fe-
rites ramosus, as in the underlying chalks, although a local abundance of
Microdinium irregulare Clarke and Verdier occurs in the Isle of Wight
succession. The assemblage described from the lateral equivalent of the
Plenus Marl in Saumur (Foucher, in Robaszynski et al., 1982) was relatively
diverse but abundances were low, and no particular form was noted as being
dominant. Nevertheless, this assemblage also contains P. infusorioides and
S. ramosus. ‘fhe anomalous composition of the dinoflagellate cyst assem-
blages noted in the Abbots Cliff Chalk, therefore, continues through the
remainder of the Cenomanian at Dover, prior to the disappearance of all
cysts near the top of the stage.
Samples from the basal 4m of the Turonian Dover Chalk Formation
at Dover proved barren, but above this Oligosphaeridium complex and
Cyclonephelium distinctum are again the dominant cysts. There are also
localised abundances of Batiacasphaera euteiches, Kallosphaeridium
ringnesiorum, Senoniasphaera microreticulata and the low ‘l’uronian indices
Hystrichosphaeridium dificile and Senoniasphaera rotundata.
By contrast, an assemblage recorded from a comparable interval on the Isle
of Wight (Clarke and Verdier 1967) is dominated by Spiniferites ramosus and
Odontochitina costata, and to a lesser extent by Cyclonephelium membrani-
phorum and Canningia colliveri. In SE Devon (Jarvis and Tocher, 1983;
Tocher and Jarvis, 1987; Jarvis et al., 1987, 1988) the most abundant
forms are Palaeohystrichophora infusorioides, S. ramosus and Cyclonephelium
distinctum. Samples from the ‘Craie marneuse’ of Saumur (Foucher, in
Robaszynski et al., 1982) are dominated by 0. operrulata and Oligosphae-
ridium complex but contain common Spiniferites and P. infusorioides,
while at Civray-de-Touraine the assemblages are also dominated by the
S. ramosus group and P. infusorioides.
Clearly, therefore, there is good evidence for significant geographical
variation in Cenomanian and Turonian dinoflagellate cyst assemblages, with
the Dover samples in particular proving atypical.
Table 5. Carbon and oxygen stable-isotope data for uppermost Cenomanian to basal Turonian chalks,
Dover. Sample numbers correspond to those in Figure 3. All data are bulk sediment determinations
(ND, not determined; Bed numbers after Jefferies [1962, 19630, b]; MRB, Melbourn Rock Beds).
Carbon isotope values decline steadily in the basal beds of the overlying
Dover Chalk (Figure 26), reaching values of 3.0x, approximately 2 m above
the base of the formation. Remaining samples in the lower Dover Chalk
exhibit values of 2.6-3.1x, 613C (Table 5). Th e carbon stable-isotopes thus
display a major ‘excursion’ occurring in the Plenus Marl and basal Dover
Chalk (Melbourn Rock Beds). The peak of the anomaly occurs at the top of
the Plenus Marl, immediately below the formational boundary and, there-
fore, also below the base of the Turonian as defined by appearance of
Mytiloides inoceramid bivalves (Section 3.4).
Comparable carbon stable-isotope data through the Plenus Marl have
recently been presented by Schlanger et al. (1987, figure 11) for shore
sections at the eastern end of Shakespeare Cliff (Figure 1). These authors
additionally sampled the uppermost 1.5 m of the Abbots Cliff Chalk and the
basal 1 m of the Dover Chalk (Melbourn Rock Beds). Schlanger et aI.‘s
(1987) data differ from ours in two respects. Firstly, their 613C values reach a
70 I. Jarvis rt t/l
3c
H. archaeocretacea
.
I
2 1c
02 oc
1 9c
-1 8C
:
;I
i -1 2:
1 1C
DC
,30
13C
70
t50
R. cushmani TRZ
:
:
:
Figure
!
26. Carbon
Cenomanian/Turonian
and oxygen
boundary
c I
stable-isotope
, I
distribution
at Dover. Major lithologies
I
I ’
and iithostratigraphy
and symbols are explained
I ’ I
across the
in Figures 5
and 6.
Microfossil Xssemblages and the Oceanic Anosic Event 71
_ 1:
1;
- 1.
l( 0
H. archaeocretacea
PRZ
c I
I 0
---___ I-
0
_- -
, 0
\
.
0
I
I
0
I I
gurt’ 27. Carbon and oxygen stable-lsotopedistrlbuti(ln and lithostratygaphy III the higher beds of the
lower Turonian at Dover. The succession is a continuation of that shwn In Figure 26. Lithologies
and sgnhol< at’r explained in P~gurr 5. 6 and 8.
maximum within the Melbourn Rock Beds (although still within the
Cenomanian), and not within the upper Plenus Marl; their values do not
show any decline within their sampled interval (despite their uppermost
sample being from - 2 m above the base of the Melbourn Rock). Secondly,
the isotopic variation in the section is greater in our samples (2.3-4.8x, 6ljC)
than that illustrated by Schlanger et al. (1987). Their data display constant
values of - 3x, 6l 3C f rom 1.5 m below the base of the Plenus Marl up to Bed
4, and then an increase to a maximum of 4.2x, in the basal Melbourn Rock
Beds. The reasons for these differences in detail are unclear, but it is
noteworthy that our carbon stable-isotope data correspond more closely with
trends displayed by associated microfossil assemblages (see below).
The oxygen stable-isotope data (Figures 26,27, Table 5) may cautiously be
interpreted as having a ‘background’ value of approximately -2.6x, 6180.
Within the Plenus Marl, the oxygen isotope values show considerable
variation, with a slight positive excursion around Beds 6-8. Within these
beds, there is a positive correlation between 613C and 6lsO values. The most
negative value is - 3 .S%, which occurs approximately 2 m above the base of
the Dover Chalk. Oxygen values return to ‘background’ levels immediately
above the Melbourn Rock Beds (Figure 27). The positive correlation of 613C
with 6180 in the upper beds of the Plenus Marl is contrary to the negative
correlation observed in German sections by Hilbrecht and Hoefs (1986).
These authors interpreted the oxygen data as indicating lower sea-water
temperatures during the latest Cenomanian. Our results correspond more
closely with those of Scholle and Arthur (1980) who noted a drop in al80
values at, or close to, the Cenomanian-Turonian boundary. At this stage,
therefore, we prefer to use the oxygen stable-isotope data solely as a means of
isolating diagenetically altered data points (cf. Jarvis et al., 1988), and to
indicate any obvious diagenetic trends. We do not as yet place any sig-
nificance on the stratigraphic variation in the 6180 curve.
Diagenesis will only alter bulk-sediment isotope values if there has been
significant interaction between the sediment and the surrounding pore-fluid
(cementation or recrystallisation). Such interaction commonly occurs during
early diagenesis in sediments composed of metastable mineralogies (arag-
onite or Mg-calcite). However, the Chalk was deposited as predominantly
low Mg-calcite and, with the possible exception of hardground horizons,
is unlikely to have undergone significant early diagenetic modification.
Furthermore, the carbon stable-isotopes of any metastable mineralogies
which were present would probably not have altered extensively since,
compared with oxygen stable-isotopes, carbon isotopes are relatively
immune to diagenetic modification. This is because the volume of carbon
in carbonate sediments vastly exceeds that in solution within their coexisting
pore waters, and the fractionation of carbon isotopes between calcium
carbonate and dissolved bicarbonate is relatively small at near surface
temperatures (Emrich et al., 1970).
Nevertheless, diagenetic modification of bulk-rock carbon stable-isotope
ratios in pelagic limestones can occur. Precipitation of carbonate cements in
Microfossil Assemblages and the Oceanic Anoxic Event 73
5
0 chalk
1 * marl
4-
.
d3C %o
(PDBI l
-4 -3 -2
Figure 2X. Carbon versus oxygen stable-isotope values for the Cenomanian-Turonian boundary
succession at Dover. Values are listed in Table S. Note the absence of any correlation between the two
sets of data.
changes have had a minimal effect on 6i3C values. Maximum changes may be
up to -0.4x, 613C ( as estimated from adjacent chalk-marl values; Figure
26), but these lie within a positive carbon isotope ‘excursion’ of - 2.5x,.
PM Plenus Marl
1:1gure 29. Biostratigraphlc summary chart for the Cenomaman-Turonian boundary succession at
I>o\-rr. The stratigraphIc ranges of representative macrofossil and microfossil species are indicated
bv the vertical lines. Dashed vertical lines indicate intermittent occurrences; the discontinuous line
w:ithin the Gacelinelln veussi G. berthelini plexus indicates a shift from dominantly G. veussi to
dominantly G. berth&G morphotypes. Lithostratigraphic units are drawn to scale. The horizontal
stippled area indicates the stratigraphic extent of the carbon stable-isotope excursion. Dtscontinuous
horizontal lines mark lithological boundaries (base of the Plenus Marl, base of the Melbourn Rock,
top of the Melbourn Rock). The continuous horizontal line indicates the base of the Turonian.
defined by the appearance of Mytifoidrs spp. inoceramid bivalves.
I
I Cenomanlan I Turontan I
(Plenus Marl Bed 3 event). Major extinctions and disappearances only occur
in the microflora (phytoplankton) at the peak if the isotope excursion in
Plenus Marl Beds 6-8. The Cenomanian-Turonian boundary occurs at a
level where 613C values are declining, typical ‘I’uronian fossil species
appearing gradually throughout the Early Turonian. It is clear, therefore,
that there is a close correspondence between the 6’ 3C curve and a sequence of
floral and fauna1 events which accompany the Cenomanian-Turonian
boundary succession.
isotope excursion are more difficult to assess. In both England and Germany
the top of the excursion lies immediately above the appearance of Mytiloides
spp. (i.e. base of the Turonian). Hilbrecht and Hoefs (1986) suggested that
this is coincident with the appearance of H. ‘helvetica’. This conclusion was
based, however, on data from attenuated successions in SE Germany, the
species being absent to the NW. In addition, the identification of H. helvetica
relies strongly on the species concepts of the individual micropalaeontolo-
gist, since it forms part of the Hedbergella delrioensis-Helvetoglobotruncana
praehelvetica-Hi. helvetica plexus. We record the appearance of H. praehel-
vetica several metres above the top of the carbon stable-isotope excursion at
Dover, while H. helvetica appears more than 20m above the base of the
Turonian, above the interval studied in detail here.
I I I
I I I
mid Cenomanian
basemen,
Figure 30. Schematic representation of the palaeoceanographic changes which may have caused the
Cenomanian-Turonian Oceanic Anoxic Event (OAE). The mid to early late Cenomanian was
characterized by low rates of oceanic turnover, and therefore only low to moderate surface
productivity. An oxygen-minimum zone was probably poorly developed in the water column, being
represented by relatively well-oxygenated water. Widespread deposition of nannofossil oozes took
place everywhere above the carbonate compensation depth (CCD). A rapid increase in the rate of
oceanic turnover during the latest Cenomanian (possibly associated with a major transgressive pulse)
caused widespread upwelling of nutrient-rich deep-ocean water, particularly along the NW
European continental margin. A consequent increase in surface productivity caused the formation of
an expanded and mtensrhed oxygen-minimum zone m the water column, and the wdesprcad
deposition of radiolarian-bearing organic-rich muds in deeper-water areas. In shelf settmgs
productivity was lower and marly nannofossil oozes (Plenus Marl facies) were deposited. At the
height of the upwelling event, the oxygen-depleted (dysaerobic) upper portion of the oxygen-
minimum zone impinged onto the Chalk Shelf, causing a widespread decline in benthonic diversity.
A fall in the rate of oceanic turnover at the end of the Cenomanian produced a waning of the
productivity event, and during the early Turonian there was a gradual return to a mid-Cenomanian-
like palaeoceanographic regime.
processes.) Neither the presence of bioturbation, nor the absence of organic-
rich sediments prove that an oxygen-minimum zone was not developed in
the water column. On the contrary, we will argue that the decline of
microfossil abundances and diversities in close association with rising 13C
values in the Plenus Marl (see Section 11.1) strongly suggests that bottom
waters became significantly depleted in oxygen (cf. Hart and Bigg, 1981;
Hart, 1985; Hart and Ball, 1986), even if not truly anoxic (Section 11.2).
11. Effects of the OAE on the Chalk Sea biota (I. J. et al.)
I I I I
I 1
I ’
1 ,
6 13c
..,.,. .,.,.:
::::::..I:
....
,.,.
:.,.:.
.,. . ‘. .:.
., ,. .,
i; .,::,j.: Melbourn Rock
:..:.......
Beds
: ,: .::.:.:.:.:,
:..::li”iiijliliiiij’& ___
-k
,,.:: :. ../._
:.S.”
.::. .‘*.x__._
%, (PDBl ,, (-jstracods ?._-,._
‘jll
2 4 5
I
0
I
10
1
20
I
0 10
I 1
20
Number of I Number of
Melbourn Rock
602’0d 10
Planktonic diversity
Microfossil Assemblages and the Oceanic Anoxic Event 87
Figure 3 1. Carbon stable-isotope stratigraphy and microfossil species diversity within the Cenomanian-
Turonian boundary succession at Dover. Discontinuous horizontal lines are lithological boundaries
(base of the Plenus Marl, base of the Melbourn Rock, top of the Melbourn Rock), the continuous
horizontal line marks the base of the Turonian. Note the decline in diversity of all groups evithin the
Plenus Marl in close association with rising 6’% values. Recoveries in diversity within the lower
Turonian arc’ more rapid in planktonic than benthonic groups.
I I 1
t Number of
613c specimens
,:::::::. :::.
.::,:: ::,,:::,
:. .A..~.~.~.
:. ::. :::. ::
::
:+,.,T...r..._ __ - - - - - - - - - __-_-______
,.,. .. ..
Melbourn Rock
Beds
I,....
:...
::.
,.:.:.
.F...
TUWllW
-------.Cenomanian
%o (PDBI .:YOstracods
I
I I
2- 4 5 - 400 0 500 1000
Figure 32. Carbon stable-isotope stratigraphy and microfossil abundance within the Cenomanian-Turonian boundary succession at Dover. Symbols its in Figure 31.
Microfossil abundances are lowest immediately following the peak of the carbon stable-isotope excursion. Abundances and diversities are other\vise not clearI)
interrelated (compare Figure 31).
Microfossil Assemblages and the Oceanic ilnoxic Event 89
sea level
thin-walled, variable
trochospire, small
number randomly
placed pustules
\
periphery
\ ; \ / 4
some
I forms evolve \ thick-walled, biconvex
I with weakly to plano-convex,
I developed .I supplementary apertures,
two keels I thick well-developed
, ‘early dicarinellids’ 1 keels which thin
towards last-formed
\ / chambers
\ /
. -_ /
depth
Figure 33. Relatiohships between test morphology and water depths inhabited by Cenomanian-
Turonian planktonic foraminifera. Migration within the water column during ontogeny is shown
schematically. Juveniles of all groups inhabited shallow water. Individuals migrated into deeper
water during ontogeny, reproducing only at the lower limit of their depth range. Deeper water forms
were, therefore, more susceptible to oxygen-depletion within deep-water masses.
92 I. Jar\ is rr rd.
Sea level
Sea level
J-Jr
‘. ” :, .’ ‘.”
_-__-__-_---------------------
____-_____---- ____~__----------
_______--------------
reached its shallowest depth at the peak of the isotope excursion (Figure
34D), immediately before the start of the Turonian, during the deposition of
Plenus Marl Beds 6-8 and the basal metre of the Melbourn Rock. Calcareous
microfossil tests display evidence of seafloor dissolution within this interval,
a feature which is also characteristic of tests collected from modern oxygen-
minimum zones (Penrose and Kennett, 1979). The occurrence of only
‘shallow-water’ planktonic foraminifera at this time suggests that the
oxygen-minimum zone may have risen to depths as shallow as 50 m (cf. Bk,
1977). The recovery of microfossil diversity associated with the stage
boundary indicates the waning and contraction of the oxygen-minimum
zone during the earliest Turonian.
this time most typical Cenomanian species had disappeared, and the
microbenthos was of low diversity, restricted to a few species of foraminifera
(Gavelinella berthelini, Lingulogavelinella globosa, Textularia chapmani,
smooth Lenticulina spp.) and platycopid ostracods (mainly Cytherella ovata).
Oxygen-depleted waters had now risen to their shallowest depth (possibly to
within 50m of the surface), leaving only reduced populations of shallow-
water hedbergellid planktonic foraminifera. All calcareous microfossils
display surface etching of their tests in these beds, indicating seafloor
dissolution of carbonate, comparable to that seen in modern oxygen-
minimum zones. Dinoflagellate cysts also display a major decline in diversity
at this time, and disappear entirely immediately following the deposition of
the Plenus Marl. Their decline is initially accompanied by a sudden
abundance of Kallosphaeridium ringnesiorum, and this is followed by a
proliferation of calcispheres, which constitute the main component of the
sediment at the height of the OAE. These changes may relate to the
establishment of abnormally stable hydrodynamic conditions at this time.
Oxygenation levels remained low during the deposition of basal beds of the
Melbourn Rock which contain faunas that are as impoverished as those in the
uppermost Plenus Marl. A decline in calcareous nannofossil diversity at the
end of the Cenomanian, however, may indicate falling nutrient levels
associated with the waning of the OAE. An initial phase of better oxygenation
immediately preceding the start of the Turonian (marked by the appearance
of Mytiloides spp. inoceramid bivalves), is evidenced by increases in both
benthonic and planktonic microfossil diversity.
The base of the Turonian is defined by the appearance of new species of
inoceramid bivalves and ammonites which were associated with increasing
oxygen levels in the Chalk Sea. New species appeared in all groups during the
early Turonian, as an increased number of habitats became reinhabitable,
and evolution filled the niches which had been left vacant by the late
CenoTanian extinctions. Appearances included new species of Areno-
bulimirza, Lingulogavelinella, and Tritaxia in the benthonic foraminifera,
and new species of podocopid ostracods (e.g. C)lthereis sp. cf. C. longaeeja,
Mosaeleberis sp. A, Curfsina senior). Planktonic foraminifera evolved new
intermediateiwater (Helvetoglobotruncana) and then deep-water (Margino-
truncana) genera, and several new species of calcareous nannofossil and
dinoflagellate cyst eventually appeared (including the biostratigraphically
important forms Q.uadrum gartneri and Senoniasphaera rotundata). Plank-
tonic groups generally recovered more rapidly than benthonic taxa, although
dinoflagellate cysts remained absent for a considerable interval of time.
Diversities never regained Cenomanian levels during the early Turonian,
despite evidence of continued increases in regional water depth. This
demonstrates the lasting impact of the OAE on the marine ecosystem.
We believe, therefore, that the Cenomanian-Turonian OAE caused a
sequence of extinctions during the latest Cenomanian. The recovery of the
biota following the event led to the evolution of the new species which define
the Turonian, enabling it to be distinguished from the under1yir.g
Cenomanian. Oceanic anoxic events provide, therefore, a mechanism ior
extinction and evolution in the marine realm which was almost certainly
operative throughout the Phanerozoic.
Acknowledgments
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