-
23. APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS FROM THE MAZAGAN
PLATEAU,NORTHWEST AFRICA (SITES 545 AND 547, DEEP SEA DRILLING
PROJECT LEG 79)1
Raimond Below, Institut für Palàontologie, Universitàt Bonn2
ABSTRACT
Eighty-eight samples of Aptian to lower Cenomanian sediments of
Sites 545 and 547, DSDP Leg 79, from the Ma-zagan Plateau area
(offshore Northwest Africa) were analyzed for palynomorphs. The
very rich dinoflagellate cyst as-semblages make it possible to
narrow shipboard age determinations and to correlate Sites 545 and
547. The distributionof 174 dinoflagellate cyst taxa is tabulated
in this study and the biostratigraphic value of selected
dinoflagellate cysts isdiscussed. Additional taxonomic remarks are
made about some species. The new dinoflagellate cyst species Aptea
al-mohadensis, Occisucysta hinzü, O. mazaganensis, and the
subspecies Maghrebinia perforata (Clarke and Verdier, 1967)Below,
1981 ssp. mirabilis are described.
INTRODUCTION
Sites 544 to 547 were drilled in the Mazagan Plateauarea of the
Moroccan Atlantic continental slope duringDSDP Leg 79. Of these,
Sites 545 and 547 were chosenfor biostratigraphic analysis of
mid-Cretaceous dinofla-gellate cyst assemblages.
Site 545, at the northwestern foot of the MazaganPlateau slope,
consists of 701 m of Jurassic to Quater-nary sediment. For this
palynological study, 278 m ofgreen, carbonaceous claystone with a
few meters of do-lomitized chalk at the base are of interest. The
age wasdetermined by shipboard scientists as early late Aptianto
middle or late Cenomanian. These strata rest uncon-formably on
Jurassic carbonates and are separated by afurther unconformity from
the overlying Neogene chalks.
Site 547, 14 km north-northeast of Site 545, is situ-ated on the
northeastern slope of a basement horst.Holes 547A and 547B
penetrate a total of 1030 m of Ju-rassic to Quaternary sediment.
The samples for thisstudy were taken from a 351-m-thick section of
Albianto Cenomanian carbonaceous clay- and mudstones. Thisinterval
is separated from the underlying Jurassic lime-stones by a major
hiatus, and from overlying debrisflows, reworked during the Late
Cretaceous and Paleo-cene, by an unconformity.
A total of 88 samples were prepared using commonpalynological
techniques. In addition to various palyno-morphs such as pollen and
spores, acritarchs, scoleco-donts, phycomycetes,
"micro-foraminifers", wood frag-ments, and cuticula, all samples
yielded rich dinoflagel-late cyst assemblages, with many
individuals and oftenvery high species diversity. Preservation of
the cysts isvery good, except for a few reworked specimens.
Analysis of these dinoflagellate cyst assemblages hasmade
possible the detailed determination of sedimentages, subdivision of
the mid-Cretaceous sequence, and
Hinz, K., Winterer, E. L., et al., Init. Repts. DSDP, 79:
Washington (U.S. Govt. Print-ing Office).
2 Address: Institut für Palàontologie, Rheinische
Friedrich-Wilhelms Universitàt Bonn,Nussallee 8, D-53OO Bonn,
Federal Republic of Germany.
correlation of Site 547 with Site 545. The results will
bediscussed in the biostratigraphic section of this chapter.Another
section deals with dinoflagellate cyst taxono-my; it includes short
remarks about selected species anddescriptions of four new
taxa.
BIOSTRATIGRAPHY
Because reliable, detailed, and universally valid
dino-flagellate cyst zones for the mid-Cretaceous Period arenot yet
formalized, age determination is based on com-parison with
dinoflagellate cyst assemblages known fromthe neighboring,
stratigraphically well-dated sections ofSite 370 (Williams, 1978)
and Moroccan onshore sec-tions (Below, 1981; Below, 1982a,
1982b).
Site 545 (33°39.86'N; 9°21.88'W;water depth 3142 m)
A total of 278 m of mid-Cretaceous sediment wasdrilled,
extending from Core 56 to Core 27. The basal 10m consist of yellow
grey dolomitic chalks resting imme-diately upon the
Jurassic/mid-Cretaceous unconformi-ty. These strata were dated as
early late Aptian by ship-board paleontologists (shore-based
study). The seriescontinues with a monotonous interval of
carbonaceousclaystones with a few layers of clay pebbles in Cores
40,35, and 34. Within this sequence, the following ship-board age
determinations were proposed, using fora-minifers and nannofossils:
Core 56, early Aptian (?lateBarremian); Cores 55 to 44, late
Aptian; Cores 44 to 37,late Albian, with problematic existence of
early Albianequivalents; Cores 36 to 34, early(?) Cenomanian;
Cores34 to 33, middle(?) Cenomanian; Cores 33 to 27, late
Ce-nomanian. These ages have subsequently been furtherrefined with
shore-based study (see chapters by Leckieand Wiegand, this volume).
The mid-Cretaceous/Neo-gene unconformity is located within Core
27.
AnalysisFrom Hole 545, 39 samples were collected: 3 in 11 m
of the ?Barremian-Aptian sequence; 18 in 110 m of up-per Aptian
sediments; 12 in 70 m of the upper Albianinterval, and 6 in 87 m of
Cenomanian age. The distri-
621
-
R. BELOW
Ag
eC
eno
man
ian
bas
al
! ea
rly
Alb
ian
late
itian
late
Ap
mid
dle
?ear
ly
Pal
In
st C
olle
ctio
n N
o
691 9
6920
6 92 1
6922
6923
6 92 4
6925
6926
6927
6928
6 929
6930
6931
6932
6933
6934
6935
6936
6937
6938
6939
6940
694 1
6942
694 3
6944
6945
6946
694 7
6948
6949
6950
6951
6952
6953
6954
6955
6956
6957
Cor
e
Sec
tion
Inte
rval
(
in
cm )
31
32
33
34
35
36
,?738
39
39
4040
4 1
4142
43
43
44
4545
4646
474 7
48
48
49
4950
50
57
5152
53
54
55
55
56
56
7
7
7
3
7
7
I7
7
4
2
6
7
4
5
7
3
2
7
4
2
4
2
6
2
6
7
5
J_67
6
2
3
3
7
6
3
6
130 - 133
029 - 033
090 - 094044 - 046
097 - 099
044 - 047
084 - 087120 - 123
093 - 096
094 - 097
116- 119057 - 06004 7 - 04 9
000 - 004016 - 019
013 - 016
017 - 020
078 - 082
059 - 062040 - 043
063 - 066
063 - 066
097 - 0990 7 2 - 0 7 5
7 7 2 - 1 1 4
016 - 01 8
03 4 - 03 7
049- 052147 - 150
056 - 058
047-- 050
066 - 069
123 - 126090 - 094
043 - 047
060 - 062071 - 073058 - 060055 - 057
Olig
osp
ha
eri
diu
m
totu
m m
ino
r 1
1
Sp
inife
rite
s ra
mo
sus
ram
osu
s 2
T
-
-
-
-
t1
•
Olig
osp
haer
idiu
m
com
ple
x 3
1
iITi
Tan
yosp
hae
rid
ium
b
ole
tum
4
i
r
II *
-
-
Cri
bro
pe
rid
iniu
m c
f. d
iap
hane
5
Sp
inife
rite
s m
uit
ibre
vis
mu
itib
revi
s 6
Occ
isuc
ysta
te
nulc
eras
7
•
Mud
ero
ngia
pe
rfo
rata
8
•
••
••
Ce
rbia
ta
bu
lata
9
Su
btil
isp
ha
era
sen
egal
ensi
s 1
0D
issi
liod
iniu
m g
lob
ulum
1
1
*
1
•
•
11
1
1
1
I
T
1
Cri
bro
pe
rid
iniu
m
ort
hoce
ras
12
|1
1
11
•
T
J
DD0D)3D
E o
II
Ifi>
-
APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS, MAZAGAN PLATEAU
Ap
tea
p
oly
mo
rph
a
31
•
I1
L
11
Kle
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ria
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ae
rid
ium
e
oin
od
es
32
•
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•
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ho
mo
sph
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ra
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ieri
3
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me
tod
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whi
te/
35
1
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iou
roq
on
yau
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sto
veri
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0
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cyst
a so
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s
•
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•
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ocα
1iòαoE
T3O
c
•
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•
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lysp
ha
eri
diu
m7 f
lexu
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m
39
1
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1
1
1
Can
nin
gia
c
f re
tic
ula
ta
4 0
•
••
•
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•
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α
IO
α
oco
•
•
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11
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•
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3
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o
D l
O
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Ca
nn
ing
io
rin
gn
esi
oru
m
4 4
|
Olig
osp
ha
eri
diu
m
totu
m
totu
m
' 5
J
•
•
11
1
1
•
•
Pte
rod
iniu
m
co
rnu
tum
4
6 j
T
+••
Olig
osp
ha
eri
diu
m
po
cu
lum
4
7 j
I
•
••••
T
•
•
Ap
teo
din
ium
m
ac
ula
tum
4
8
T
1
1
•
i
1
• |
|
Cle
/sto
spha
erid
ium
da
vulu
m
4 9
I
011̂o
^5DiΦ
ε3
•Q
OiD
αIOo
D
T
1j1
| 1
Co
ron
ife
ra
oc
ea
nic
a
5 /
]
•
••
••
r
I
|
ε3O
.w
"5E3
O
•
—-p̂
F
lore
nti
nia
m
an
telli
i 5
31T1.
i
I
Od
on
toc
hiti
na
o
pe
rcu
lata
5
4 \
Sp
inif
eri
tes
an
co
rife
rus
a
nc
ori
feru
s 5
5 |
I • 1
S
pin
ife
rite
s le
nzi
i 5
6 |
1•11
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I
1I
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* II
111T
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"3εw
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ch
om
osp
ha
era
n
ep
tun
i 5
8 \
j_
1 C
rib
rop
eri
din
ium
cf
ort
ho
ce
ras
59
|
1iI1I
to
u3D
O-Q
ΦO
I1I
I
11
1
Sys
tem
ato
ph
ora
si
lyb
a
-
R. BELOW
Olig
osp
hae
rid
ium
ve
rru
cosu
m
80
Cπ
bro
din
ium
e
dw
ard
sii
81
•
f-
Sp
inif
eri
tes
ram
osu
s re
tic
ula
tus
82
•••
Ca
ssic
ulo
sph
ae
rid
ia r
eti
cu
lata
8
3
••
••
1
Olig
osp
ha
eri
diu
m
div
idu
um
8
4
>
>
•
•
•
Pro
toe
llip
so
idin
ium
se
ghi
re
seg
hire
8
5
1
•
•1
00
ε3
COo
ε3"50)0
αo
o
c
0
in
uo
α>•QO
•
11
|
Ca
rpo
din
ium
gra
nu
latu
m
9 8
\
•
•
Occ
isu
cyst
a h
inzi
i 9
9 \
Ba
tia
ca
sph
ae
ra
said
en
sis
1 0
0
•
•
Pse
ud
oc
era
tiu
m g
oc
hti
i 1
01
\
•
Su
btil
isp
ha
era
d
efo
rma
ns
1 0
2
•
1
oc~>3
3n3
5
3
3
>
3
5J
1
i
| H
ystr
ich
od
iniu
m
pu
lch
rum
1
04
\
•
•
•
•
•
•
1
—"|
P
ala
eo
pe
rid
iniu
m
cre
tac
eu
m
10
5
|
Ch
lam
ydo
ph
ore
lla
nye
i 1
06
| E
nd
osc
rmiu
m c
am
pa
nu
la
nic
han
10
7
I
111
Tj _
1
\ P
tero
din
ium
m
am
ou
nia
1
08
Cri
bro
pe
rid
iniu
m
dia
ph
an
e
10
9P
tero
din
ium
b
ab
1
1 0
Co
do
nie
lla
psy
gm
a
1 1
1
•
Olig
osp
ha
eri
diu
m
alb
ert
en
se
11
2
-
Hys
tric
ho
din
ium
co
mp
act
um
1
13
E3o
L.ΦQ_
ε3
O13
δin
g
c
1
u->
α>
ε3coOicQ
•
Po
lysp
ha
eri
diu
m
sp.
sens
u B
elo
w
1 1
6J
Sys
tem
ato
ph
ora
cr
eta
cea
1
1 7
1
00
oco3Cn3
-C
|
Ojo-c
o
o
•
•1S
pin
ife
rite
s c
orn
utu
s c
orn
utu
s 1
1 9
•
•
His
tio
cys
ta
ou
tan
ane
nsi
s 1
20
•
•
Co
do
nie
lla
ca
mp
an
ula
1
21
•
II
•I
Hys
tric
ho
sph
ae
rin
a s
chin
de
wo
lfii
122
I
Occ
isu
cyst
a
cf.
te
nu
ice
ras
123
Olig
osp
ha
eri
diu
m
po
rosu
m
12
4
••
Sys
tem
ato
ph
ora
c
om
plic
ata
1
25
1
Ne
mat
osp
hae
rop
sis?
sin
gu
lari
s 12
6
•
Wa
llod
iniu
m
lun
a
12
7 CO
ε3cΦ
Eoin
cv
3
13
ΦO
'αinO
- c
ò>5C
•
Pro
toel
lipso
dini
um s
pino
sum
12
9
I
Pro
toelli
pso
idin
ium
to
uile
to
uile
13
0
|•
Figure 1. (Continued).
for example, Site 547 (this chapter), Site 370 (Williams,1978),
and onshore sections (Below, 1981; Below, 1982a,1982b). Only
selected samples were examined, although,in contrast to various
published studies of dinoflagellatecyst assemblages, they were
taken at very short intervals.Nevertheless, it is probable that
many occurrences of"short-ranging" species in strata between sample
pointswere by chance not found. Only continuous trench sam-pling
would document all faunal horizons.
With this in mind, this chapter discusses only selectedcysts,
not all 160 taxa occurring at Site 545. For more de-tailed
information about other species, the reader is re-ferred to the
dinoflagellate cyst distribution chart, Fig-ures 1 and 2.
The few individuals from a poor dinoflagellate cystassemblage in
Sample 545-56-6, 55-57 cm give little evi-dence for age
determination. The four species, Oligo-sphaeridium complex, O.
totum ssp. minor, Spiniferites
624
-
APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS, MAZAGAN PLATEAU
Eyr
ea
ne
bu
losa
13
1 \
|
•
Su
btil
isp
ha
era
che
it 1
32 \
1
•I
Olig
osp
haeridiu
m
pulc
herr
imum
7
33 |
•
Lito
sphaeridiu
m
coni
spin
um
7 34
|
1i••
Chi
amyd
opho
reila
d
iscr
eta
7
35
1
•
Litosphaeridiu
m s
ipho
niph
orum
1
36 \
IT
•
Dl
C
•
Hys
tric
ho
sph
ae
rid
ium
a
run
du
m
1 38
\K
ioka
nsiu
m c
oro
llum
1
39 \
Maqhre
bin
ia p
erf
ora
ta m
ira
bili
s 14
0\
1| |
Oo
dn
ad
att
ia
tub
erc
ula
ta
141
1
•1
Cyc
lonepheliu
m c
f m
embr
anip
horu
m
7 42
|
J_
Occ
isuc
ysta
m
aza
ganensi
s 14
3 |
Pala
eohys
tric
hophora
inf
usor
ioid
es
14 4
I
Xen
ascu
s p
lote
i 1
45 j
•1
Flo
ren
tinia
res
ex
146
•I
Ma
gh
reb
inia
ch
teuh
1
4 7
I
Cyc
lonepheliu
m
vann
opho
rum
7
48
|
•
Co
nya
ula
cyst
a
cass
idata
1 4
9\
Ach
omos
phae
ra r
am
ulif
era
7
50
|X
enas
cus
cera
tioid
es
151
]
•
•
•O
odnadattia
al
ata
1 52
|
1I
IC
yclo
nepheliu
m
chab
aca
7 53
|F
lore
ntin
ia
ste
llata
15
4
Ii
Ma
gh
reb
inia
pe
rfo
rata
perf
ora
ta
1 55
]
|
Ovo
idin
ium
ve
rruco
sum
7
56 |
Iii
Ach
omos
phae
ra
sage
na
7 5
7J
Ep
eiid
osp
ha
erid
ia
spin
osa
7 5
8 |
1
7 - 3 specimens4-20
21 -5051-100> 700
Flo
rentm
ia
laci
nia
ta la
cin
iata
7
59
|
1
Hys
tric
hosp
haeridiu
m
bow
erb
anki
i 1
60 |
I
1i
Cor
e
Se
ctio
n
Inte
rval
(in
cm
)
31
32
33
34
35
36
37
38
3939404041
4142
434344
454546464747
4848494950
50575752
53
54
55555656
7
7
7
3
7
7
2
7
7
4
2
6
1
4
5
7
3
2
7
4
2
4
2
6
2
6
1
5
7
6
7
6
2
3
3
7
6
3
6
7 3 0 - 7 3 3
029 - 033090 - 094044 - 0 4 6
097 - 099
044 - 047
084 - 087120 • 123093 - 096094 - 097
7 76 - 7 79
057 - 060047 - 049
000 - 004016 - 019
013 - 01601 7 - 02 0078 - 082
059 - 062040 - 043063 - 066063 - 066097 - 0990 72 - 07 5
112 - 11401 6 - 07 8034 - 03 7049 - 05274 7 - 750
056 - 058047 - 050
066 - 069123 - 126090 - 094
043 - 047060 - 062071 - 073058 - 060055 - 057
Pal
. In
st.
Col
lect
ion
No.
69196920692169226923692469256926692769286929693069316932693369346935693669376938693969406
94 169426 94 36944694
5694669476948694969506951695269536954695569566957
Age
Cen
oman
ian
basa
l (e
arly
Alb
ian
late
2a
c
Q.
mid
dle
?ear
ly
Figure 1. (Continued).
ramosus ssp. ramosus, and Tanyosphaeridium boletum,are all
long-ranging taxa, known from Lower and UpperCretaceous sediments.
This sample and Sample 545-56-3,58-60 cm were taken from a sequence
of crushed andfaulted green claystone, dated as (?Barremian) Aptian
bythe shipboard party (subsequently dated as early lateAptian by
foraminifers; Leckie, this volume). Becauseof the stratigraphically
insignificant dinoflagellate cystassociation this age determination
can neither be veri-
fied nor rejected for Sample 545-56-6, 55-57 cm. How-ever, an
Aptian—even an early Aptian age—can be pro-posed for Sample
545-56-3, 58-60 cm, because charac-teristic dinoflagellate cyst
populations found in middleAptian sediments of Moroccan onshore
sections do notoccur below Core 53. The population of Sample
545-56-3, 58-60 cm comprises 21 species, of which three are
ofstratigraphic interest: Aptea eisenackii (peak), Occisu-cysta
tenuiceras, and Cerbia tabulata. Both of the form-
625
-
Figure 2. Stratigraphic ranges of selected dinoflagellate cysts
in Hole 545, between Cores 30 and 56. X = dinoflagellate cyst taxa
also present in the Albian (Alb) and/or Cenomanian (C)of Holes 547A
and 547B.
-
APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS, MAZAGAN PLATEAU
er are limited to the Aptian of the onshore Ida ouTanane section
(Below, 1981), whereas Cerbia tabulatashows equal frequency in the
Barremian and Aptian ofthis onshore locality. Species with base
ranges in thissample and maxima during the Aptian are
Cribroperidi-nium orthoceras, Cyclonephelium maugaad, and
Kiokan-sium hydra. The base range of Florentinia
radiculatacorresponds to its first occurrence in the Aptian of
Site370 (Williams, 1978).
Sample 545-55-6, 71-73 cm yielded a very rich popu-lation, both
in species diversity and number of individ-uals. Of 48
dinoflagellate species, 38 have their site-spe-cific base ranges in
this sample. Sample 545-55-6, 71-73cm contains dinoflagellate cysts
that are regarded as typ-ical of Aptian and/or younger sediments,
such as Acho-mosphaera verdieri, Aptea eisenackii, A. polymorpha,C.
maugaad, F. radiculata, O. sousensis, O. tenuiceras,and
Prolixosphaeridium parvispinum. Most of thesespecies are very
abundant in this sample and thus clearlycorrespond to their known
peak in the Aptian. Williams(1978) has demonstrated that the
characteristic P. par-vispinum (as P. granulosum) has its base
range withinAptian sediments at Site 370. Cribroperidinium
ortho-ceras, K. hydra, and Rhynchodiniopsis aptiana, whichalso
occur in Sample 545-55-6, 71-73 cm, are typical ofboth Barremian
and Aptian ages.
Danea chibanis, in Sample 545-54-3, 43-47 cm, is wellknown in
the middle Aptian of Moroccan onshore sec-tions (Below, 1981).
Protoellipsodinium touile ssp. touilein Sample 545-53-3, 90-94 cm
and P. seghire ssp. seghirein Sample 545-52-2, 123-126 cm
strengthen the impres-sion that these are all middle Aptian
dinoflagellate cystpopulations. This last sample also yields the
first ap-pearance of the morphologically very characteristic
andshort-ranging Oligosphaeridium verrucosum.
Only a few species are introduced as new componentsin the
dinoflagellate cyst associations in the interval be-tween Samples
545-51-6, 66-69 cm and 545-50-1, 147-150 cm. Of these, Hapsocysta
peridyctia and Systema-tophora sp. MAZ may in the future prove to
be of strati-graphic value for middle Aptian time. In this
sampleinterval, low species diversity is correlated with the
ab-sence of dominant species with high numbers of speci-mens.
This is obviously different in Sample 545-49-5, 49-52cm, which
shows high species diversity and great num-bers of dinoflagellate
cyst individuals. Eleven species oc-cur in this sample for the
first time. Of these, Aptea al-mohadensis, Chlamydophorella nyei,
Coronifera tubu-losa, and Subtilisphaera deformans immediately
becomedominant species, but only for a short period. It is inthis
sample that Cassiculosphaeridia tazadensis has itsgreatest
frequency; this is a significant dinoflagellatecyst from the middle
Aptian in the Ida ou Tanane on-shore section (Below, 1981).
Morphologically character-istic species with base ranges are
Carpodinium granula-tum and Occisucysta hinzü.
Many species are new in the dinoflagellate cyst as-semblages
between Samples 545-48-6, 16-18 cm and 545-44-2, 78-82 cm. Some of
these are unknown in the Mo-roccan lower and middle Aptian:
Codoniella campanu-
la, C. psygma, Kleithriasphaeridium? sarmentum,
Ne-matosphaeropsis? singularis, and O. cf. tenuiceras. Sincethey
are associated with dinoflagellate cyst populationsin which typical
Aptian species dominate, it is possiblethat they may indicate a
late Aptian age for this strati-graphic interval. This is supported
by the many Hystri-chosphaerina schindewolfii specimens in Sample
545-46-4, 63-66 cm. Their presence makes it possible to connectthis
sample with the Systematophora schindewolfii Zoneof the late Aptian
(Williams, 1975, 1978). Other species,such as Ellipsoidictyum
imperfectum, Oligosphaeri-dium indicum, and S. cretacea, which have
base rangesin the sequence between Samples 545-47-6, 72-75 cmand
545-46-2, 63-66 cm, do not occur in the Moroccanmaterial before the
Albian. Some species typical for theBarremian-Aptian have their top
ranges within the in-terval between Samples 545-46-2, 63-66 cm and
545-41-4, 0-4 cm: Cassiculosphaeridia tazadensis, Cerbia auc-da, C.
tabulata, Occisucysta tentoria, and Spiniferitesmagnoserratus.
Species of the genus Cerbia seem to begood guide fossils, for they
never extend above the Apti-an/ Albian boundary (Williams, pers.
comm., 1981). Nev-ertheless, stratigraphically reliable species
indicate thatthe Aptian extends as high as Sample 545-40-6,
57-60cm. Species having their top range within this sampleand the
somewhat older Sample 545-41-1, 47-49 cm areCassiculosphaeridia
reticulata, Cribroperidinium ortho-ceras, Cyclonephelium maugaad,
K. eoinodes, Meiouro-gonyaulax stoveri, O. sousensis,
Protoellipsodinium seg-hire ssp. seghire, Pterodinium aliferum, and
Rhyncho-diniopsis aptiana.
Several thin beds of claystone conglomerates havebeen recorded
within Core 40. Shipboard scientists pro-posed a very minor hiatus.
Changes in the compositionof dinoflagellate cyst populations
demonstrate it to be amajor hiatus. Sample 545-40-6, 57-60 cm, 80
cm belowthe first pebble conglomerate, is of Aptian age,
whereasSample 545-40-2, 116-119 cm yields dinoflagellate
cyststypical for the very late Albian. Most species which havebase
ranges from Sample 545-40-2, 116-119 cm to Sam-ple 545-31-1,
130-133 cm characterize the Albian/Ceno-manian boundary. No
dinoflagellates significant for thelower or middle Albian from
onshore sections were rec-ognized. Cyclonephelium chabaca,
Litosphaeridium si-phoniphorum, Maghrebinia chleuh, M. perforata
ssp.mirabilis, M. perforata ssp. perforata, and
Palaeohystri-chophora infusorioides occur within the Ida ou
Tananeonshore section in the highest Vraconian and are knownto
range into the Cenomanian (Below, 1981).
There are problems in determining the exact positionof the
Albian/Cenomanian boundary at Site 545 (see al-so discussion of
Site 547, later). Sample 545-39-1, 93-96cm contains the species P.
infusorioides, the base rangeof which, according to Williams
(1978), marks the be-ginning of the Cenomanian at Site 370. This is
in agree-ment with the work of Clarke and Verdier (1967),
whoerected the Hystrichosphaeridium siphoniphorum Zoneof Cenomanian
to ?Turonian age with three subzones.The base of this zone and its
basal subzone, respectively,are marked by the first appearance of
P. infusorioides.Palynologically, this can be taken as the base of
the Ce-
627
-
R. BELOW
nomanian (Clarke and Verdier, 1967). Nevertheless, Flo-rentinia
berran, F. resex, Maghrebinia chleuh, M. perfo-rata ssp. perforata,
and Oodnadattia alata, well knownin upper Albian onshore sections
(Below, 1981; Below,1982b), occur within Site 545 with base ranges
that areyounger than the first appearance of P. infusorioides.
Acomparison of dinoflagellate cyst assemblages from well-dated
onshore sections with those from Site 545 seemsto indicate that
Xenascus ceratioides is, at least region-ally, a more reliable
marker for the beginning of the Ce-nomanian; therefore, it is used
here to establish the Albi-an/Cenomanian boundary between Samples
545-38-1,120-123 cm and 545-37-2, 84-87 cm.
According to Williams (1978), the abundant Epeli-dosphaeridia
spinosa in Sample 545-30-1, 130-133 cmdemonstrate that this stratum
is not younger than earli-est Cenomanian. The sequence between the
first occur-rence of E. spinosa and Sample 545-37-2, 84-87 cm,
whichcontains the base range of X. ceratioides, would, there-fore,
be of basal Cenomanian age (Williams's term).This conclusion would
be supported by the presence ofKiokansium corollum, whose range,
according to Wil-liams (1978, as Cleistosphaendium polypes ssp.
A),should not extend over the basal/early Cenomanianboundary.
Site 547 (Holes 547A and 547B) (33°46.84'N;09°20.98'W; water
depth 3938 m)
A total of 351 m of mid-Cretaceous sediment werecored in Hole
547A between Cores 39 and 73 and in thereentry Hole 547B between
Cores 1 and 7.
The sequence between Cores 547B-6 and 547A-39 con-sists of
grayish green claystone and mudstone with com-mon, intraformational
flat-pebble mudstone conglomer-ate layers. It was dated by
shipboard paleontologists asranging from the late Albian to middle
Cenomanian. Itrests with a hiatus on Upper Jurassic to Lower
Creta-ceous nodular limestone and terminal claystone. Thetop of the
mid-Cretaceous section is marked by an un-conformity upon which
Upper Cretaceous and LowerTertiary strata were recorded.
Within the sequence studied here, age determinationsproposed in
the shipboard report using nannofossils were:547A-39-1, 140 cm to
547A-40,CC, middle and late Al-bian to middle Turonian; 545A-41,CC
to 545A-62,CC,early Cenomanian to early Turonian; 547A-63,CC
through547B-6-1, 107 cm, late Albian. Ages using foraminiferswere:
Cores 547A-39 to 547A-46, late Cenomanian;547A-47 to 547A-62, early
Cenomanian; 547A-63through 547B6-1, 107 cm, late Albian.
Redeposition with-in the section masks clearly defined
boundaries.
Analysis, Hole 547B
Thirteen core samples were collected from a 48-m up-per Albian
interval in Hole 547B (with high sample den-sity, e.g., 10 samples
from 9.5 m of Core 5). Ninety taxawere recorded in Hole 547B; their
distribution is shownin Figures 3 and 5.
All 10 samples of Core 547B-5 yielded rich dinofla-gellate cyst
populations. Dominant species are Cyclone-phelium brevispinatum,
Litosphaeridium conispinum,Odontochitina operculata,
Oligosphaeridium complex,
Ag
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A
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6874
6875
6876
6877
6878
6879
6880
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APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS, MAZAGAN PLATEAU
Pervosphaeridiwn truncatum, and Spiniferites multi-brevis ssp.
seghiris. Important components are Chlamy-dophorella nyei,
Coronifera oceanica, Exochosphaeri-dium phragmites, Florentinia
mantellii, Gonyaulacystahelicoidea, Kiokansium corollum,
Maghrebinia perfora-ta ssp. mirabilis, O. irregulare, P.
pseudohystrichodinium,S. ancoriferus ssp. ancoriferus, and S. cf.
wetzelii. Incomparison with dinoflagellate cysts from
biostratigraph-ically well dated onshore sections, the association
L.siphoniphorum, M. chleuh, M. perforata ssp. mirabilis,and M.
perforata ssp. perforata characterize earliest Ce-nomanian and/or
late Albian. This is supported by thepresence of F berran and F.
resex in the somewhatyounger Core 547B-2. However, the late Albian
age ismore probable and is proposed here, because
Xenascusceratioides, with a base range at the beginning of
theCenomanian (see later discussion) is absent in Hole547B. It
appears for the first time in Core 547A-69.
It is difficult to correlate Core 547B-5 with the se-quence of
Hole 547A. Theoretically, such correlationshould be easy by
comparing the distinctive base rangesof the species named earlier
in Hole 545 with their baseranges in the continuous strata of Hole
547A, which areclear. However, these species all make their first
appear-ance within the deepest sample in Hole 547B, Sample547B-5-7,
40-43 cm. This first record does not of courseimply that the sample
is their biostratigraphic base range.
Some characteristic Aptian taxa are recorded in Hole547B
samples: Callaiosphaeridium trycherium, Cassicu-losphaeridia
reticulata, Muderongia perforata, Occisu-cysta sousensis, and
Systematophora silyba. These are
almost all single specimens, a fact that indicates rework-ing of
older sediments.
Analysis, Hole 547A
Eleven core samples were collected from an 85-m up-per Albian
interval in Hole 547A, and 25 samples froma 236-m Cenomanian
section. An additional sample,547A-21-4, 110-113 cm, from a
conglomerate layer withinEocene sediment, was analyzed for
palynomorphs. InHole 547A 91 taxa were recorded; their distribution
isshown in Figures 4 and 5.
Cores 547A-72 and 547A-73 overlap with Cores 547B-1to 547B-3,
and the dominant species in the dinoflagel-late cyst assemblages of
both core intervals correspond.The dinoflagellate cyst populations
of nearly all sam-ples collected from Hole 547A are characteristic
of theAlbian/Cenomanian boundary section (see earlier dis-cussion).
If one assumes that the base range of Xenas-cus ceratioides begins
with the Cenomanian, then theinterval from Samples 547A-73-1,
148-150 cm to 547 A-70-2, 70-93 cm, where X. ceratioides is absent,
can bedated as Albian, whereas the first record of this speciesin
Sample 547A-69-2, 108-110 cm implies that the sam-ple is
Cenomanian.
The section from Sample 547A-69-2, 108-110 cm toSample
547A-44-2, 0-2 cm belongs to the CenomanianCleistosphaeridium
polypes Assemblage Zone of Wil-liams (1975), erected on the basis
of dinoflagellate cystmaterial from Scotian shelf sections. Besides
other spe-cies listed by Williams, Kiokansium corollum ( = C.
po-lypes sp. A sensu Williams, 1978) does not range into
Occ
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6894
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6899
6900
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6906
6907
6908
6909
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R. BELOW
Figure 5. Stratigraphic ranges of selected dinoflagellate cysts
in Hole 547A, between Cores 37 and 73 and Hole 547B, be-tween Cores
1 and 5. X = dinoflagellate cyst taxa also present in the Aptian
(Apt), Albian (Alb), and Cenomanian (C)of Hole 545.
632
-
APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS, MAZAGAN PLATEAU
late cyst populations found in the interval from
Sample547A-69-2, 108-110 cm to 547A-44-2, 0-2 cm, and theclast is
therefore dated as earliest Cenomanian.
CONCLUSIONSTo summarize the results of this biostratigraphic
chap-
ter, age determinations based on dinoflagellate cyst analy-sis
are, for Holes 547,547A, and 547B:
545-32-1, 29-33 cm to 545-31-1, 130-133 cm: early
Cenomanian545-37-2, 84-87 cm to 545-33-1, 90-94 cm: earliest
Cenomanian545-40-2, 116-119 cm to 545-38-1, 120-123 cm: late
Albian545-48-6, 16-18 cm to 545-40-6, 57-60 cm: late
Aptian545-53-3, 90-94 cm to 545-49-1, 34-37 cm: middle
Aptian545-56-6, 55-57 cm to 545-54-3, 43-47 cm: early (?)
Aptian
547A-69-2, 108-110 cm to 547A-44-2, 0-2 cm: earliest
Cenoma-nian
547A-73-1, 148-150 cm to 547A-70-2, 90-93 cm: late Albian
547B-5-7, 40-43 cm to 547B-2-2, 0-2 cm: late Albian
Sites 545 and 547 are correlated by comparing the
Al-bian/Cenomanian boundary, marked by the base rangeof X.
ceratioides at the beginning of the Cenomanian.Core 545-37 is
comparable to Core 547A-69, since bothcores include this
boundary.
ADDENDUM
At the postcruise meeting for Leg 79 it was decided that 18
addi-tional samples of Hole 547B should be analyzed for
dinoflagellatecysts. These samples are:
547B-6-2, 6-7 cm; 547B-6-3, 57-58 cm; 547B-6-4, 76-77 cm;
547B-7-4, 54-56 cm; 547B-7-3, 124-126 cm; 547B-7-3, 94-96 cm;
547B-7-2,133-135 cm; 547B-7-1, 42-43 cm; 547B-8-3, 26-27 cm;
547B-8-2, 118-120 cm; 547B-8-1, 89-91 cm; 547B-10-4, 50-51 cm;
547B-10-3, 9-10cm; 547B-10-2, 125-126 cm; 547B-11-4, 80-81 cm;
547B-11-3, 3-4 cm;547B-11-2, 138-139 cm; 547B-11-1, 139-140 cm. The
samples weretaken from limestones with clay interbeds located
immediately belowthe section studied in this chapter and dated by
shipboard paleontolo-gists as Early Cretaceous to Middle Jurassic.
All these samples werepalynologically barren.
TAXONOMY
This section is divided into two parts. The first lists names of
alldinoflagellate cyst taxa encountered during this study, arranged
in al-phabetical order of genus names, with references to plate and
figurenumbers where applicable; numbers in parentheses refer to the
posi-tion of the species in the distribution charts for Holes 545,
547B, and547A (Fig. 1,3, and 4, respectively). A zero indicates
that the speciesis absent from that particular hole. Species marked
with an asteriskare discussed in more detail in the second part,
which provides mor-phologic descriptions or systematic
classifications.
Illustrated specimens are housed in the micropaleontological
col-lection of the Institut für Palàontologie, Rheinische
Friedrich-WilhelmsUniversitàt Bonn, Germany.
Dinoflagellate Cyst Taxa
Achomosphaera neptuni (Eisenack, 1958) Davey and Williams,
1966.(58-0-0)
Achomosphaera ramulifera (Deflandre, 1937) Evitt, 1963; Plate
1,Fig. 1. (150-45-48)
Achomosphaera sagena Davey and Williams, 1966; Plate 1, Fig
2.(157-0-82)
Achomosphaera triangulata (Gerlach, 1961) Davey and
Williams,1969; Plate 1, Fig. 3. (43-47-72)
Achomosphaera verdieri Below (1982); Plate 1, Fig. 4.
(34-0-0)Aptea almohadensis n. sp.; Plate 1, Figs. 5, 6, 7.
(96-0-0)Aptea eisenackii (Davey, 1969) Davey and Verdier, 1974;
Plate 1, Fig 8,
9.(17-54-68)Aptea polymorpha Eisenack, 1958; Plate 1, Fig. 10,
(31-0-0)
Apteodinium maculatum Eisenack and Cookson, 1960; Plate 1,
Fig.11. (48-0-0)
Batiacasphaera saidensis Below, 1981; Plate 1, Fig. 12.
(100-0-0)Callaiosphaeridium trycherium Duxbury, 1980.
(78-82-0)Canningia reticulata Cookson and Eisenack, 1960.
(40-0-0)Canningia ringnesiorum Manum and Cookson, 1964.
(44-69-40)Carpodinium granulatum Cookson and Eisenack, 1962; Plate
1, Figs.
13, 14.(98-0-0)Cassiculosphaeridia magna Davey, 1974.
(0-0-88)Cassiculosphaeridia reticulata Davey, 1969; Plate 2, Fig.
2. (83-64-36)Cassiculosphaeridia tazad ensis Below, 1981.
(92-0-0)Cerbia aucda Below, 1981. (60-0-0)Cerbia tabulata (Davey
and Verdier, 1974) Below, 1981; Plate 2, Fig. 2.
(9-0-0)Chlamydophorella discreta Clarke and Verdier, 1967.
(135-67-70)Chlamydophorella nyei Cookson and Eisenack, 1958.
(106-30-15)Cleistosphaeridium clavulum (Davey, 1969) Below (1982);
Plate 2, Fig.
3. (49-20-85)Cleistosphaeridium? huguoniotii (Valensi, 1955)
Davey, 1978. (118-0-
0)Cleistosphaeridium multispinosum (Singh, 1964) Brideaux,
1971;
Plate 2, Fig. 4. (0-57-0)Codoniella campanula (Cookson and
Eisenack, 1960) Downie and
Sarjeant, 1965, emend. Davey, 1979; Plate 2, Fig. 5.
(121-23-32)Codoniella psygma Davey, 1979; Plate 2, Figs. 6, 7.
(111-0-0)Cometodinium? whitei (Deflandre and Courteville, 1939)
Stover and
Evitt, 1978; Plate 2, Fig. 8. (35-21-64)Coronifera oceanica
Cookson and Eisenack, 1958; Plate 2, Fig. 9.
(51-24-10)Coronifera tubulosa Cookson and Eisenack, 1974; Plate
2, Fig. 10.
(103-70-0)Cribroperidinium cooksoniae Norvick, 1976.
(87-48-33)Cribroperidinium diaphane (Cookson and Eisenack, 1958)
Stover and
Evitt, 1978. (109-0-0)Cribroperidinium cf. diaphane (Cookson and
Eisenack, 1958) Stover
and Evitt, 1978. (5-0-0)Cribroperidinium edwardsii (Cookson and
Eisenack, 1958) Davey,
1969. (81-0-0)Cribroperidinium orthoceras Eisenack, 1958; Plate
2, Fig. 11. (12-83-
61)Cribroperidinium cf. orthoceras Eisenack, 1958 sensu Below,
1981.
(59-0-0)Cribroperidinium tensiftense Below, 1981.
(62-16-56)Cyclonephelium brevispinatum (Millioud, 1969) Yun, 1981;
Plate 2,
Fig. 12. (21-5-16)Cyclonephelium chabaca Below, 1981; Plate 2,
Fig. 13. (153-0-38)Cyclonephelium cf. chabaca Below, 1981.
(0-0-69)Cyclonephelium compactum Deflandre and Cookson, 1955; Plate
2,
Fig. 14; Plate 3, Fig. 1. (0-44-0)Cyclonephelium distinctum
Deflandre and Cookson, 1955; Plate 3,
Figs. 2, 3. (52-63-7)Cyclonephelium hystrix (Eisenack, 1958)
Davey, 1978; Plate 3, Fig. 4.
(19-0-0)Cyclonephelium maugaad Below, 1981; Plate 3, Fig. 5.
(13-0-0)Cyclonephelium cf. membraniphorum Cookson and Eisenack,
1962;
Plate 3, Fig. 6. (142-55-1)Cyclonephelium paucimarginatum
Cookson and Eisenack, 1962;
Plate 3, Fig. 7. (25-79-65)Cyclonephelium vannophorum Davey,
1969. (148-0-26)Cyclonephelium cf. vannophorum Davey, 1969.
(26-0-0)Danea chibanis Below, 1981. (70-0-0)Dingodinium albertii
Sarjeant, 1966; Plate 3, Fig. 8. (115-0-0)Dissiliodinium globulum
Drugg, 1978; Plate 3, Figs. 9, 10, 11. (11-
49-0)*Ellipsoidictyum imperfectum (Brideaux and Mclntyre, 1975)
Lentin
and Williams, 1977; Plate 3, Fig. 12. (114-0-0)Endoceratium
ludbrookii (Cookson and Eisenack, 1958) Loeblich and
Loeblich, 1966. (0-0-83)Endoscrinium campanula (Gocht, 1959)
Vozzhennikova, 1967 ssp. ni-
chan Below, 1981. (107-0-0)Epelidosphaeridia spinosa (Cookson
and Hughes, 1964) Davey, 1969;
Plate 3, Fig. 13. (158-0-0)Exochosphaeridium bifidum (Clarke and
Verdier, 1967) Clarke, Da-
vey, Sarjeant, and Verdier, 1968; Plate 3, Fig. 14.
(15-50-0)Exochosphaeridium phragmites Davey, Downie, Sarjeant, and
Wil-
liam, 1966. (63-56-29)
633
-
R. BELOW
Eyrea nebulosa Cookson and Eisenack, 1958; Plate 3, Fig. 15.
(131-65-51)
Florentinia berran Below (1982). (0-84-59)Florentinia laciniata
ssp. laciniata Davey and Verdier, 1973; Plate 4,
Fig. 1. (159-0-73)Florentinia laciniata Davey and Verdier, 1973
ssp. seghiris Below (in
press). (66-0-0)Florentinia mantellii (Davey and Williams, 1966)
Davey and Verdier,
1973; Plate 4, Fig. 2. (53-25-39)Florentinia radiculata (Davey
and Williams, 1966) Davey and Verdier,
1973, emend. Davey and Verdier, 1976; Plate 4, Fig. 3.
(20-6-17)Florentinia resex Davey and Verdier, 1976; Plate 4, Fig.
4. (146-85-42)Florentinia stellata (Maier, 1959) Below (1982);
Plate 4, Fig. 5. (154-
17-57)Fromea amphora Cookson and Eisenack, 1958; Plate 4, Fig.
6. (75-
60-0)Gonyaulacysta cassidata (Eisenack and Cookson, 1960)
Sarjeant,
1966. (149-0-80)Gonyaulacysta cretacea (Neale and Sarjeant,
1962) Sarjeant, 1969;
Plate 4, Fig. 7. (41-22-49)Gonyaulacysta helicoidea (Eisenack
and Cookson, 1960) Sarjeant,
1966; Plate 4, Fig. 8. (22-26-12)Hapsocysta dictyota Davey,
1979; Plate 4, Fig. 9. (0-31-52)Hapsocysta peridyctia (Eisenack and
Cookson, 1960) Davey 1979.
(95-58-0)Heterosphaeridium heteracanthum (Deflandre and Cookson,
1955)
Eisenack and Kjellström, 1971. (16-0-2)Hexagonifera cf.
chlamydata Cookson and Eisenack, 1962; Plate 4,
Fig. 10. (68-59-78)Histiocysta outananensis Below, 1981.
(120-0-0)Hystrichodinium compactum Alberti, 1961.
(113-0-0)Hystrichodinium pulchrum Deflandre, 1935.
(104-71-3)Hystrichosphaeridium arundum Eisenack and Cookson, 1960;
Plate
4, Fig. 11. (138-3-0)Hystrichosphaeridium cf. arundum Eisenack
and Cookson, 1960;
Plate 4, Fig. 12. (0-74-0)*Hystrichosphaeridium atlasense Below
(1982); Plate 4, Fig. 13. (0-77-
0)Hystrichosphaeridium bowerbankii Davey and Williams, 1966;
Plate
5, Fig. 1. (160-0-5)Hystrichosphaerina schindewolfii Alberti,
1961. (122-0-0)Kiokansium corollum (Hasenboehler, in press); Plate
5, Fig. 2, 3, 4.
(139-38-22)Kiokansium hydra (Duxbury, 1979) Below (1982); Plate
5, Fig. 5. (14-
0-0)Kiokansium polypes (Cookson and Eisenack, 1964) Below (in
press);
Plate 5, Fig. 5. (74-86-0)Kleithriasphaeridium eoinodes
(Eisenack, 1958) Davey, 1974; Plate 5,
Fig. 7. (32-0-0)Kleithraisphaeridium? sarmentum (Davey, 1979)
Below (1982); Plate
5, Fig. 8. (128-0-0)Litosphaeridium conispinum Davey and
Verdier, 1973; Plate 5, Fig. 9.
(134-4-6)Litosphaeridium siphoniphorum (Cookson and Eisenack,
1958) Da-
vey and Williams, 1966; Plate 5, Figs. 10, 11
(136-78-43)Maghrebinia chleuh Below, 1981; Plate 6, Fig. 1.
(147-2-31)Maghrebiniaperforata (Clarke and Verdier, 1967) Below,
1981 ssp. mi-
rabilis n.ssp.; Plate 6, Fig. 2. (140-32-9)*Maghrebinia
perforata (Clarke and Verdier, 1967) emend. Below, 1981
ssp. perforata; Plate 6, Fig. 3. (155-27-58)Meiourogonyaulax cf.
bulloidea (Cookson and Eisenack, 1960) Sar-
jeant, 1969; Plate 6, Fig. 4. (67-0-0)Meiourogonyaulax stoveri
Millioud, 1969. (36-0-0)Muderongia perforata Alberti, 1961; Plate
6, Figs. 5, 6. (8-87-0)Nematosphaeropsis? singularis Davey, 1979.
(126-0-0)Occisucysta coummia Below, 1981. (0-39-62)Occisucysta
hinzii n.sp.; Plate 6, Figs. 7, 8. (99-0-0)*Occisucysta
mazaganensis n.sp.; Plate 6, Fig. 9; Plate 7, Fig. 1. (143-
33-0)*Occisucysta sousensis Below, 1981; Plate 7, Fig. 2
(37-72-0)Occisucysta tentoria Duxbury, 1977. (94-0-0)Occisucysta
tenuiceras (Eisenack, 1958) Below, 1981. (94-0-0)Occisucysta cf.
tenuiceras (Eisenack, 1958) Below, 1981. (123-0-0)Odontochitina
cribropoda Deflandre and Cookson, 1955. (0-88-0)Odontochitina
operculata (O. Wetzel, 1933) Deflandre and Cookson,
1955; Plate 7, Fig. 3. (54-7-18)
Oligosphaeridium albertense Pocock, 1962.
(112-43-0)Oligosphaeridium asterigerum (Gocht, 1959) Davey and
Williams
1969; Plate 7, Fig. 4. (28-34-86)Oligosphaeridium complex
(White, 1842) Davey and Williams, 1966;
Plate 7, Fig. 5. (3-9-19)Oligosphaeridium dividuum Williams,
1978; Plate 7, Fig. 6. (84-0-84)Oligosphaeridium djenn Below (in
press). (27-0-0)Oligosphaeridium indicum (Jain, 1977) Below (1982);
(124-0-89)Oligosphaeridium irregulare (Pocock, 1962) Davey and
Williams,
1969;Oligosphaeridium poculum Jain, 1977; Plate 7, Fig. 8.
(47-51-30)Oligosphaeridium pulcherrimum (Deflandre and Cookson,
1955) Da-
vey and Williams, 1966; Plate 7, Fig. 9.
(133-76-60)Oligosphaeridium totum Brideaux, 1971 ssp. totum.
(45-42-91)Oligosphaeridium totum Brideaux 1971 ssp. minor
(Brideaux, 1971)
Lentin and Williams, 1973; Plate 7. Fig. 10.
(1-40-81)Oligosphaeridium verrucosum Davey, 1979; Plate 7, Figs.
11, 12, 13.
(80-0-0)*Oodnadattia alata (Cookson and Eisenack, 1962) Below,
1981; Plate
8, Figs. 1, 2. (152-1-20)Oodnadattia tuberculata Eisenack and
Cookson, emend, Below, 1981;
Plate 8, Figs. 3,4. (141-62-11)Operculodinium? hirsutum
(Ehrenberg, 1838) Lentin and Williams,
1973.(71-0-0)Ovoidinium diversum? Davey, 1979; Plate 8, Fig. 5
(0-90-0)Ovoidinium implanum Davey, 1979; Plate 8, Fig. 6.
(0-0-79)Ovoidinium scabrosum (Cookson and Hughes, 1964) Davey,
1970;
Plate 8, Fig. 7, 8. (69-68-76)Ovoidinium verrucosum (Cookson and
Hughes, 1964) Davey, 1970;
Plate 8, Figs. 9, 10. (156-0-50)Palaeohystrichophora
infusorioides Deflandre, 1935; Plate 8, Fig. 11.
(144-0-0)Palaeoperidinium cretaceum Pocock, 1962; Plate 8, Fig.
12. (105-52-
67)Pervosphaeridium pseudohystrichodinium (Deflandre, 1937)
Yun,
1981; Plate 8, Fig. 13. (0-36-21)Pervosphaeridium truncatum
(Davey, 1969) Below (1982). Plate 8,
Fig. 14. (23-8-13)Phoberocysta neocomica Gocht, 1957.
(97-0-0)Polysphaeridium duma Below (1982). (33-0-0)Polysphaeridium?
flexuosum (Davey, Downie, Sarjeant, and Wil-
liams) Below (in press). (39-0-0)Polysphaeridium sp. sensu Below
(1982); Plate 8, Fig. 15. (116-75-0)Polysphaeridium warrenii Habib,
1975; Plate 9, Fig. 1. (64-37-90)Prolixosphaeridium conulum Davey,
1969; Plate 9, Fig. 2. (86-19-71)Prolixosphaeridium parvispinum
(Deflandre, 1937) Davey and Wil-
liams, 1969 ssp. deirense (Davey, Downie, Sarjeant, and
Williams,1966) Below (1982); Plate 9, Fig. 3. (93-0-0)
Prolixosphaeridium parvispinum (Deflandre, 1937) Davey and
Wil-liams, 1969 ssp. parvispinum; Plate 9, Fig. 4. (42-66-14)
Protoellipsodinium seghire Below, 1981 ssp. seghire.
(85-0-0)Protoellipsodinium spinosum Davey and Verdier, 1971.
(129-0-0)Protoellipsodinium touile Below, 1981 ssp. touile.
(76-0-0)Pseudoceratium gochtii Neale and Sarjeant, 1962.
(101-0-0)Pterodinium aliferum Eisenack, 1958. (77-0-0)Pterodinium
agadirense Below, 1981. (88-0-0)Pterodinium bab Below, 1981.
(110-0-0)Pterodinium cingulatum (O. Wetzel, 1933) Below, 1981 ssp.
cingula-
tum; Plate 9, Fig. 5 (65-53-46)Pterodinium cornutum Cookson and
Eisenack, 1962; Plate 9, Figs. 6,
7.(46-81-41)Pterodinium mamounia Below, 1981.
(108-0-0)Rhynchodiniopsis aptiana Deflandre, 1935.
(38-0-0)Spiniferites ancoriferus Cookson and Eisenack, 1974, ssp.
ancori-
ferus; Plate 9, Fig. 8. (55-28-23)Spiniferites ancoriferus
Cookson and Eisenack, 1974 ssp. ghiran Be-
low (1982). (137-0-0)Spiniferites cornutus (Gerlach, 1961)
Sarjeant, 1970; Plate 9, Fig. 9.
(119-0-87)Spiniferites dentatus (Gocht, 1959) Lentin and
Williams, 1973,
emend. Duxbury, 1977. (73-0-0)Spiniferites hyperacanthus
(Deflandre and Cookson, 1955) Cookson
and Eisenack, 1974; Plate 9, Fig. 10. (29-0-37)Spiniferites
lenzii Below (1982); Plate 9, Fig. 11. (56-73-34)Spiniferites
magnoserratus (Cookson and Eisenack, 1962) Stover and
Evitt, 1978. (30-0-0)
634
-
APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS, MAZAGAN PLATEAU
Spiniferites multibrevis (Davey and Williams, 1966) ssp. ghiran
Below(1982); Plate 9, Fig. 12. (57-41-75)
Spiniferites multibrevis (Davey and Williams, 1966) Below (1982)
ssp.multibrevis; Plate 9, Fig. 13. (6-78-45)
Spiniferites multibrevis (Davey and Williams, 1966) Below (1982)
ssp.seghiris Below (1982); Plate 9, Fig. 14. (24-11-24)
Spiniferites ramosus (Ehrenberg, 1838) Loeblich and Loeblich,
1966ssp. ramosus. (2-89-55)
Spiniferites ramosus (Ehrenberg, 1838) Loeblich and Loeblich,
1966ssp. reticulatus (Davey and Williams, 1966) Lentin and
Williams,1973.(82-0-74)
Spiniferites cf. wetzelii (Deflandre, 1937) Sarjeant, 1970;
Plate 9, Fig.15. (89-46-0)*
Subtilisphaera deformans (Davey and Verdier, 1973) Stover and
Evitt,1978; Plate 10, Fig. 1. (102-0-0)
Subtilisphaera cheit Below, 1982. (132-10-35)Subtilisphaera
scabrata Jain and Millepied, 1973. (79-29-27)Subtilisphara
senegalensis Jain and Millepied, 1973. (10-0-4)Surculosphaeridium
trunculum Davey, 1979. (90-12-77)Systematophora? sp. MAZ; Plate 10,
Fig. 2. (91-0-63)*Systematophora complicata Neale and Sarjeant,
1962. (125-0-0)Systematophora cretacea Davey, 1979; Plate 10, Figs.
3, 4. (117-0-0)Systematophora silyba Davey, 1979.
(61-14-0)Tanyosphaeridium boletum Davey, 1974; Plate 10, Fig. 5.
(4-0-53)Tanyosphaeridium regulare Davey and Williams, 1966.
(50-61-66)Trabeculodinium quinquetrum Duxbury, 1980; Plate 10, Fig.
6. (0-
15-28)Trichodinium castanea (Deflandre, 1935) Clarke and
Verdier, 1967;
Plate 10, Fig. 7. (72-13-25)Wallodinium luna (Cookson and
Eisenack, 1960) Lentin and Wil-
liams, 1973; Plate 20, Fig. 8. (127-0-0)Xenascus ceratioides
(Deflandre, 1937) Lentin and Williams, 1973;
Plate 10, Figs. 9-12. (151-0-47)Xenascus plotei Below, 1981.
(145-0-44)
Description of Selected Dinoflagellate Cyst Taxa
Class DINOPHYCEAE Fritsch, 1929Order PERIDINIALES Haeckel,
1894
Genus APTEA Eisenack 1958, emend. Davey and Verdier, 1974
Junior synonyms. Aptea Eisenack, 1958, emend. Dörhöfer
andDavies, 1980; Doidyx Sarjeant, 1966.
Type species. Apteapoiymorpha Eisenack, 1958.Diagnosis.
Eisenack, 1958, p . 393; Davey and Verdier, 1974, p. 640.
Aptea almohadensis n.sp.(Plate 1, Figs. 5, 6, 7)
Derivatio nominis. After the dynasty of the Almohavids, kings of
theMaghreb in the eleventh century.Diagnosis. Proximate, lenticular
cyst, shape subsphaeroidal to asym-
metrical; with a left antapical bulge and occasionally with a
right lat-eral bulge in the region of the postcingular paraplate
girdle. Apexrounded or with a narrow bulge. Paratabulation
presumably ceratia-cean—not completely indicated: 1 ' - 4 ' , 1 " -
6 " ' as, Xc, X " ' , Xs,X " " . Archeopyle apical, with simple,
free, asymmetrical opercu-lum, type 4 A ^ ^ , . Autophragma of
varying thickness, very finelyreticulated, rarely smooth. Cyst
ornamentation very sparse and inmost cases without rules of
orientation. Ornaments as low, solid, flat-tened, interconnected
ledges, isolated short spines with distally acumi-nated or blunted
ends, or simple cones forming tubercles. Sometimesthese ornaments
are arranged to pandasutural lines.
Holotype. Slide 11-6946: 49.6/100.1Type locality. Leg 79, Sample
545-49-5, 49-52 cm.Dimensions. Holotype length (without apical
calotte) 58 µm, breadth
60 µm, maximal height of ornaments 2 µm; other specimens:
length50-60 µm, breadth 60-75 µm, maximal height of ornaments 1-4
µm.
Description. The cyst shape is highly variable because of the
devel-opment of an apical, right lateral and an asymmetrically
arranged an-tapical bulge. Specimens with an asymmetrical shape
caused by thepresence of a left antapical bulge and therefore
typical of the genusAptea are present, as well as cysts with a
subsphaeroidal or sphaeroidaloutline typical of the genus
Cyclonephelium. The additional develop-ment of a right lateral
bulge shows the close connection of these speci-mens to individuals
of the genus Pseudoceratium. Reconstruction of
paratabulation is only indirectly possible for the epicyst by
studyingthe archeopyle suture and breakages of opercula and
archeopyle mar-gin. There are four apical paraplates. 1' and 2 '
are compact and po-lygonal, whereas 3 ' and 4 ' are typically
elongated and thus cause thevery asymmetrical operculum shape. The
precingular paraplate seriesis divided into six precingulars and
the plate as. Paratabulation of thehypocyst is unknown. The sparse
distribution of low ornaments suchas spines or tubercles makes any
consideration of penitabular align-ment along parasutures
impossible. Because of this pandasuturalzones are also not
developed. In few cases, ornamentation is absent.
Comparison. This new species is related to Aptea eisenackii
(Da-vey, 1969) Davey and Verdier, 1974. The sparse cyst
ornamentationcharacteristic of A. almohadensis is a result of the
strong reduction ofornaments in A. eisenackii.
Occurrence. See Figure 1, No. 96: Sample 545-49-5, 49-52 cm
toSample 545-47-2, 97-99 cm; middle to late Aptian.
Genus DISSILIODINIUM Drugg, 1978
Type species. Dissiliodinium globulum Drugg, 1978.Diagnosis.
Drugg, 1978, p. 45.
Dissiliodinium globulum Drugg, 1978(Plate 3, Figs. 9, 10,
11)
Remarks. Research results of archeopyle type and epicystal
parap-late patterns compiled by Below (1981) are verified by the
rich DSDPmaterial. Nevertheless, a few remarks about the existence
of specialapical structures that can be interpreted as reflections
of preapical platesof motile dinoflagellate theca must be added.
The structure, a small,smooth apical area, is somewhat raised
against the surrounding cystwall. In other cases, boundaries are
marked as faint lines that resultfrom a fusion of the small
granulae of the cyst wall. This interestingstructure, interpretated
as reflections of 1PR and 2PR, is illustrated inPlate 3, Fig.
10.
Occurrence. See Figure 1, No. 11: Sample 545-56-3, 58-60 cm
toSample 545-41-4, 0-4 cm, Aptian; Figure 3, No. 49: Sample
547B-5-5,100-103 cm to Sample 547B-4-3, 21-24 cm; late Albian.
Genus HYSTRICHOSPHAERIDIUM Deflandre, 1937,emend. Davey and
Williams, 1966
Type species. Hystrichosphaeridium tubiferum (Ehrenberg, 1838)
De-flandre, 1937, emend. Davey and Williams, 1966.
Diagnosis. Deflandre, 1937, p. 68; Davey and Williams, 1966b, p
. 55.
Hystrichosphaeridium cf. arundum Eisenack and Cookson,
1960(Plate 4, Fig. 12)
Remarks. The single specimen from the sample 547B-5-2, 47-50
cmis characterized by elongate, buccinate appendages that abruptly
wid-en distally to funnels having twice the diameter of the
process. In thisprocess morphology, the cyst differs from the
tubiferous appendagesof a typical Hystrichosphaeridium arundum
Eisenack and Cookson.The processes of H. cf. arundum with a length
of 0.5 cyst diameter aresignificantly longer than the processes of
H. arundum, with a lengthof 0.25 of a cyst diameter.
Occurrence. See Figure 3, No. 74: Sample 547B-5-2, 47-50 cm;
lateAlbian.
Genus MAGHREBINIA Below, 1981
Type species. Maghrebinia perforata (Clarke and Verdier 1967)
Below,1981.
Diagnosis. Below, 1981, p . 22.
Maghrebinia perforata (Clarke and Verdier, 1967)Below 1981 ssp.
mirabiüs n. ssp.
(Plate 6, Fig. 2)
Derivatio nominis. Latin mirabilis, beautiful.Diagnosis.
Chorate, somewhat lenticular cyst with bitrapezoidal size.
Divided into a small epicyst and a greater hypocyst by an
equatorialplanispiral intratabular membrane. Paraplates bordered by
high, dis-tally denticulated, perforated or unperforated membranes,
which areoften irregularly and deeply slit. An apical area with few
intratabularprocesses as well as the antapical, four precingular,
and three post-cingular paraplates are bordered by these septa.
Nevertheless, the
635
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R. BELOW
paraplate scheme could be 3 ' - ? 4 ' , 5 " , Oc, 4 " ' , 1 " "
because thelarge ventral area between polar plates is
undifferentiated. The rightend of the paracingulum extends into
this area. The cyst surface is mi-cro frangate (see Below, 1981, p.
23). Endophragmal gonal membranalthickening is often perforated.
Archeopyle type A.
Holotype. The specimen shown as Maghrebinia perforata (Clarkeand
Verdier, 1967) Below, 1981 in Plate 1, Figs, la, lb, lc, in
Below1981; slide VIII-4744 1/8.
Type locality. Section Ida ou Tanane in the north of Agadir,
Mo-rocco, section-meter 152.
Type level, basal Cenomanian.Dimensions. Holotype: length 80 µm;
paratypes: length 62-96 µm,
breadth 60-90 µm.Remarks. The subspecies Maghrebinia perforata
(Clarke and Ver-
dier, 1967) Below, 1981 ssp. mirabilis n.ssp. differs from M.
perforata(Clarke and Verdier, 1967) Below, 1981 ssp. perforata by
its coarse anddeep, irregular splitting of the high parasutural
membranes. Septures ofthe letter subspecies are lower, are nearly
constant in height, and arefinely distally denticulated.
Two types of parasutural septa variations are now known: (1)
thesuccessive reduction of membrane height from M. perforata ssp.
mi-rabilis to M. perforata ssp. perforata; (2) the second type is
the inter-ruption of membranes by deep splits and consequently the
total ab-sence of membranes between gonal thickenings, as occurs in
the rangeof variation between M. perforata ssp. mirabilis and M.
chleuh Below,1981.
Occurrence. See Figure 1, No. 140: Sample 545-39-4, 94-97 cm
toSample 545-31-1, 130-133 cm, late Albian, basal and early
Cenoma-nian; Figure 3, No. 32: Sample 549B-5-6, 50-53 cm to Sample
549B-2-2,0-2 cm, late Albian; Figure 4, No. 9, Sample 549A-73-1,
148-150 cm toSample 547A-48-2, 75-77 cm, late Albian to basal
Cenomanian.
Genus OCCISUCYSTA Gitmez, 1970
Type species. Occisucysta balios Gitmez, 1970.Junior synonym.
Diacanthum Habib, 1972.Diagnosis. Gitmez 1970, p. 267.
Occisucysta hinzii n.sp.(Plate 6, Figs. 7, 8; text, Fig. 6)
Derivatio nnminis. In honor of Dr. Karl Hinz, Co-Chief Scientist
ofLeg 79.Diagnosis. Large, proximate, two-layered, cornucavate,
oval cyst
with broad, pyramidal apical horn. Periphragma irregularly
thickenedto alveolate or forming a reticulum with muri and soli of
variablewidth and shape. Very sparsely granulated. Endophragma
thin, finelyperforated. Paratabulation goniaulacacean ?PR, 4 ' , 6
" , Xc, as, ra,Is, rs, pv, 6 " ' , 1 " " . Parasutures marked by
low, thick ridges thatcan expand to low septa. The hypocyst is
distorted against the epicys-tal plate series so that 3 " V 4 " '
parasuture is mid-dorsal. Intratabu-lar pseudosutures occasionally
present. Archeopyle 2P2"_3"
Holotype. Slide IV 6938: 41.4/101.Type locality. DSDP Leg 79,
545-45-4, 40-43 cm, Mazagan Pla-
teau, off Northwest Africa.Type level, late Aptian.Dimensions.
Holotype: length 78 µm, breadth 67 µm, length of en-
docyst 60 µm, height of septa up to 3 µm; other specimens:
length72-104 µm, length of endocyst 60-86 µm, width 60-84 µm,
height ofsepta up to 4 µm.
Description. The large, two-layered cyst has a basically oval
ambi-tus with an obvious, broad-based, hollow, equilateral
pyramidal apicalhorn, which is positioned slightly off the anterior
cyst pole and dis-placed in ventral direction (Plate 6, Fig. 8).
The two cyst wall layers areclosely attached, except at the apex
where the outer layer forms thecornucavate apical protuberance. The
periphragma is irregularly thickenedand so gives rises to uneven,
broad and high muri around soli of varia-ble size and form (Fig.
6). Under light microscopy, the cyst wall ap-pears alveolate. A few
specimens show special thickenings of the murito small
intraparatabular pseudosutures, as described for Occisucystabrixii
Below (1982a), or thickenings of intrabular areas as shown
forOccisucysta evittii (Dodekova, 1969) Gitmez, 1970 by Below
(1982a).General reduction of periphragmal structures is possible.
Occasionalspecimens show nodes and tubercles as relicts of muri on
a smooth orfinely reticulated periphragma. The endophragma is
smooth to finely re-ticulated. The cyst wall has low, thick
parasutural ridges or—on a few
2µ,m
Figure 6. Occisucysta hinzii n.sp. detail of cystwall; E,p =
perforatedendophragma; P,s = solid periphragma; g = granulae; m =
muri; s= solum.
specimens—narrow, perforated parasutural septa that are best
devel-oped around the antapex. The paratabulation is typically
goniaulaca-cean. Preapical paraplates cannot be observed with
absolute assur-ance, but there are faint lines on the apical horn
that imply the exist-ence of a top preapical 1PR and a second 2PR
on the dorsal side ofthe horn. Four apical paraplates can be
distinguished, a small, long1' , pentagonal 2 ' and 3 ' and the
rhombic 4' that fills the angle be-tween 5 ' ' and 6 ' ' . The
precingular series are trapezoidal or pentago-nal and of equal
size, except for the smaller 6 ' ' . The equatorially situ-ated
paracingulum is offset by its width at the parasulcus with its
firstand last paraplate bent posteriorly. The postcingular series
begins witha small rectangular 1 " ' . This is poorly separated
from the parasulcusby a parasutural rib, but differs by its
alveolate texture from thesmooth Is. The postcingular series
continues with the equilaterally tri-angular 2 ' " and the
trapezoidal 3 ' " to 6 " ' . The pv paraplate isvery elongated and
poorly defined against pr, ps, and Is, because ofthe absence of
parasutural ridges. In the parasulcus it is possible todistinguish
the elongated smooth Is, a long rs and an arrowshaped psthat is
bent into a small antapical 1 " " . Paraplate series of epicystand
hypocyst are distorted as in Cribroperidinium (Gocht, 1979; Be-low,
1981), so that the 3 " V 4 " ' boundary is mid-dorsal. The
ar-cheopyle-type is 2P 2"_ 3" , but in some specimens paraplate 1 "
and/or 4 " are lost also.
Comparison. The irregularly thickened cyst wall with
variousshapes of muri and soli is unique among species of the genus
Occisu-cysta.
Occurrence, see Figure 1, No. 99: Sample 545-49-5, 49-52 cm
toSample 545-44-2, 78-82 cm; middle to late Aptian.
Occisucysta mazaganensis n.sp.(Plate 6, Fig. 9; Plate 7, Fig. 1;
text, Fig. 7)
Derivatio nominis. Mazagan, town in Morocco. DSDP Leg 79
Sitesare situated near submarine Mazagan Plateau.Diagnosis. Large,
proximate, spheroidal, two-layered, acavate or
cornucavate cyst with a low apical bulge. Endophragma
smooth,periphragma reticulated, bearing solid, irregularly
interconnected spinesthat are strung together on parasutures.
Paratabulation goniaulaca-cean ?XPR, 4 ' , 6 " , Xc, as, rs, Is,
ps, pv, 6 " ' , 1' " ' , with torsion ofhypocyst against epicyst so
that the 3 ." V 4 " ' paraplate boundary ismid-dorsal. One or two
additional, isolated, solid spines per parap-late. Archeopyle type
2P2- - .3" .
Holotype. Slide III-6879: 40.6/107.7.Type locality. DSDP Leg 79,
547B-5-5, 100-103 cm; Mazagan Pla-
teau, off Northwest Africa.Type level, late Albian.Dimensions.
Holotype: length 86 µm, breadth 72 µm, length of
spines 6 µm to 10 µm; other specimens: length 78-95 µm,
breadth68-81 µm, length of spines 2-12 µm.
Description. The large, two-layered cyst has an overall oval to
sub-spheroidal ambitus. It tapers apically, forming an apical horn
or a low,apical bulge. The smooth endophragma and the reticulated
peri-
636
-
APTIAN TO CENOMANIAN DINOFLAGELLATE CYSTS, MAZAGAN PLATEAU
E,s
2µm
Figure 7. Occisucysta mazaganensis n.sp., detail of cystwall;
E,s = solid endophragma; P,p = perforated periphragma; pS =
parasuturalspine row.
phragma are attached. Only in a few cases do they form a
cornucavatepericoel in the apical region. The cyst wall gives rise
to solid acuminatespines of varying height and width, which are
arranged on parasu-tures. Spines are irregularly connected at
different heights (Fig. 7).Generally, there are one or two solid,
acuminate intratabular spines inthe center of most paraplates. A
goniaulacacean paratabulation is de-fined: ?XPR, 4 ' , 6 " , Xc,
as, rs, Is, ps, pv, 6 " ' , 1 " " . The preapicalplates are
difficult to distinguish because the apical horn builds a
com-plicated trestle, formed by the junctions of the parasutural
spine-rowsbetween apical paraplates. Nevertheless, the crownlike
morphology ofthe apical horn tip implies the existence of such
preapical paraplates.The apical series is comprised of a narrow,
long 1' , pentagonal 2' and3 ' and a rhombic 4 ' . Paraplate 4 '
fills the angle between 5 ' ' and 6 ' ' ,as is typical for the
species of the genus Occisucysta. The paraplates ofthe precingular
series are trapezoidal or pentagonal and equally large,except for
the smaller pentagonal 6 ' ' . The equatorially situated
para-cingulum is offset by its width at the parasulcus. In contrast
to otherparaplates, the paracingulum bottom is built up of a very
coarseperiphragmal net. The postcingular series begins with a small
rectan-gular 1 ' ' ' and continues with the equilateral triangular
2 ' ' ' and trap-ezoidal 3 " ' to 6' " . The pv plate is elongate
and polygonal. In theparasulcus, it is possible to observe the
anterior/posteriorly elongatedrs and Is and an arched ps. The
trapezoidal 1 " " is indented by thisps paraplate. Epicyst and
hypocyst series are distorted so that the3 ' ' ' / 4 " ' boundary
is mid-dorsal. Archeopyle type 2P2< ' . 3 " .
Comparison. The irregularly connected rows of parasutural
spines,which look like a crown of thorns, are typical for this
species, whichresembles only Occisucysta paucispina (Eisenack and
Cookson, 1960)Below, 1981. This species has few parasutural spines
which are notconnected or are only basally interconnected.
Occurrence. See Figure 1, No. 143: Sample 545-39-1, 93-96
cm,late Albian; Figure 3, No. 33: Sample 547B-5-6, 50-53 cm to
Sample547B-2-2, 0-2 cm, late Albian.
Genus OLIGOSPHAERIDIUM Davey and Williams, 1966
Type species. Oligosphaeridium complex (White, 1842) Davey
andWilliams 1966.
Diagnosis. Davey and Williams, 1966b, p. 70.
Oligosphaeridium verrucosum Davey, 1979(Plate 7, Figs. 11, 12,
13)
Remarks. Oligosphaeridium verrucosum, a very large species,tends
to show a greater degree of variability in cyst wall ornamenta-tion
than other species of the genus Oligosphaeridium. Within the
same population one can observe specimens with either dense or
sparsedistribution of large granulae; or individuals with a very
dense butfine granulation; or specimens with a scabrate or smooth
cyst wall.
Occurrence. See Figure 1, No. 80: Sample 545-52-2, 123-126 cm
toSample 545-47-6, 72-75 cm; middle to late Aptian.
Genus SPINIFERITES Mantell, 1850, emend. Sarjeant, 1970
TVpe species. Spiniferites ramosus (Ehrenberg, 1838) Loeblich
andLoeblich, 1966.
Diagnosis. Mantell, 1850, p. 191: Sarjeant, 1970, p. 75.
Spiniferites cf. wetzelii (Deflandre, 1937) Sarjeant, 1970(Plate
9, Fig. 15)
Remarks. This species is characterized by high parasutural
septabetween the gonal processes. These processes are only slightly
higherthan the parasutural membranes and are distally bifurcated or
trifur-cated. In contrast to the typical Spiniferites wetzelii with
distally smoothsepta, the membranes of the species discussed here
have broad but bi-furcate appendages.
Occurrence. See Figure 1, No. 89: Sample 545-51-1, 47-50
cm,middle Aptian; Figure 3, No. 46: Sample 547B-5-5, 100-103 cm to
Sam-ple 547B-5-1, 0-3 cm, late Albian.
Genus SYSTEMATOPHORA Klement, 1960
Type species. Systematophora areolata Klement, 1960.Diagnosis.
Klement, 1960, p. 61.
Systematophoral sp. MAZ(Plate 10, Fig. 2: text, Fig. 8)
Description. Spheroidal, two-layered chorate cyst.
Endophragmasmooth, periphragma finely fibrous, especially where
forming pro-cesses. Processes of one individual are all of equal
length—corre-sponding to half the cyst body diameter but of
somewhat unequal di-ameter. The processes are hollow and are either
tubiform or distallysomewhat widened. The process wall is made up
of very fine fiberswhich are irregularly connected to a fine-meshed
net. The distal pro-cess margins are deeply split. A few thin,
simple, solid appendages arepresent. The arrangement of processes
is unknown and reconstructionof paratabulation is therefore
impossible. Archeopyle unknown.
Remarks. There are too few well preserved and favorably
orientedspecimens to formally erect a new species. Nevertheless,
this strikingcyst is a typical component of some Aptian and
Cenomanian dinofla-gellate cyst assemblages.
637
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R. BELOW
Figure 8. Systematophora? sp. MAZ, fibrous processes of variable
morphology.
Occurrence: See Figure 1, No. 91: Sample 545-50-6, 56-58 cm
toSample 545-33-1, 90-94 cm, middle and late Aptian, basal
Cenoma-nian; Figure 4, No. 63: Sample 547-63-2, 0-2 cm, basal
Cenomanian.
ACKNOWLEDGMENTS
I wish to acknowledge Prof. H. K. Erben and Prof. K. J.
Müller,directors of the Institut für Palàontologie, Rheinische
Friedrich-WilhelmsUniversitàt Bonn, who made working facilities and
material availableto me. Acids used in the palynological
preparation were donated byBAYER AG, Leverkusen.
I would also express my thanks to Dr. H. Gocht, Tubingen, andDr.
J. Jansonius, Calgary, for critically reading the manuscripts and
toMrs. B. Martin, Bonn, for revising the English.
Financial support was given by the Heinrich-Hörlein-Stiftung
ofthe Fachgruppe Erdwissenschaften, Rheinische Feridrich-Wilhelms
Un-iversitàt Bonn.
REFERENCES
Alberti, G., 1961. Zur Kenntnis mesozoischer und alt-tertiafer
Dinofla-gellaten und Hystrichosphaerideen von Nord- und
Mitteldeutsch-land sowie einigen anderen europáischen Gebieten.
Palaeontographi-ca, A, 116:1-58.
Below, R., 1981. Dinoflagellaten-Zysten aus dem oberen Hauterive
bisunteren Cenoman Süd-West-Marokkos. Palaeontographica, B,
176:1-145.
, 1982a. Scolochorate Zysten der Gonyaulacaceae (Dinophy-ceae)
aus der Unterkreide Marokkos. Palaeontographica, B, 182:1-51.
., 1982b. Dinoflagellate cysts from the Valanginian to
lowerHauterivian of sections near Ait Hamouch/Morocco. Rev
Espan.Micropaleontol, 14:23-52.
Brideaux, W. W., 1971. Palynology of the Lower Colorado
Group,central Alberta, Canada, I. Introductory remarks. Geology
andmicroplankton studies. Palaeontographica, B, 135:53-114.
Brideaux, W. W., and Mclntyre, D. J., 1975. Miospores and
micro-plankton from Aptian-Albian rocks along Horton River,
Districtof Mackenzie. Geol. Surv. Can. Bull., 252:1-85.
Clarke, R. F. A., Davey, R. J., Sarjeant, W. A. S., and Verdier,
J.-P.,1968. A note on the nomenclature of some Upper Cretaceous
andEocene dinoflagellate taxa. Taxon, 17:181-183.
Clarke, R. F. A., and Verdier, J.-P., 1967. An investigation of
micro-plankton assemblages from the Chalk of the Isle of Wight,
En-gland. K. Nederl. Akad. Wetensch. Afd. Nat. Eerste Reeks,
24:1-96.
Cookson, I. C , and Eisenack, A., 1958. Microplankton from
Austra-lian and New Guinea Upper Mesozoic sediments. Proc. R.
Soc.Victoria, 70:19-79.
, 1960. Upper Mesozoic microplankton from Australia andNew
Guinea. Palaeontology, 2:243-261.
., 1962. Additional microplankton from Australian
Cretaceoussediments. Micropaleontology, 8:485-507.
_, 1974. Mikroplankton aus Australischen Mesozoischen
undTertiàren Sedimenten. Palaeontographica, B, 148:44-93.
Cookson, I. C , and Hughes, N. F., 1964. Microplankton from
theCambridge Greensand (mid-Cretaceous). Palaeontology,
7:37-59.
Davey, R. J., 1969. Non-calcareous microplankton from the
Cenoma-nian of England, northern France and North America, Part
I.Bull. Brit. Mus. Nat. Hist., Geol., 17:1-11.
, 1974. Dinoflagellate cysts from the Barremian of theSpeeton
clay, England. Symp. Stratig. Palynol. Birbal Sahni
Inst.Palaeobot., Spec. Publ., 3:41-75.
., 1978. Marine Cretaceous palynology of Site 361, DSDPLeg 40,
off southwest Africa. In Bolli, H. M., Ryan, W. B. F , et al.,Init.
Repts. DSDP, 4
