GeolCarp_Vol64_No3_209

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA

, JUNE 2013, 64, 3, 209—230 doi: 10.2478/geoca-2013-0015

Introduction

The intramontane Vienna Basin has produced by far the larg-
est  volumes  of  hydrocarbons  and  also  provided  Austria’s
earliest petroleum production (Hamilton et al. 2000). Oil and
gas production in the Vienna Basin has come from Neogene
basin-fill sandstones (termed the ‘First Floor’) and from un-
derlying  allochthonous  Triassic  dolomites  of  the  Northern
Calcareous  Alps  and  units  of  the  Rhenodanubian  Flysch
Zone (RFZ, ‘Second Floor’). Reservoirs within the underly-
ing subthrust zone comprise mainly Jurassic carbonates and
Cretaceous-Paleogene  sandstones  (‘Third  Floor’).  Some  ex-
ploration  of  deep,  autochthonous,  mainly  Jurassic-age,  sub-
thrust reservoirs below the Neogene of the Vienna Basin has
also been conducted (Hamilton et al. 2000). A commercial gas
field (Höflein NW of the city of Vienna, Fig. 1), reservoired in
Middle  Jurassic  cherty  sandstones,  has  been  discovered  in
autochthonous units below the RFZ (e.g. Hamilton et al. 2000).
The stratigraphy of the RFZ within one of the wells (Höflein 6
of OMV) at the gas field Höflein is the topic of this paper.

The RFZ, which constitutes a 500-km-long, imbricated

thrust pile, trends E-W to ENE-WSW, parallel to the northern
margin of the Eastern Alps. To the south of Lake Chiemsee, it
is interrupted for a short distance and consequently it has been
subdivided  into  eastern  and  western  parts  (e.g.  Egger  &
Schwerd  2008).  Investigations  of  this  study  concentrated  on
the  easternmost  part  of  the  RFZ,  the  Wienerwald  area  near
Vienna, and the subcrop in the Vienna Basin.

The sedimentary succession of the RFZ consists of deep-

water deposits, which have been considered a lithostrati-
graphic supergroup (Mattern 1999; Wortmann et al. 2004) or
group (Egger & Schwerd 2008). It is subdivided into a num-
ber of formations, from the Lower Cretaceous up to the

Organic-walled dinoflagellate cyst biostratigraphy of the

Well Höflein 6 in the Cretaceous—Paleogene Rhenodanubian

Flysch Zone (Vienna Basin, Austria)

OMAR MOHAMED

1,2

and MICHAEL WAGREICH

Geology Department, Faculty of Sciences, Minia University, El-Minia, Egypt; omar.mohamed@mu.edu.eg

Department of Geodynamics and Sedimentology, Center for Earth Sciences, University of Vienna, Althanstrasse 14, A-1090 Vienna, Austria

(Manuscript received May 24, 2012; accepted in revised form March 14, 2013)

Abstract: Palynological analysis of the Rhenodanubian Flysch Zone section recovered from Well Höflein 6 north of
Vienna allows the successful application of non-calcareous dinoflagellate biostratigraphy to the deep-water sediments
of the Greifenstein Nappe. All 62 cuttings samples contained organic-walled dinoflagellate cysts (dinocysts) and some
of them allow age-assessment. The results corroborated the presence of two thrust slices. The upper thrust unit A
comprises a Campanian to Lower Eocene succession including, from old to young, the Röthenbach Subgroup, Perneck
Formation, Altlengbach Formation and Greifenstein Formation. The lower thrust unit B contains in addition a pre-
Campanian base, probably the Wolfpassing Formation of Early to mid-Cretaceous age.

Key words: Cretaceous, Paleogene, Vienna Basin, Rhenodanubian Flysch Zone, biostratigraphy, organic-walled
dinoflagellate cysts.

Fig. 1.  Simplified  geological  map  of  the  study  area  at  the  eastern
margin  of  the  Eastern  Alps,  showing  the  location  of  the  Höflein
area  including  Well  Höflein 6  within  the  Rhenodanubian  Flysch
Zone (RFZ).

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GEOLOGICA CARPATHICA, 2013, 64, 3, 209—230; Electronic Supplement, i—xii

Eocene. Classically, biostratigraphy in these deep-water clas-
tic flysch deposits is hindered by the fact that deposition was
mainly  below  the  calcite  compensation  depth  (e.g.  Egger  &
Schwerd  2008;  Wagreich  2008).  Consequently,  stratigraphi-
cally useful macrofossils are missing almost completely, and
planktonic foraminifera are very rare. Calcareous nannoplank-
ton and agglutinated foraminifera are the main sources so far
for  biostratigraphic  subdivision  of  the  RFZ.  Organic-walled
microplankton  such  as  dinoflagellates  have  been  used  only
rarely (e.g. Kirsch 2000, 2003).

Planktonic foraminifera and calcareous nannoplankton are

rare in the Paleocene and Eocene successions in the Wiener-
wald area, such as the flysch units of the Höflein gas field.
Therefore, a reassessment of a well within the Höflein area,
Höflein 6, was carried out in the present study using organic
dinoflagellate  cysts  (dinocysts).  Dinocysts  provide  a  high
resolution chronostratigraphic framework for this part of the
RFZ. Tectonic interpretations regarding thrust units, hitherto
largely based on heavy mineral studies, can be confirmed by
dinoflagellate  biostratigraphy.  This  paper  presents  the  first
palynological study of Well Höflein 6, illustrating the distri-
bution of the dinocysts and their relative abundances. Thus,
dinocysts  are  useful  means  for  the  solution  of  stratigraphic
problems in flysch sediments despite problems of reworking
and borehole downfall.

Stratigraphy

The OMV Höflein 6 Well, situated at the longitude of

E 16°18

’38.13” and latitude N 48°19’43.77” (Fig. 1), was

analysed  stratigraphically  using  cuttings  samples  by  OMV
in-house lab based on a few biostratigraphic data by benthic
foraminifera and rare nannoplankton beside sediment-petro-
graphic  methods.  The  total  thickness  of  the  Rhenodanubian
Flysch  sediments  within  Well  Höflein 6  measures  ca.
2565 m. Studies of heavy minerals from the well have iden-
tified  different  contents  especially  of  zircon  and  garnet,  on
the basis of which several lithostratigraphic units can be dis-
tinguished.  According  to  unpublished  OMV  in-house  data
(R. Sauer, pers. comm. 2010), the RFZ section of Höflein 6
can be divided into two thrust units, an upper thrust unit A and
a  lower  thrust  unit B  (Table 1).  The  upper  thrust  unit A  is
composed from top to bottom by the Greifenstein Formation
(zircon-dominated  heavy  mineral  assemblages),  Altlengbach
Formation  (garnet-dominated  heavy  mineral  assemblages),
Perneck  Formation  (characteristic  red  shaly  interval)  and  the
Röthenbach  Subgroup  (garnet-dominated  heavy  mineral  as-
semblages). The lower thrust unit B is composed by a succes-
sion, from top to bottom, by again the Greifenstein Formation
(zircon-dominated  heavy  mineral  assemblages),  Altlengbach
Formation  (garnet-dominated  heavy  mineral  assemblages)
and  Wolfpassing  Formation  (zircon-dominated  or  mixed
heavy  mineral  assemblages).  The  Wolfpassing  Formation
may  form  a  separate  thrust  unit  below  thrust  unit B  (e.g.
Schnabel  1992)  but  is  herein  regarded  as  stratigraphically
connected to thrust unit B.

The modern lithostratigraphic subdivision of the RFZ is

based on Schnabel (1992), Faupl (1996), Wagreich (2008)

and Egger & Schwerd (2008). Biostratigraphic data from out-
crops in the Wienerwald and wells apart from Well Höflein 6
indicate  in  general  a  Late  Paleocene  to  Early  Eocene  age  of
the Greifenstein Formation (Thanetian—Ypresian, NP9—NP13,
see  Schnabel  1992),  although  a  significant  diachronism  was
noted  already  by  Hekel  (1968).  The  Altlengbach  Formation
ranges  from  Late  Campanian—Maastrichtian  up  to  the  Paleo-
cene  (CC22—NP8;  Schnabel  1992;  Egger  &  Schwerd  2008).
The  Perneck  Formation  (former  “Oberste  Bunte  Schiefer”,
e.g. Sauer et al. 1992) has a Late Campanian age to the west of
the  Wienerwald  area  (CC21—22a;  Egger  &  Schwerd  2008).
The  Röthenbach  Subgroup  (former  Zementmergelserie  and
partly Kahlenberg Formation, e.g. Sauer et al. 1992) is mainly
Campanian in age (CC18—CC21/22; Egger & Schwerd 2008).
For  the  Wolfpassing  Formation  a  Barremian  to  Aptian  and
questionable  Albian  age  was  reported  by  Grün  et  al.  (1972;
see also Sauer et al. 1992).

Material and methods

A total of 62 cuttings samples was selected on the basis of

variation  in  lithology  and  to  sample  especially  formation
boundaries  for  palynological  analysis  from  the  Well  Höf-
lein 6,  giving  a  mean  sample  interval  of  ca.  50 m  for  the
2565 m thick well section.

30 grams of dry sediment were crushed and treated with

cold  35%  HCl  for  one  day  in  order  to  remove  carbonates.
Adding  water  and  subsequent  decantation  was  carried  out
twice with a minimum interval of six hours. Then, the sam-
ples were treated with 38% HF for 1—2 days to remove sili-
cates. Adding water and decantation twice with a minimum
interval of seven hours followed the HF treatment. A small
amount  of  35%  HCl  was  added  again  to  the  samples  to  re-
move gel which may have formed during the previous step.
Water was added to samples for the last time and the samples
were put in an ultrasonic device for 10—30 seconds and then
sieved over 15 and 30 µm nylon meshes. A part of the resi-
due  was  mounted  in  glycerin  jelly  on  2  or  3  microscope
slides after extensive mixing to obtain homogeneity and then
covered  by  a  slide  cover  (20 40 mm).  One  of  these  slides
holds  the  residue  particles  over  15 µm  and  the  other  slides

Table 1: Lithostratigraphic units of the RFZ and their inferred ages
in Well Höflein 6 based on OMV internal reports (see also Sauer et
al. 1992). Originally, the Perneck Formation was termed “Bunte
Schiefer” and the Röthenbach Subroup was termed “Kahlenberg
Formation”.

Depth [m]

Lithologic units

Age

10–322

Greifenstein Formation

Late Paleocene–Early
Eocene

322–785

Altlengbach Formation

Late Campanian–Early

Paleocene

785–ca.1017 Perneck

Formation

Late

Campanian

1017–1210

Röthenbach Subgroup Campanian

er t

hru

(

)

1210–1520? Greifenstein Formation Late Paleocene–Early

Eocene

1520–2480

Altlengbach Formation Late Campanian–Early

Paleocene

2480–2561

Wolfpassing Formation Early Cretaceous?

er t

hru

uni

t (B)

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GEOLOGICA CARPATHICA, 2013, 64, 3, 209—230; Electronic Supplement, i—xii

hold the residue particles over 30 µm. The slides are stored
at the Department of Geodynamics and Sedimentology, Cen-
ter for Earth Sciences, University of Vienna, Austria.

All samples were scanned for dinocyst taxa and were

counted  for  allowing  identification  of  acmes,  better  correla-
tions within units, and to get some information on reworking
and/or  downhole  contamination.  Taxonomy  of  dinocysts  is
generally  based  on  Fensome  et  al.  (1993)  and  dinocyst  no-
menclature follows Fensome et al. (2008) – see Appendix A.
Most of biostratigraphically significant dinocyst taxa are doc-
umented in Figs. 3 to 12. The relative abundances of dinocysts
are shown in electronic Appendix B and C.

Flysch sediments, comprising turbidites and other deep-

water  mass-flow  deposits,  are  prone  to  ample  reworking  of
older sediments, thus last occurrences of species have to be
used  with  caution.  In  addition,  using  cuttings  samples  also
involves possible downhole contamination of younger strata
into  older  cuttings,  making  first  occurrences  questionable
and  hard  to  interpret.  Thus,  stratigraphic  interpretation  is
done  with  care,  using  assemblage  counts  and  acmes,  and
also  lithostratigraphic  correlations  to  evaluate  and  corrobo-
rate  stratigraphic  results.  In  the  following,  first  occurrence
(FO)  and  last  occurrence  (LO)  are  used  in  the  connotation
from old to young as used for stratigraphic (outcrop) sections
(and not as it may be used for a borehole drilled from young
to old), so that FO denotes the stratigraphic base and LO de-
notes the top of the stratigraphic range of a taxon.

Results

Dinocyst preservation

Of the 62 samples from the Well Höflein 6 most samples

were productive yielding dinocysts; only 9 samples were low
productive  and  yielded  less  than  50  specimens.  Marine  pa-
lynomorphs  dominate  most  samples,  while  the  sporomorph
component  is  composed  mainly  of  bisaccate  pollen  and
spores  with  a  very  low  relative  abundance  in  all  samples
(electronic  Appendix B,  C).  Preservation  of  dinocysts  is
moderate to good in most productive samples. The dinocysts
show a high diversity in most samples, with up to 82 species
per  sample  at  the  depth  of  300 m.  A  total  of  292  dinocyst
species  and  subspecies  have  been  identified  from  the  Well
Höflein 6 and are listed in Appendix A.

Dinocyst bioevents

In the upper thrust unit A, the FO of Trithyrodinium evittii

(Fig. 8.13,14)  is  recorded  at  the  depth  of  1210 m,  the  bio-
stratigraphically  lowest  event  within  the  Röthenbach  Sub-
group.  The  FOs  of  Chatangiella  granulifera  (Fig. 7.2,3),
Palynodinium  grallator  (Fig. 3.7,8)  and  Cannosphaeropsis
utinensis  (Fig. 4.14,15)  are  present  at  the  depth  of  1170 m.
The  FO  of  Membranilarnacia?  tenella  (Fig. 11.1—3)  and
Odontochitina  operculata  (Fig. 10.3,4)  are  recorded  at  the
depth  of  1120 m.  The  FOs  of  Alterbidinium  acutulum
(Fig. 9.9,10),  Corradinisphaeridium  horridum  (Fig. 5.1,2),
Chatangiella  robusta  (Fig. 7.1),  Xenascus  gochtii  and  Tra-

beculidium  quinquetrum  (Fig. 11.9)  occur  at  the  depth  of
1070 m.  The  LOs  of  Odontochitina  porifera  (Fig. 10.1,2)
and Chatangiella granulifera occur at the depth of 1120 m.
Pervosphaeridium  pseudhystrichodinium  (Fig. 12.7)  and
Palaeoperidinium  pyrophorum  (Fig. 8.6)  acmes  are  present
at  the  depths  of  1120 m  and  1070 m  respectively  (Fig. 2).
The FO of Xenascus sarjeantii (Fig. 10.10—12) is recorded at
the depth of 1020 m, still within the Röthenbach Subgroup.

At the base of the overlying Perneck Formation Xenascus

gochtii  and  Palynodinium  grallator  have  their  LOs  at  the
depth  of  1010 m.  Within  the  Perneck  Formation,  the  FO  of
Diphyes  colligerum  (Fig. 6.1,2)  and  Manumiella  seelandica
(990 m) (Fig. 9.5) and the LO of Odontochitina operculata,
Florentinia mayi  (Fig. 5.11)  and  Trabeculidium  quinquetrum
are recorded (890 m).

Eisenackia circumtabulata (Fig. 6.7—9) first occurred at

the  base  of  the  overlying  Altlengbach  Formation  (780 m).
The LOs of Alterbidinium acutulum and Xenascus sarjeantii
are recorded at the depth of 790 m (Fig. 2). The FOs of Car-
patella  cornuta  (Fig. 5.3—6)  and  Senoniasphaera  inornata
(Fig. 5.7—9)  are  recorded  at  the  depth  of  680 m  and  Damas-
sadinium californicum (Fig. 5.12) first occurred at the depth
of  630 m,  within  the  Altlengbach  Formation.  The  FO  of
Glaphyrocysta exuberans (Fig. 6.12—15) occurs at the depth
of  430 m.  Cerodinium  diebelii  (Fig. 8.1)  and  Riculacysta
amplexa—Glaphyrocysta  semiticta  acmes  (Fig. 11.11—12;
Fig. 6.11) are recorded at the depth of 680 m and 430 m re-
spectively (Fig. 2).

Within the Greifenstein Formation, the FO of Apectodinium

homomorphum  (Fig. 3.5,6),  Diphyes  ficusoides  (Fig. 6.3,4)
and Deflandrea phosphoritica (Fig. 8.7,8) and the last occur-
rence  of  Membranilarnacia?  tenella,  Eisenackia  circumta-
bulata and Manumiella seelandica are recorded in the sample
at the depth of 300 m (Fig. 2). Particularly high abundances of
certain  taxa,  namely  the  Areoligera  coronata—Areoligera
senonensis  acme  (Fig. 3.9—11),  Glaphyrocysta  exuberans—
Glaphyrocysta  ordinate  acme  and  Paralecaniella  indentata
acme  (Fig. 11.13—15),  are  recorded  at  250,  200  and  100 m
depth respectively.

In the lower thrust unit B, the LOs of Muderongia extensive

(Fig. 10.5), Muderongia australis (Fig. 10.6), Leberidocysta?
laticaudata and Pseudoceratium pelliferum (Fig. 10.8) are re-
corded at the depth of 2530 m, within the Wolfpassing Forma-
tion. A Surculosphaeridium longifurcatum—Oligosphaeridium
complex acme (Fig. 12.1—2) was found at the same depth
(Fig. 2). The FOs of Areoligera coronata, Chatangiella ditis-
sima (Fig. 7.4,6), Alterbidinium acutulum, Cannosphaeropsis
utinensis and Cerodinium diebelii are present at the depth of
2480 m, at the boundary of the Wolfpassing Formation to the
overlying Röthenbach Subgroup.

Near the base of the Röthenbach Subgroup of the lower

thrust unit, the FOs of Areoligera senonensis, Trithyrodinium
evittii and Hystrichosphaeropsis obscura are recorded at the
depth of 2380 m. The FOs of Xenascus sarjeantii and Corra-
dinisphaeridium  horridum  are  recorded  at  the  depth  of
2130 m  and  2080 m  respectively.  The  LO  of  Corradini-
sphaeridium  horridum  is  recorded  at  the  depth  of  2030 m.
The LO of Odontochitina porifera, Alterbidinium acutulum,
Xenascus  sarjeantii  and  Palaeohystrichophora  infusorioides

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Fig. 2. Stratigraphic
distribution  of  se-
lected dinocyst spe-
cies  and  bioevents
in  the  Well  Höf-
lein 6.

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Fig. 3.  Characteristic  and  strati-
graphically important dinoflagel-
late  cysts  of  Well  Höflein 6.
The species name is followed by
the sample number and England
Finder coordinates (for localiza-
tion  of  the  specimen  on  the
slide);  all  scale  bars  are  20 µm.
1, 2 – Apectodinium augus-
tum  (Harland,  1979)  Lentin  &
Williams, 1981; H6-1520A, U4/2.
3, 4 – Apectodinium quinquela-

(Fig. 9.20) occur at the depth of 1930 m. The Areoligera se-
nonensis—Trithyrodinium  evittii  acme  and  Hystrichodinium
pulchrum acme are recorded at the depth of 2130 and 1930
respectively (Fig. 2). Within the Altlengbach Formation, the
FOs  of  Membranilarnacia?  tenella  and  Diphyes  colligerum
occur at the depth of 1880 m, followed by the FOs of Manu-
miella  seelandica  and  Manumiella  druggii  (Fig. 9.6)  at
1780 m  and  the  Glaphyrocysta  exuberans  acme  and  Cero-
dinium diebelii acme at the depth of 1730 m and 1680 m re-

spectively (Fig. 2). The FO of Damassadinium californicum
and Senoniasphaera inornata are recorded at the depth of
1630 m and 1580 m respectively.

The base of the Greifenstein Formation of the lower thrust

unit is characterized by the FO of Apectodinium augustum
(Fig. 3.1,2) at the depth of 1520 m. Manumiella druggii last
occurred at the depth of 1520 m. An acme of Palaeoperidinium
pyrophorum is recorded at the depth of 1580 m (Fig. 2). The
LO of Damassadinium californicum is recorded at the depth

tum (Williams & Downie, 1966) Costa & Downie, 1979; H6-2180A, F33/1. 5, 6 – Apectodinium homomorphum (Deflandre & Cookson,
1955) Lentin & Williams, 1977; emend. Harland, 1979; H6-300B, Q42. 7, 8 – Palynodinium grallator Gocht, 1970

;

H6-1170A, K2.

9 – Areoligera senonensis Lejeune-Carpentier, 1938; H6-1520A, P19/1. 10 – Areoligera coronata (Wetzel, 1933b) Lejeune-Carpentier,
1937; H6-1470A, S30/3. 11 – Areoligera guembelii Kirsch, 1991; H6-250A, V29. 12 – Areoligera coronata (Wetzel, 1933b) Lejeune-
Carpentier, 1937; H6-1470A, Q33/3. 13, 14 –

Areosphaeridium diktyoplokum (Klumpp, 1953) Eaton, 1971; emend. Eaton, 1971; H6-2530B,

F7/4. 15 – Wetzeliella symmetrica Weiler, 1956; H6-1830A, K33.

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Fig. 4. Characteristic and strati-
graphically  important  dinofla-
gellate cysts of Well Höflein 6.
The  species  name  is  followed
by the sample number and En-
gland  Finder  coordinates  (for
localization of the specimen on
the  slide);  all  scale  bars  are
20 µm.  1  –  Achomosphaera
regiensis

Corradini,

1973;

H6-1120A, G24/4. 2, 3 – Acho-

of 1470 m, Membranilarnacia? tenella at 1370 m, Apecto-
dinium  homomorphum  and  Diphyes  colligerum  at  1270 m,
and  Senoniasphaera  inornata,  Diphyes  colligerum  and
Manumiella  seelandica  at  1220 m.  The  Areoligera  corona-
ta—Areoligera  senonensis  and  Palaeoperidinium  pyropho-
rum  acmes  are  recorded  at  the  depth  of  1470 m,  and  the
Glaphyrocysta  exuberans—Glaphyrocysta  ordinate  acme  at
1420 m.  A  Paralecaniella  indentata  acme  is  present  at  the

depth of 1320 m and 1270 m (Fig. 2), within the Greifen-
stein Formation.

Interpretation

According to the above results, dinocysts provide a useful

tool for biostratigraphic studies in RFZ sediments. In the

mosphaera cf. alcicornu (Eisenack, 1954) Davy & Williams, 1966; H6-300A, B16/1. 4, 5 – Adnatosphaeridium tutulosum (Cookson &
Eisenack, 1960) Morgan, 1980; H6-20B, J9/4. 6 – Alisogymnium euclaense (Cookson & Eisenack, 1970) Lentin & Vozzhennikova, 1990;
H6-990B, J11. 7 – Dinogymnium acuminatum Evitt et al., 1967; H6-480B, B19. 8, 9 – Dinogymnium denticulatum (Alberti, 1961) Evitt
et al., 1967; H6-890B, Q15/4. 10 – Dinogymnium sp.; H6-1220B, L31. 11 – Batiacasphaera micropapillata Stover, 1977; H6-1220A,
A6. 12 – Batiacasphaera sp.; H6-300B, B16/3. 13 – Canningia reticulata Cookson & Eisenack, 1960; emend. Below, 1981; H6-1020A,
D33/2. 14 – Cannosphaeropsis utinensis Wetzel, 1933; emend. May, 1980; H6-1170A, D3. 15 – Cannosphaeropsis utinensis Wetzel,
1933; emend. May, 1980; H6-790A, H22.

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Well  Höflein 6,  despite  the  problems  of  reworking  and
downfall  contamination,  ages  and  stage  boundaries  can  be
indicated with some confidence by a combination of the first
(earliest)  and  last  (latest)  stratigraphic  occurrences  (FOs,
LOs) of some selected marker species based on comparisons
with world wide dinocyst studies.

The FOs of Apectodinium homomorphum, Diphyes ficu-

soides and Deflandrea phosphoritica are biostratigraphically

important species that are found at the depth of 300 m within
the Greifenstein Formation (Fig. 2). Previously, Apectodinium
homomorphum was recorded in the Upper Paleocene to Lower
Eocene  in  New  Zealand  (Wilson  1988)  and  in  St.  Pankraz,
Austria (Hofmann et al. 2011), in the Lower Eocene of north-
ern Spain (Caro 1973), Belgium (N) (de Coninck et al. 1983),
Germany  (Fechner  &  Mohr  1988;  Köthe  &  Piesker  2007),
north  Egypt  (El-Beialy  &  Shahin  1990),  English  Channel

Fig. 5.  Characteristic  and  strati-
graphically important dinoflagel-
late cysts of Well Höflein 6. The
species  name  is  followed  by  the
sample  number  and  England
Finder  coordinates  (for  localiza-
tion  of  the  specimen  on  the
slide);  all  scale  bars  are  20 µm.
1,  2  –  Corradinisphaeridium
horridum  (Deflandre,  1937b)
Masure,  1986;  emend.  Masure,
1986; H6-2080B, N38. 3, 4 – Carpatella cornuta Grigorovitch, 1969; emend. Fechner & Mohr, 1986; H6-580A, B15. 5, 6 – Carpatella
cornuta Grigorovitch, 1969; emend. Fechner & Mohr, 1986; H6-580A, K31/1. 7—9 – Senoniasphaera inornata (Drugg, 1970) Stover &
Evitt, 1978; H6-20A, C22/2. 10 – Hystrichokolpoma bulbosum (Ehrenberg, 1838) Morgenroth, 1968; emend. Morgenroth, 1968; H6-2530A,
J2. 11 – Florentinia mayi Kirsch, 1991; H6-1010A, V8. 12 – Damassadinium californicum (Drugg, 1967) Fensome et al., 1993; H6-320B,
A18/2. 13 – Disphaerogena carposphaeropsis Wetzel, 1933; emend. Sarjeant, 1985; H6-1470B, Y6/3. 14, 15 – Turnhosphaera hypoflata
(Yun Hyesu, 1981) Slimani, 1994; emend. Slimani, 1994; H6-1020A, G17/2.

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Fig. 6. Characteristic and strati-
graphically  important  dinofla-
gellate cysts of Well Höflein 6.
The  species  name  is  followed
by  the  sample  number  and
England Finder coordinates (for
localization of the specimen on
the  slide);  all  scale  bars  are
20 µm.  1,  2  –  Diphyes  col-
ligerum (Deflandre & Cookson,
1955)  Cookson,  1965;  emend.
Cookson,

1965;

H6-300B,

B16/3. 3, 4 – Diphyes ficusoides Islam, 1983; H6-300A, B16/1. 5, 6 – Eisenackia margarita (Harland, 1979) Quattrocchio & Sarjeant,
2003; H6-790B, L1/2. 7—9 – Eisenackia circumtabulata Drugg, 1967; H6-780A, D17. 10 – Glaphyrocysta perforata Hultberg &
Malmgren, 1985; H6-2130A, U6. 11 – Glaphyrocysta semiticta (Bujak in Bujak et al., 1980) Lentin & Williams, 1981; H6-1930A, M37/4.
12 – Glaphyrocysta exuberans (Deflandre & Cookson, 1955 ex Eaton, 1976) Stover & Evitt, 1978; emend. Sarjeant, 1986; H6-1170A, E29.
13 – Glaphyrocysta exuberans (Deflandre & Cookson, 1955 ex Eaton, 1976) Stover & Evitt, 1978; emend. Sarjeant, 1986; H6-1420A, X5.
14 – Glaphyrocysta exuberans (Deflandre & Cookson, 1955 ex Eaton, 1976) Stover & Evitt, 1978; emend. Sarjeant, 1986; H6-1420A, G3.
15 – Glaphyrocysta exuberans (Deflandre & Cookson, 1955 ex Eaton, 1976) Stover & Evitt, 1978; emend. Sarjeant, 1986; H6-1470B, Y5/4.

(Auffret & Gruas-Cavagnetto 1975) and in the Middle to Up-
per  Eocene  of  northern  Germany  (Costa  &  Martini  1981),
northern  France  (Châteauneuf  1980).  Williams  &  Bujak
(1985) documented that Apectodinium homomorphum ranges
from  the  Upper  Paleocene  to  the  Middle  Eocene.  Diphyes
ficusoides  is  recorded  in  the  Lower  Eocene  in  Virginia
(Edwards  1989),  Lower  to  Middle  Eocene  in  Germany

(Heilmann-Clausen  &  Costa  1989;  Köthe  &  Piesker  2007)
and  in  southern  England  (Islam  1983),  and  ranges  from  the
Middle to the Upper Eocene according to Stover et al. (1996).
Deflandrea phosphoritica is recorded in the Lower Eocene in
Germany  (Köthe  &  Piesker  2007).  The  co-occurrence  of  A.
homomorphum,  D.  ficusoides  and  D.  phosphoritica  in  Well
Höflein 6 at the depth of 300 m indicates a stratigraphic posi-

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Fig. 7. Characteristic and stratigraphically important dinoflagellate cysts of Well Höflein 6. The species name is followed by the sample
number and England Finder coordinates (for localization of the specimen on the slide); all scale bars are 20 µm. 1 – Chatangiella? robusta
(Benson,  1976)  Stover  &  Evitt,  1978;  H6-1070A,  U31/4.  2  –  Chatangiella  granulifera  (Manum,  1963)  Lentin  &  Williams,  1976;
H6-1730A, T23. 3 – Chatangiella granulifera (Manum, 1963) Lentin & Williams, 1976; H6-1170A, B17. 4 – Chatangiella ditissima
(McIntyre,  1975)  Lentin  &  Williams,  1976;  H6-250A,  H7.  5  –  Chatangiella  hexacalpis  Harker  &  Sarjeant  in  Harker  et  al.,  1990;
H6-1930C,  K18.  6  –  Chatangiella  ditissima  (McIntyre,  1975)  Lentin  &  Williams,  1976;  H6-1120A,  G27.  7  –  Subtilisphaera  terrula
(Davey, 1974) Lentin & Williams, 1976; emend. Harding, 1986; H6-1120A, O11/1. 8 – Subtilisphaera perlucida (Alberti, 1959) Jain &
Millepied, 1973; H6-300B, L9/2. 9 – Isabelidinium sp.; H6-890B, V8. 10 – Isabelidinium cooksoniae (Alberti, 1959) Lentin & Williams,
1977; H6-1120A, U16/4. 11 – Isabelidinium cooksoniae (Alberti, 1959) Lentin & Williams, 1977; H6-840B, B19/1. 12 – Magallanesium
densispinatum (Stanley, 1965) Quattrocchio & Sarjeant, 2003; H6-1470A, X13. 13, 14 – Magallanesium macmurdoense Wilson, 1967;
H6-300A,  A9.  15,  16  –  Spinidinium  sp.;  H6-1930A,  L14.  17  –  Magallanesium  macmurdoense  Wilson,  1967;  H6-250A,  V29.
18 – Spinidinium echinoideum (Cookson & Eisenack, 1960) Lentin & Williams, 1976; emend. Sverdlove & Habib, 1974; H6-1170B, F13.
19, 20 – Spinidinium echinoideum subsp. rhombicum (Cookson & Eisenack, 1974) Lentin & Williams, 1976; H6-2080B, T12/4.

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Fig. 8. Characteristic and stratigraphically important dinoflagellate cysts of Well Höflein 6. The species name is followed by the sample num-
ber and England Finder coordinates (for localization of the specimen on the slide); all scale bars are 20 µm. 1 – Cerodinium diebelii (Alberti,
1959) Lentin & Williams, 1987; H6-1220B, B21. 2 – Cerodinium obliquipes (Deflandre & Cookson, 1955) Lentin & Williams, 1987;
H6-153B, T21/4. 3, 4 – Cerodinium speciosum subsp. glabrum (Gocht, 1969) Lentin & Williams, 1987; H6-730A, D15. 5 – Cerodinium
speciosum (Alberti, 1959) Lentin & Williams, 1987; H6-1220C, P30. 6 – Palaeoperidinium pyrophorum (Ehrenberg, 1838 ex Wetzel, 1933)
Sarjeant, 1967; emend. Sarjeant, 1967; H6-1470B, V5/1. 7, 8 – Deflandrea phosphoritica Eisenack, 1938; H6-300A, E22/3. 9 – Deflandrea
antarctica Wilson, 1967; H6-50A, B8/2. 10—12 – Deflandrea cygniformis Pöthe de Baldis, 1966; H6-300A, E23. 13 – Trithyrodinium evittii
Drugg, 1967; H6-1220B, C34/2. 14 – Trithyrodinium evittii Drugg, 1967; H6-300A, E2/1. 15 – Trithyrodinium cf. evittii Drugg, 1967;
H6-1120A, U15. 16 – Trithyrodinium robustum Benson, 1976; H6-2480A, H20/3. 17 – Trigonopyxidia ginella (Cookson & Eisenack, 1960)
Downie & Sarjeant, 1965; H6-320B, J7. 18 – Trigonopyxidia ginella (Cookson & Eisenack, 1960) Downie & Sarjeant, 1965; H6-300B,
W11/2. 19 – Palaeotetradinium silicorum Deflandre, 1936; emend. Deflandre & Sarjeant, 1970; H6-320B, A11/3. 20 – Palaeotetradinium
silicorum Deflandre, 1936; emend. Deflandre & Sarjeant, 1970; H6-320B, O42.

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Fig. 9. Characteristic and stratigraphically important dinoflagellate cysts of Well Höflein 6. The species name is followed by the sample
number and England Finder coordinates (for localization of the specimen on the slide); all scale bars are 20 µm. 1 – Lejeunecysta communis
Biffi & Grignani, 1983; H6-1220B, E36. 2 – Lejeunecysta hyalina (Gerlach, 1961) Artzner & Dörhöfer, 1978; emend. Sarjeant, 1984;
H6-2130A, W33. 3 – Lejeunecysta hyalina (Gerlach, 1961) Artzner & Dörhöfer, 1978; emend. Sarjeant, 1984; H6-300A, B35. 4 – Lejeunecysta
hyalina (Gerlach, 1961) Artzner & Dörhöfer, 1978; emend. Sarjeant, 1984; H6-2480A, C5/3. 5 – Manumiella seelandica (Lange, 1969)
Bujak & Davies 1983; emend. Firth, 1987; H6-790A, E29/2. 6 – Manumiella druggii (Stover, 1974) Bujak & Davies, 1983; H6-300B,
Q40/3. 7 – Manumiella? hemmoorensis Marheinecke, 1992; H6-1680A, O27/1. 8 – Manumiella? cf. cretacea (Cookson, 1956) Bujak &
Davies, 1983; H6-300B, A12/4. 9 – Alterbidinium acutulum (Wilson, 1967) Lentin & Williams, 1985; H6-790B, A13. 10 – Alterbidinium
acutulum (Wilson, 1967) Lentin & Williams, 1985; H6-1020B, U31/1. 11 – Phelodinium magnificum (Stanley, 1965) Stover & Evitt,
1978; H6-300B, K25/2. 12 – Phelodinium magnificum (Stanley, 1965) Stover & Evitt, 1978; H6-300A, C11/3. 13 – Phelodinium tricuspe
(Wetzel, 1933) Stover & Evitt, 1978; emend. Lejeune-Carpentier & Sarjeant, 1981; H6-2030B, D20. 14 – Andalusiella polymorpha (Malloy,
1972) Lentin & Williams, 1977; H6-680B, L4. 15 – Andalusiella polymorpha (Malloy, 1972) Lentin & Williams, 1977; H6-1120B, G3.
16 – Palaeocystodinium golzowense Alberti, 1961; H6-300A, D24/3. 17, 18 – Biconidinium reductum (May, 1980) Kirsch, 1991; emend.
Kirsch, 1991; H6-1120B, B32/1. 19 – Kenleyia lophophora Cookson & Eisenack, 1965; H6-300B, J11/4. 20 – Palaeohystrichophora
infusorioides Deflandre, 1935; H6-1930B, Q36/4.

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Fig. 10. Characteristic and strati-
graphically important dinoflagel-
late cysts of Well Höflein 6. The
species  name  is  followed  by  the
sample  number  and  England
Finder  coordinates  (for  localiza-
tion  of  the  specimen  on  the
slide);  all  scale  bars  are  20 µm.
1,  2  –  Odontochitina  porifera
Cookson, 1956; H6-1170A, H2/1.
3  –  Odontochitina  operculata
(Wetzel, 1933) Deflandre & Cookson, 1955; H6-990A, A19. 4 – Odontochitina operculata (Wetzel, 1933) Deflandre & Cookson, 1955;
H6-890A, B11/4. 5 – Muderongia extensiva Duxbury, 1977; H6-2530A, M20. 6 – Muderongia australis Helby, 1987;  emend. Monteil,
1991;  H6-2530B,  W20.  7  –  Muderongia  sp.;  H6-1120A,  G10.  8,  9  –  Pseudoceratium  pelliferum  Gocht,  1957;  emend.  Dörhöfer  &
Davies, 1980; H6-2530A, J23. 10 – Xenascus sarjeantii (Corradini, 1973) Stover & Evitt, 1978; H6-790A, R5/1. 11, 12 – Xenascus
sarjeantii (Corradini, 1973) Stover & Evitt, 1978; H6-1020B, B18. 13 – Xenascus sp.; H6-1930B, O14/2. 14 – Xenascus   cf. asperatus
Stover & Helby, 1987; H6-1930B, O14/2. 15 – Xiphophoridium alatum (Cookson & Eisenack, 1962) Sarjeant, 1966; emend.

Sarjeant,

1966; H6-20A, Y28.

tion not lower than Lower Eocene and the Paleocene/Eocene
boundary probably may be present below 300 m depth
(Fig. 2). Associated with the Paleocene-Eocene thermal maxi-
mum (PETM) warming is a > 2.5 % negative carbon isotope
(

C) excursion (CIE) measured on terrestrial and marine

sedimentary carbon (Kennett & Stott 1991; Koch et al. 1992;
Thomas et al. 2002; Pagani et al. 2006). The position of the
CIE termination is correlated to the concomitant LO of the di-

nocyst species Apectodinium augustum, which has only been
recorded from the CIE (Bujak & Brinkhuis 1998; Steurbaut et
al. 2003; Sluijs et al. 2007; Hofmann et al. 2011; Speijer et al.
2012). In Well Höflein 6, Apectodinium augustum occurs at
the depth of 1520 m which indicates equivalence to the upper-
most Paleocene PETM-interval for this horizon directly below
the Paleocene/Eocene boundary in the lower thrust unit B at
the base of the Greifenstein Formation (Fig. 2).

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Fig. 11. Characteristic and strati-
graphically important dinoflagel-
late cysts of Well Höflein 6. The
species name is followed by the
sample  number  and  England
Finder  coordinates  (for  localiza-
tion  of  the  specimen  on  the
slide);  all  scale  bars  are  20 µm.
1 – Membranilarnacia? tenella
Morgenroth,  1968;  H6-300A,
A14.  2  –  Membranilarnacia?
tenella

Morgenroth,

1968;

H6-940A, P11/3. 3 – Membranilarnacia? tenella Morgenroth, 1968; H6-1120A, D16/2. 4—6 – Thalassiphora delicata Williams & Downie,
1966; emend. Eaton, 1976; H6-1370A, N9. 7 – Thalassiphora pelagica (Eisenack, 1954) Eisenack & Gocht, 1960; emend.  Benedek &
Gocht, 1981; H6-300B, C4/1. 8 – Thalassiphora inflata Heilmann-Clausen in Thomsen & Heilmann-Clausen, 1985;  H6-2530B, S14/3.
9  – Trabeculidium quinquetrum Duxbury, 1980; H6-1020A, E32/3.  10  – Trichodinium ciliatum (Gocht, 1959)  Eisenack & Klement,
1964; H6-2530B, Q32/2. 11 – Riculacysta amplexa Kirsch, 1991; H6-1470B, M20/4. 12 – Riculacysta amplexa Kirsch, 1991; H6-1470B,
J35/3. 13 – Paralecaniella indentata (Deflandre & Cookson, 1955) Cookson & Eisenack, 1970; emend. Elsik, 1977; H6-1270A, T40/2.
14 – Paralecaniella indentata (Deflandre & Cookson, 1955) Cookson & Eisenack, 1970; emend. Elsik, 1977; H6-1270B, N20. 15 – Para-
lecaniella indentata (Deflandre & Cookson, 1955) Cookson & Eisenack, 1970; emend. Elsik, 1977; H6-1420A, D31.

Carpatella cornuta is a typical Danian dinoflagellate spe-

cies (e.g. Williams et al. 2004). In the Northern Hemisphere
at the El-Kef GSSP (Global Boundary Stratotype Section
and Point) section, Brinkhuis et al. (1998) documented that
the lowest global occurrence of Carpatella cornuta is slight-
ly above the Cretaceous/Paleogene (K/Pg) boundary. In the

Southern Hemisphere, Carpatella cornuta is recorded above
the K/Pg boundary, two sporadic occurrences of Carpatella
cornuta below the boundary are interpreted as the result of
intense bioturbation (Ferrow et al. 2011). This Danian species
and Senoniasphaera inornata are reported to occur slightly
above the boundary both in the Northern and Southern Hemi-

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Fig. 12.

Characteristic

and

stratigraphically  important  di-
noflagellate  cysts  of  Well  Höf-
lein 6.  The  species  name  is
followed by the sample number
and England Finder coordinates
(for localization of the specimen
on  the  slide);  all  scale  bars  are
20 µm. 1  –  Oligosphaeridium
complex (White, 1842) Davey &
Williams, 1966; H6-300B, Q42.
2 – Oligosphaeridium complex (White, 1842) Davey & Williams, 1966; H6-2530B, F7. 3, 6 – Hystrichosphaerina schindewolfii Alberti,
1961;  H6-1170B,  E12.  4,  5  –  Circulodinium  distinctum  (Deflandre  &  Cookson,  1955)  Jansonius,  1986;  H6-300A,  C27.  7  –  Pervo-
sphaeridium pseudhystrichodinium (Deflandre, 1937) Yun Hyesu, 1981; emend. Davey, 1969; H6-300A, D27/3. 8 – Hystrichodinium pulchrum
Deflandre, 1935; H6-790A, H18/1. 9 – Hystrichodinium pulchrum subsp. areatum Marheinecke, 1992; H6-1730A, W35/1. 10 – Hystricho-
sphaeridium tubiferum subsp. brevispinum (Davey & Williams, 1966) Lentin & Williams, 1973; emend. Marheinecke, 1992; H6-790B, A4/2.
11 – Hystrichosphaeridium tubiferum (Ehrenberg, 1838) Deflandre, 1937; emend. Davey & Williams, 1966; H6-790B, R32. 12 – Hystricho-
sphaeridium recurvatum (White, 1842) Lejeune-Carpentier, 1940; H6-940A, W25. 13 – Surculosphaeridium? basifurcatum Yun Hyesu,
1981; H6-1010B, F15/2. 14 – Spiniferites cf. bulloideus (Deflandre & Cookson, 1955) Sarjeant, 1970; H6-200A, Q23. 15 – Spiniferites
membranaceus (Rossignol, 1964) Sarjeant, 1970; H6-890B, L37.

spheres by Williams et al. (2004). In West Greenland, N

hr-

Hansen  &  Dam  (1997)  recognized  the  LO  of  Senonia-
sphaera  inornata  in  the  K/Pg  boundary  clay  layer.  In  the
Gams Basin, Austria, Senoniasphaera inornata and Damas-
sadinium  californicum  occur  first  directly  above  the  K/Pg
boundary  and  Carpatella  cornuta  occurs  30 cm  above

(Mohamed  et  al.  2012).  In  the  upper  thrust  unit  A  of  Well
Höflein 6, the FO of Damassadinium californicum, also typ-
ical for the Danian (e.g. Williams et al. 2004), is recorded at
the  depth  of  630 m.  Both,  Carpatella  cornuta  and  Senonia-
sphaera inornata are recorded at the depth of 680 m suggest-
ing  the  presence  of  the  K/Pg  boundary  directly  below  the

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depth of 680 m (Fig. 2), within the Altlengbach Formation. In
the  lower  thrust  unit B,  Carpatella  cornuta  is  not  recorded.
The  FOs  of  Damassadinium  californicum  and  Senonia-
sphaera  inornata  are  recorded  at  the  depth  of  1630 m  and
1580 m  respectively,  suggesting  the  position  of  the  K/Pg
boundary directly below the depth of 1630 m (Fig. 2).

The FO of Manumiella seelandica is recorded at the Cam-

panian-Maastrichtian boundary in the southern mid latitudes
and in the Upper Maastrichtian in the northern mid latitudes
(e.g.  Williams  et  al.  2004).  In  Well  Höflein 6,  the  FO  of
Manumiella  seelandica  occurs  at  the  depth  of  990 m  in  the
upper  thrust  unit A  (Perneck  Formation)  and  at  1780 m
depth  in  the  lower  thrust  unit B  (Altlengbach  Formation)
(Fig. 2). The occurrence of this species at this level indicates
a  stratigraphic  position  not  lower  than  Maastrichtian.  The
highest occurrence of Palaeohystrichophora infusorioides is
recorded  directly  above  the  Campanian-Maastrichtian
boundary  in  the  southern  mid  latitudes  and  occurs  at  the
depth of 1930 m in Well Höflein 6. The most significant bio-
stratigraphic event within this interval is the LO of Odonto-
chitina  porifera.  The  highest  occurrence  of  this  species  is
restricted  to  the  Campanian  in  the  northern  mid  latitudes
(e.g. Williams et al. 2004). Kirsch (1991) reported this species
from the Lower—Middle Campanian in the Helvetic and Ultra-
helvetic realm of Germany and Austria. In Well Höflein 6 the
LO  of  Odontochitina  porifera  is  recorded  at  the  depth  of
1120 m  in  the  upper  thrust  unit A  (Röthenbach  Subgroup)
and  1930 m  in  the  lower  thrust  unit B  (Altlengbach  Forma-
tion?). Therefore, a Campanian age can be inferred for these
depth intervals, and formations are either diachronous or for-
mation  interpretation  of  the  lower  thrust  unit  has  to  be  re-
fined.  Corradinisphaeridium  horridum  occurs  in  three
samples (depth of 1070, 2030, 2080 m) (Fig. 2). This species
was originally reported by Corradini (1973) from the Campa-
nian  of  northern  Italy,  from  the  type  Campanian  (Charente,
France) by Masure (1986) and from mid-Campanian strata in
southern  Germany  by  Kirsch  (1991).  Schi

ler & Wilson

(2001) regard this species as an important Middle—Upper
Campanian marker with a LO close to the Campanian-Maas-
trichtian boundary. Based on these data the Campanian-
Maastrichtian boundary can be positioned above the depth of
1070 m and 1930 m in the Well Höflein 6.

Further biostratigraphically important events are recorded

at  the  boundary  of  the  Röthenbach  Subgroup  (Altlengbach
Formation according to OMV internal report) to the underly-
ing  Wolfpassing  Formation,  namely  the  LOs  of  Muderongia
extensiva,  Muderongia  australis,  Leberidocysta?  laticaudata
and  Pseudoceratium  pelliferum  at  2530 m  depth,  and  the
FOs  of  Areoligera  coronate,  Chatangiella  ditissima  and
Cerodinium  diebelii  at  2480 m  depth  (Fig. 2).  Kirsch  (1991)
reported the FO of Areoligera coronata from the Lower Cam-
panian  in  southern  Germany.  Previous  studies  demonstrated
that the FO of the genus Areoligera is not recorded below the
Campanian  (e.g.  Williams  &  Bujak  1985;  Williams  et  al.
2004).  The  FO  of  Cerodinium  diebelii  was  recorded  as  a
Maastrichtian event in central and northern Europe by Górka
(1963), Wilson (1974), Kirsch (1991) and Smelror & Riegraf
(1996),  from  the  Lower  Maastrichtian  of  West  Greenland
(N

hr-Hansen 1996) and NW Germany (Marheineck 1992),

and  from  the  Upper  Campanian  in  the  North  Sea  area  by
Costa  &  Davey  (1992).  C.  diebelii  was  also  found  in  the
Maastrichtian  of  the  Outer  Carpathian  Flysch  Zone  by
Skupien  &  Mohamed  (2008),  and  probably  occurs  before  in
the  Late  Campanian  (see  Skupien  et  al.  2009).  Schi

ler &

Wilson  (2001)  interpreted  this  species  as  an  important  bio-
stratigraphic  marker  situated  directly  below  the  Campanian-
Maastrichtian  boundary.  Leberidocysta?  laticaudata  was
recorded in the Santonian of the USSR (Vozzhennikova 1967)
and NW Germany (Yun Hyesu 1981). Thus, in the Höflein 6
Well  the  lower  boundary  of  the  Campanian  occurs  between
2480 and 2530 m depth.

The succession of the Wolfpassing Formation below the

Lower  Campanian  part  of  thrust  unit B  in  Well  Höflein 6  is
regarded as older than Santonian, although the age evidence
is not straightforward and rather contradictory. Muderongia
extensiva  was  recorded  in  the  Lower  Valanginian—Lower
Hauterivian  in  England  (Duxbury  1977),  Valanginian-basal
Hauterivian in Denmark (Heilmann-Clausen 1987), Valangin-
ian/Hauterivian—Upper Aptian in Norway (Thusu 1978), Up-
per  Berriasian—Upper  Valanginian  in  the  Gulf  of  Mexico
(Riley  &  Fenton  1984)  and  Upper  Barremian  in  France
(Srivastava 1984). Muderongia australis was recorded in the
Hauterivian—Barremian in Australia (Helby 1987), basal Hau-
terivian in NW Germany (Prauss 1990) and Lower Barremian
in  east  Greenland  (N

hr-Hansen 1993). Stover et al. (1996)

documented  that  the  range  of  this  species  is  Hauterivian—
Aptian.  On  the  other  hand,  the  species  Palaeoperidinium
pyrophorum  and  Palaeotetradinium  silicorum  (sample 2530;
Fig. 8.19—20) have their FO probably in the Cenomanian, and
Surculosphaeridium  longifurcatum  (samples 2560,  2530)  has
its  FO  in  the  Middle  Albian  according  to  some  authors  (e.g.
Skupien  et  al.  2009),  however,  it  was  also  recorded  in  older
sediments, e.g. Gedl (2007) in the Aptian to Early Campanian
in  Poland.  Thus,  in  the  Well  Höflein 6  the  interval  from
2530 m to 2565 m can most probably be correlated to the in-
terval from the Hauterivian to the Cenomanian. Because of the
presence of the genus Muderongia, a Hauterivian—Barremian to
Aptian age is most likely, which is in accordance with the gen-
eral age assignment for this formation (e.g. Sauer et al. 1992).

Reworking and downfall materials

A few specimens of Cretaceous species such as Florentinia

aculeata,  Tanyosphaeridium  xanthiopyxides,  Cannosphae-
ropsis  utinensis,  Chatangiella  spp.,  Palaeotetradinium  sili-
corum  and  Surculosphaeridium  longifurcatum  are  recorded
in the Paleocene samples indicating a high rate of reworking
(electronic Appendix B, C). The presence of single specimen
of  Areosphaeridium  diktyoplokum  (Fig. 3.13,14)  and  Cero-
dinium diebelii at 2530 m depth are probably downfall from
the  upper  succession.  A  few  specimens  of  Cretaceous  spe-
cies such as Cannosphaeropsis utinensis, Chatangiella spp.,
Tanyosphaeridium  xanthiopyxides,  Palaeotetradinium  sili-
corum,  Surculosphaeridium?  basifurcatum  and  Xenascus
sarjeantii are recorded in the Paleocene samples at the depth
of ca. 1520 m indicating again increased reworking (electronic
Appendix B, C). The occurrence of one damaged specimen of

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GEOLOGICA CARPATHICA, 2013, 64, 3, 209—230; Electronic Supplement, i—xii

Odontochitina porifera at the depth of 890 m indicates a re-
working from older sediments.

The FO of Glaphyrocysta exuberans at the depth of

1730 m  may  be  due  to  downhole  contamination  since  this
species is only known from the Paleocene and Eocene (e.g.
Köthe & Piesker 2007). The LO of the Paleocene taxa Car-
patella  cornuta,  Damassadinium  californicum  are  recorded
at the depth of 250 m and the LO of Senoniasphaera inornata
is recorded in 20 m depth, indicating a reworking of Paleo-
cene  into  Eocene  strata.  A  few  specimens  (one  or  two  per
sample) of Cretaceous species such as Chatangiella ditissima,
Chatangiella madura, Chatangiella spp., Cannosphaeropsis
utinensis,  Florentinia  aculeata,  Manumiella?  cf.  cretacea,
Surculosphaeridium  longifurcatum,  Dinogymnium  denticu-
latum,  Tanyosphaeridium  xanthiopyxides,  Palaeotetradinium
silicorum,  Trigonopyxidia  ginella,  Stanfordella  fastigiata
and Xiphophoridium alatum are present in the Eocene sam-
ples of the lower and upper thrust units indicating a high de-
gree of reworking (electronic Appendix B, C).

Stratigraphic correlations of Flysch units from

borehole to outcrop

The dinoflagellate-based ages as interpreted from the cut-

tings  samples  of  the  Höflein 6  Well  compare  well  with  age
data  derived  from  the  corresponding  lithostratigraphic  units
from the succession of the Wienerwald area. The Wolfpassing
Formation at the base of the lower thrust unit B in Höflein 6
(2490—2530 m)  gives  indications  for  a  Hauterivian  to  Aptian
age, in accordance with published data from the type section
of the Wolfpassing Formation of the Wienerwald area (mainly
Barremian—Aptian; Grün et al. 1972). Stratigraphically, a gap
between Aptian and overlying Campanian strata may indicate
another thrust or fault plane separating the Wolfpassing For-
mation  from  the  overlying  younger  strata.  The  Röthenbach
Subgroup from the upper thrust unit A (1017—1210 m) ranges
from the Campanian up to the basal Maastrichtian according
to  our  dinocyst  data.  Similar  Campanian  ages  are  reported
from  Egger  &  Schwerd  (2008)  from  Bavaria,  Germany.  The
Perneck Formation from the upper thrust unit A (785—1017 m)
has an Early Maastrichtian age according to our data in con-
trast to the Late Campanian age reported from Upper Austria
and  Bavaria  (Egger  1993;  Egger  &  Schwerd  2008).  Further
biostratigraphic data is needed to evaluate whether the forma-
tion is diachronous from west to east within the RFZ, or if di-
noflagellate  biostratigraphic  datums  and  problems  due  to
reworking and downfall result in an imprecise biostratigraphic
result  for  Well  Höflein 6.  Some  uncertainties  probably  also
exist with the lithostratigraphic interpretation of the Höflein 6
borehole as the Perneck Formation, normally only up to 50 m
thick, attains a (non-dip corrected) thickness of 120 m.

The Altlengbach Formation is present both in the upper

unit A (322—785 m) and in the lower unit B (1520—2470 m).
Within the upper unit, dinoflagellate ages compare well with
the  known  stratigraphic  ages  of  Maastrichtian  to  Paleocene
of  the  Altlengbach  Formation  (up  to  NP9;  Schnabel  1992;
Egger 1995). In the lower unit B, some ambiguity seems to
exit  about  the  attribution  of  the  lower  part  of  the  interval,

mainly the Campanian part, to the Altlengbach Formation as
is also indicated by the high thickness of 940 m. At least the
lower part of this Campanian section interval from 2470 to
1930 m may be alternatively attributed to the Röthenbach
Subgroup (and the Perneck Formation on top) in accordance
with the succession within the upper thrust unit A.

The Greifenstein Formation of Well Höflein 6 straddles the

Paleocene-Eocene  boundary  and  ranges  up  into  the  Lower/
Middle  Eocene.  This  correlates  well  with  data  from  Hekel
(1968:  NP9  to  NP13,  Thanetian—Ypresian)  and  chronostrati-
graphic  ages  summarized  by  Schnabel  (1992)  and  Faupl
(1996)  for  the  Greifenstein  Formation  of  the  Greifenstein
Nappe.

Conclusions

Despite the fact that only cuttings samples from Well Höf-

lein 6 within the Rhenodanubian Flysch Zone have been inves-
tigated a concise and detailed biostratigraphic succession and
zonation based on organic-walled dinoflagellates is possible.

In the upper thrust unit A, an Eocene age is indicated in the

upper part of the Greifenstein Formation. A Paleocene age is
indicated for the upper part of the Altlengbach Formation (and
in the overlying lowermost Greifenstein Formation) between
the depths of ca. 680 m and ca. 322 m. A Maastrichtian age is
interpreted for the uppermost Röthenbach Subgroup, the Per-
neck Formation (Oberste Bunte Schiefer of OMV internal re-
ports) and the lowermost Altlengbach Formation, between the
depths of ca. 1020 m and ca. 730 m. A Campanian age is in-
ferred for the Röthenbach Subgroup, between the depths of ca.
1210 m and ca. 1017 m.

In the lower thrust unit B, an Eocene age is inferred for the

upper part of the Greifenstein Formation. A Paleocene age is
recorded in the uppermost Altlengbach Formation and in the
lowermost Greifenstein Formation between the depths of ca.
1630 m and ca. 1520 m. A Maastrichtian age is inferred for
the Altlengbach Formation between the depths of ca. 1880 m
and ca. 1680 m. A Campanian age is inferred between the
depths of ca. 2480 m and ca. 1930 m (corresponding mainly
to the lowermost part of the Altlengbach Formation and
probably parts of the Röthenbach Subgroup).

The Wolfpassing Formation of inferred Early Cretaceous

age is only reported from the lower thrust unit B (see Table 1),
between the depths of ca. 2561 m and 2470 m. A stratigraphic
gap probably indicates the presence of a thrust or fault be-
tween the Wolfpassing Formation and the overlying Campa-
nian strata.

Acknowledgments:  We  express  our  thanks  to  Werner  Piller
(Graz University, Austria) who helped and offered us the op-
portunity  to  prepare  the  palynological  slides  of  the  studied
samples  in  the  palynological  laboratory  of  the  Institute  of
Earth Sciences at Graz University. We thank OMV E & P, in
particular Philipp Strauss, for providing samples and access to
unpublished  reports,  and  Roman  Sauer  from  OMV  Lab  for
well  information.  Andrea  Schicker  (University  of  Vienna)  is
thanked for help in sampling. This paper benefited significantly
from critical reviews by Przemysław Gedl and Petr Skupien.

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GEOLOGICA CARPATHICA, 2013, 64, 3, 209—230; Electronic Supplement, i—xii

References

Auffret J.P. & Gruas-Cavagnetto C. 1975: Les formations paléo-

nes sous-marines de la Manche orientale données pa-

lynologiques. Bull. Soc. Geol. France 7e sér. 17, 5, 641—655.

Brinkhuis H., Bujak J.P., Versteegh G.J.M. & Visscher H. 1998:

Dinoflagellate-based sea surface temperature reconstructions
across the Cretaceous-Tertiary boundary. Palaeogeogr. Palaeo-
climatol. Palaeoecol. 141, 67—83.

Bujak J.P. & Brinkhuis H. 1998: Global warming and dinocyst

changes  across  the  Paleocene/Eocene  epoch  boundary.  In:
Aubry  M.-P.  et  al.  (Eds.):  Late  Paleocene—Early  Eocene  cli-
matic  and  biotic  events  in  the  marine  and  terrestrial  records.
Univ. Press, New York, Columbia, 277—295.

Caro Y. 1973: Contribution à la connaissance des dinoflagellés du

Paléocène-Eocène inférieur des Pyrenées espagnoles.

Rev. Esp.

Micropaleont. 5, 3, 329—372.

Châteauneuf J.J. 1980: Palynostratigraphie et paléoclimatologie de

l’Eoc

ne supérieur et de l’Oligoc

ne du Bassin de Paris. Mém.

Bureau Rech. Géol. Min. (BRGM) 116, 1—360.

Corradini D. 1973: Non-calcareous microplankton from the Upper

Cretaceous of the northern Apennines. Boll. Soc. Paleont. Ital.
11, 119—197.

Costa L.I. & Davey R.J. 1992: Dinoflagellate cysts of the creta-

ceous system. In: Powell A.J. (Ed.): A stratigraphic index of
dinoflagellate cysts. BMS Occasional Publ. Ser., Chapman &
Hall, London, 99—155.

Costa L.I. & Martini E. 1981: Dinoflagellaten-Gemeinschaften aus

dem höheren Eozän der Bohrung Kressenberg 7 (Bayern) und
Korrelation zu den Nannoplankton Zonen. Geol. Bavaria 82,
315—318.

De Coninck J., Geets S. & Willems W. 1983: The Mont-Heribu

Member: Base of the Ieper Formation in the Belgian Basin.
Tertiary Res. 5, 2, 83—104.

Duxbury S. 1977: A palynostratigraphy of the Berriasian to Barre-

mian of the Speeton Clay of Speeton, England. Palaeonto-
graphica, Abt. B 160, 1—3, 17—67.

Edwards L.E. 1989: Dinoflagellate cysts from the Lower Tertiary

Formations, Haynesville Cores, Richmond County, Virginia
Geology and Paleontology of the Haynesville Cores – North-
eastern Virginia Coastal Plain. U.S. Geol. Surv. Prof. Pap.,
1489-C, United States Government Printing Office, Washington,
1—23.

Egger H. 1993: Zur Nannoplankton-Stratigraphie der Seisenburger

Schichten (Coniac? – frühes Campan) in der Rhenodanubis-
chen Flyschzone (Ostalpen) östlich des Inn. Zitteliana 20, 59—65.

Egger H. 1995: Die Lithostratigraphie der Altlengbach-Formation

und der Anthering-Formation im Rhenodanubischen Flysch (Ost-
alpen, Penninikum). Neu. Jb. Geol. Paläont. Abh. 196, 69—91.

Egger H. & Schwerd K. 2008: Stratigraphy and sedimentation rates

of Upper Cretaceous deep-water systems of the Rhenodanubian
Group (Eastern Alps, Germany). Cretaceous Research 29,
405—416.

El Beialy S.Y. & Shahin A.M. 1990: Planktonic foraminifera and

dinoflagellate cysts across the Cretaceous/Tertiary boundary in
the Nile Delta area, Egypt. Neu. Jb. Geol. Paläont. Abh. 180, 1,
117—137.

Faupl P. 1996: Tiefwassersedimente und tektonischer Bau der Flysch-

zone des Wienerwaldes. Ber. Geol. Bundesanst. 33 (1996), A2:
1—32.

Fechner G. & Mohr B. 1988: Early Eocene spores, pollen and mi-

croplankton assemblages from the Fehmarn Island, Northern
Germany. Tertiary Res. 9, 1—4, 147—168.

Fensome R.A., Taylor F.J.R., Norris G., Sarjeant W.A.S., Wharton

D.I. & Williams G.L. 1993: A classification of living and fos-
sil dinoflagellates. Micropaleont., Spec. Publ. 7, 1—351.

Fensome R.A., MacRae R.A. & Williams G.L. 2008: DINOFLAJ2,

Version 1. Amer. Assoc. Stratigr. Palynologists, Data Ser. No. 1,
1—939.

Ferrow E., Vajda V., Bender Koch C., Peucker-Ehrenbrink B. &

Willumsen P. 2011: Multiproxy analysis of a new terrestrial
and a marine Cretaceous-Paleogene (K-Pg) boundary site from
New Zealand. Geochim. Cosmochim. Acta, 657—672.

Gedl P. 2007: Organic-walled dinoflagellate cysts from some Juras-

sic and Cretaceous strata of the Grajcarek Unit at Haluszowa,
Pieniny Klippen Belt (West Carpathians, Poland). Stud. Geol.
Pol. 127, 101—117.

Górka H. 1963: Coccolithophoridés, dinoflagellés, hystricho-

sphaeridés et microfossiles incertae sedis du Crétacé supérieur
de Pologne. Acta Palaeont. Pol. 8, 1, 1—83, 1—11.

Grün W., Kittler G., Lauer G., Papp A. & Schnabel W. 1972: Studien

in der Unterkreide des Wienerwaldes. Jb. Geol. Bundesanst.
115, 103—186.

Hamilton W., Wagner L. & Wessely G. 2000: Oil and gas in Aus-

tria. Mitt. Österr. Geol. Gesell. 92 (1999), 235—262.

Heilmann-Clausen C. 1987: Lower Cretaceous dinoflagellate bio-

stratigraphy in the Danish Central Trough. Danmarks Geol.
Unders. A17, 1—89.

Heilmann-Clausen C. & Costa L.I. 1989: Dinoflagellate zonation of

the uppermost Paleocene? to Lower Miocene in the Wurster-
heide Research Well, NW Germany. Geol. Jb. 111, 431—521.

Hekel H. 1968: Nannoplanktonhorizonte und tektonische Strukturen

in der Flyschzone nördlich von Wien (Bisamberg). Jb. Geol.
Bundesanst. 111, 293—338.

Helby R. 1987: Muderongia and related dinoflagellates of the latest

Jurassic to Early Cretaceous of Australasia. In: Jell P.A. (Ed.):
Studies in Australian Mesozoic palynology. Mem. Assoc. Austra-
lasian Palaeont. 4, 297—336.

Hofmann C., Mohamed O. & Egger H. 2011: A new terrestrial pa-

lynoflora from the Palaeocene/Eocene boundary in the north-
western Tethyan realm (St. Pankraz, Austria). Rev. Palaeobot.
Palynol. 16, 295—310.

Islam M.A. 1983: Dinoflagellate cyst taxonomy and biostratigraphy

of the Eocene Bracklesham Group in southern England. Micro-
paleont. 29, 328—353.

Kennett J.P. & Stott L.D. 1991: Abrupt deep-sea warming, palae-

oceanographic changes and benthic extinctions at the end of
the Palaeocene. Nature 353, 225—229.

Kirsch K.H. 1991: Dinoflagellatenzysten aus der Oberkreide des

Helvetikums und Nordultrahelvetikums von Oberbayern.
Münchner Geowiss. Abh., Reihe A, Geol. Paläont. 22, 1—306.

Kirsch K.-H. 2000: Dinoflagellatenzysten aus der höheren Oberkre-

ide des Rhenodanubischen Flysches. 1. Kalkgrabenschichten
vom Schlierse/Oberbayern. Mitt. Bayer. Staatslg. Paläont.
Hist. Geol. 40, 3—9.

Kirsch K.H. 2003: Dinoflagellatenzysten-Zonierung der höheren

Unterkreide des Rhenodanubischen Flysches. Zitteliana A43,
143—158.

Koch P.L., Zachos J.C. & Gingerich P.D. 1992: Correlation between

isotope records in marine and continental carbon reservoirs near
the Palaeocene/Eocene boundary. Nature 358, 319—322.

Köthe A. & Piesker B. 2007: Stratigraphic distribution of Paleogene

and Miocene dinocysts in Germany. Rev. Paléobiol. 26, 1, 1—39.

Marheinecke U. 1992: Monographie der Dinozysten, Acritarcha und

Chlorophyta des Maastrichtium von Hemmoor (Niedersachsen).
Palaeontographica, Abt. B 227, 1—6, 1—173.

Masure E. 1986: Corradinisphaeridium, nouveau genre de di-

noflagellés du Sénonien d’Italie et de France. Rev. Micro-
paléont. 29, 109—119.

Mattern F. 1999: Mid-Cretaceous basin development, paleogeogra-

phy, and paleogeodynamics of the western Rhenodanubian
Flysch (Alps). Z. Dtsch. Geol. Gesell. 150, 89—132.

226

MOHAMED and WAGREICH

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 209—230; Electronic Supplement, i—xii

Mohamed O., Piller W.E. & Egger H. 2012: The dinocyst record

across the Cretaceous/Palaeogene boundary of a bathyal mid-
latitude Tethyan setting: Gosau Group, Gams Basin, Austria.
Cretaceous Research 35, 143—168.

hr-Hansen H. 1993: Dinoflagellate cyst stratigraphy of the Bar-

remian to Albian, Lower Cretaceous, north-east Greenland.
Gr

nlands Geol. Unders. 166, 1—171.

hr-Hansen H. 1996: Upper Cretaceous dinoflagellate cyst strati-

graphy, onshore west Greenland. Gr

nlands

Geol. Unders.

170, 1—104.

hr-Hansen H. & Dam G. 1997: Palynology and sedimentology

across a new marine Cretaceous-Tertiary boundary section on
Nuussuaq, West Greenland. Geology 25, 851—854.

Pagani M., Pedentchouk N., Huber M., Sluijs A., Schouten S.,

Brinkhuis H., Sinninghe Damste’ J.S., Dickens G.R. & Expe-
dition-Scientists 2006: Arctic hydrology during global warm-
ing at the Palaeocene-Eocene thermal maximum. Nature 442,
671—675.

Prauss M. 1990: Palynofazielle Untersuchungen in der hodsoni-Zone

(Ober-Bathonium, Dogger) von Lechstedt bei Hildesheim. Geol.
Jb., A 121, 275—291.

Riley L.A. & Fenton J.P.G. 1984: Palynostratigraphy of the Berria-

sian to Cenomanian sequence at Deep Sea Drilling Project Site
535,  Leg  77,  southeastern  Gulf  of  Mexico.  In:  Buffler  R.T.,
Schlager  W.,  Bowdler  J.L.,  Cotillon  P.H.,  Halley  R.B.,
Kinoshita H., Magoon L.B., III, McNulty Ch.L., Patton J.W.,
Premoli Silva I., Suarez O.A., Testarmata M.M., Tyson R.V.,
Watkins  D.K.  &  Pisciotto  K.A.  (Eds):  Initial  reports  of  the
Deep  Sea  Drilling  Project,  covering  Leg  77  of  the  cruises  of
the drilling vessel Glomar Challenger. Ft. Lauderdale, Florida
to San Juan, Puerto Rico, December 1980-February 1981. Texas
A & M University, Ocean Drilling Program, College Station,
TX, U.S. 77, 675-690.

Sauer R., Seifert P. & Wessely G. 1992: Guidebook to Excursions

in the Vienna Basin and the adjacent Alpine-Carpathian
Thrustbelt in Austria. Excursion I. Mitt. Österr. Geol. Gesell.
85, 100—137.

Schi

ler P. & Wilson G.J. 2001: Dinoflagellate biostratigraphy

around the Campanian-Maastrichtian boundary at its type sec-
tion  (Tercis  Quarry,  southwest  France).  In:  Odin  G.S.  (Ed.):
The  Campanian-Maastrichtian  boundary:  characterisation  and
correlation  from  Tercis  (Landes,  SW  France)  to  Europe  and
other continents. IUGS Spec. Publ., Monograph Ser., 36, De-
velopments  in  Palaeontology  and  Stratigraphy,  19,  Elsevier
Science Publishers, Amsterdam, 233—246.

Schnabel W. 1992: New data on the Flysch Zone of the Eastern

Alps in the Austrian sector and new aspects concerning the
transition to the Flysch Zone of the Carpathians. Cretaceous
Research 13, 405—419.

Skupien P. & Mohamed O. 2008: Campanian to Maastrichtian pa-

lynofacies and dinoflagellate cysts of the Silesian Unit, Outer
Western Carpathians, Czech Republic. Bull. Geosci. 83, 2,
207—224.

Skupien P., Bubík M., Švábenická L., Mikuláš R., Vašíček Z. &

Matýsek D. 2009: Cretaceous Oceanic Red Beds in the Outer
Western Carpathians of the Czech Republic. In: Hu X., Wang
Ch., Scott R.W., Wagreich M. & Jansa L. (Eds.): Cretaceous
Oceanic Red Beds: Stratigraphy, composition, origins, and pa-
leoceanographic and paleoclimatic significance. SEPM Spec.
Publ. 91, 99—109.

Sluijs A., Brinkhuis H., Schouten S., Bohaty S.M., John C.M.,

Zachos J.C., SinningheDamste’ J.S., Crouch E.M. & Dickens
G.R. 2007: Environmental precursors to light carbon input at
the Paleocene/Eocene boundary. Nature 450, 1218—1221.

Smelror M. & Riegraf W. 1996: Dinoflagellate cyst distribution at

the lower-upper Campanian boundary beds in Westphalia
(NW-Germany). Newslett. Stratigr. 34, 109—126.

Speijer R.P., Scheibner C., Stassen P. & Morsi A.M.M. 2012: Re-

sponse of marine ecosystems to deep-time global warming: a
synthesis of biotic patterns across the Paleocene-Eocene ther-
mal maximum (PETM). Austrian J. Earth Sci. 105, 1, 6—16.

Srivastava S.K. 1984: Barremian dinoflagellate cysts from south-

eastern France. Cahiers Micropaléont. 1984—2, 1—90.

Steurbaut E., Magioncalda R., Dupuis C., Van Simaeys S., Roche E.

& Roche M. 2003: Palynology, paleoenvironments, and organic
carbon isotope evolution in lagoonal Paleocene-Eocene bound-
ary settings in north Belgium. In: Wing S.L. et al. (Eds.):
Causes and consequences of globally warm climates in the
Early Paleogene. Geol. Soc. Amer., Spec. Pap. 369, 291—317.

Stover L.E., Brinkhuis H., Damassa S.P., De Verteuil L., Helby

R.J.,  Monteil  E.,  Partridge  A.D.,  Powell  A.J.,  Riding  J.B.,
Smelror  M.  &  Williams  G.L.  1996:  Mesozoic-Tertiary  di-
noflagellates, acritarchs and prasinophytes. In: Jansonius J. &
McGregor  D.C.  (Eds.):  Palynology:  principles  and  applica-
tions. Amer. Assoc. Stratigr. Palynol. Found. 2, 64—750.

Thomas D.J., Zachos J.C., Bralower T.J., Thomas E. & Bohaty S.

2002: Warming the fuel for the fire: Evidence for the thermal
dissociation of methane hydrate during the Paleocene-Eocene
thermal maximum. Geology 30, 1067—1070.

Thusu B. (Ed.) 1978: Distribution of biostratigraphically diagnostic

dinoflagellate cysts and miospores from the Northwest Euro-
pean continental shelf and adjacent areas. Continental Shelf
Institute (Trondheim, Norway), Publ. 100, 1—111.

Vozzhennikova T.F. 1967: Fossil peridineae from Jurassic, Creta-

ceous and Paleogene deposits of the USSR. [Iskopaemye pe-
ridinei Yurskikh, Melovykh i Paleogenovykh otlozheniy
SSSR.] Izdatelstvo Nauka, Moscow, 1—347.

Wagreich M. 2008: Lithostratigraphic definition and depositional

model of the Hütteldorf Formation (Upper Albian—Turonian,
Rhenodanubian Flysch Zone, Austria). Austrian J. Earth Sci.
101, 70—80.

Williams G.L. & Bujak J.P. 1985: Mesozoic and Cenozoic di-

noflagellates. In: Bolli H.M., Saunders J.B. & Perch-Nielsen
K. (Eds.): Plankton stratigraphy. Cambridge University Press,
New York, 847—964.

Williams G.L., Brinkhuis H., Pearce M.A., Fensome R.A. & Weegink

J.W. 2004: Southern Ocean and global dinoflagellate cyst
events compared: index events for the late Cretaceouse-Neo-
gene. Proc. Ocean Drilling Progr., Scientific Res. 189, 1—98.

Wilson G.J. 1974: Upper Campanian and Maastrichtian dinoflagel-

late cysts from the Maastricht region and Denmark. Ph.D. Dis-
sert., Nottingham Univ., 1—608.

Wilson G.J. 1988: Paleocene and Eocene dinoflagellate cysts from

Waipawa, Hawkes Bay, New Zealand. New Zealand Geol.
Surv. Paleont. Bull. 57, 1—96.

Wortmann U.G., Herrle J.O. & Weissert H. 2004: Altered carbon

cycling  and  coupled  changes  in  early  Cretaceous  weathering
patterns:  evidence  from  integrated  carbon  isotope  and  sand-
stone  records  of  the  western  Tethys.  Earth  Planet.  Sci.  Lett.
220, 69—82.

Yun Hyesu 1981: Dinoflagellaten aus der Oberkreide (Santon) von

Westfalen. Palaeontographica, Abt. B 177, 1—89.

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Achilleodinium biformoides (Eisenack, 1954) Eaton, 1976
Achomosphaera cf. alcicornu (Eisenack, 1954) Davy & Williams,

1966

Achomosphaera neptuni (Eisenack, 1958) Davey & Williams, 1966;

emend. Duxbury, 1983

Achomosphaera ramosasimilis (Yun Hyesu, 1981) Londeix et al., 1999
Achomosphaera ramulifera (Deflandre, 1937) Evitt, 1963
Achomosphaera ramulifera subsp. ramulifera Deflandre, 1937
Achomosphaera regiensis Corradini, 1973
Achomosphaera spp.
Actinotheca aphroditae Cookson & Eisenack, 1960
Adnatosphaeridium filiferum (Cookson & Eisenack, 1958) Williams &

Downie, 1969

Adnatosphaeridium multispinium Williams & Downie, 1966
Adnatosphaeridium tutulosum (Cookson & Eisenack, 1960) Morgan,

1980

Alisogymnium euclaense (Cookson & Eisenack, 1970) Lentin &

Vozzhennikova, 1990

Alterbidinium acutulum (Wilson, 1967) Lentin & Williams, 1985
Alterbidinium longicornutum Roncaglia et al., 1999
Alterbidinium spp.
Alterbidinium? distinctum (Wilson, 1967) Lentin & Williams, 1985
Andalusiella polymorpha (Malloy, 1972) Lentin & Williams, 1977
Apectodinium augustum (Harland, 1979) Lentin & Williams, 1981
Apectodinium homomorphum (Deflandre & Cookson, 1955) Lentin

& Williams, 1977; emend. Harland, 1979

Apectodinium quinquelatum (Williams & Downie, 1966) Costa &

Downie, 1979

Apectodinium spp.
Apteodinium deflandrei (Clarke & Verdier, 1967) Lucas-Clark, 1987
Apteodinium granulatum Eisenack, 1958; emend. Sarjeant, 1985
Areoligera coronata (Wetzel, 1933) Lejeune-Carpentier, 1937
Areoligera gippingensis Jolley, 1992
Areoligera guembelii Kirsch, 1991
Areoligera medusettiformis Wetzel, 1933 ex Lejeune-Carpentier,

1938

Areoligera senonensis Lejeune-Carpentier, 1938
Areoligera spp.
Areoligera volata Drugg, 1967
Areosphaeridium diktyoplokum (Klumpp, 1953) Eaton, 1971;

emend. Eaton, 1971

Avellodinium? hauteriviense Prössl, 1990
Batiacasphaera grandis Roncaglia et al., 1999
Batiacasphaera micropapillata Stover, 1977
Batiacasphaera spp.
Biconidinium reductum (May, 1980) Kirsch, 1991; emend. Kirsch,

1991

Canningia reticulata Cookson & Eisenack, 1960; emend. Below,

1981

Canningia senonica Clarke & Verdier, 1967
Cannosphaeropsis hughesii Harding, 1990 ex Harding in Williams et

al., 1998

Cannosphaeropsis utinensis Wetzel, 1933; emend. May, 1980
Carpatella cornuta Grigorovitch, 1969; emend. Fechner & Mohr,

1986

Cassidium fragile (Harris, 1965) Drugg, 1967
Cauveridinium membraniphorum (Cookson & Eisenack, 1962)

Masure in Fauconnier & Masure, 2004

Cerodinium diebelii (Alberti, 1959) Lentin & Williams, 1987

Cerodinium obliquipes (Deflandre & Cookson, 1955) Lentin &

Williams, 1987

Cerodinium speciosum (Alberti, 1959) Lentin & Williams, 1987
Cerodinium speciosum subsp. glabrum (Gocht, 1969) Lentin &

Williams, 1987

Cerodinium spp.
Chatangiella ditissima (McIntyre, 1975) Lentin & Williams, 1976
Chatangiella granulifera (Manum, 1963) Lentin & Williams, 1976
Chatangiella hexacalpis Harker & Sarjeant in Harker et al., 1990
Chatangiella madura Lentin & Williams, 1976
Chatangiella spp.
Chatangiella? robusta (Benson, 1976) Stover & Evitt, 1978
Circulodinium brevispinatum (Millioud, 1969) Fauconnier in

Fauconnier & Masure, 2004

Circulodinium distinctum (Deflandre & Cookson, 1955) Jansonius,

1986

Circulodinium spp.
Cladopyxidium velatum Below, 1987
Cleistosphaeridium diversispinosum Davey et al., 1966; emend.

Eaton et al., 2001

Cleistosphaeridium placacanthum (Deflandre & Cookson, 1955)

Eaton et al., 2001; emend. May, 1980

Cleistosphaeridium spp.
Codoniella campanulata (Cookson & Eisenack, 1960) Downie &

Sarjeant, 1965; emend. Davey, 1979

Conosphaeridium striatoconum (Deflandre & Cookson, 1955)

Cookson & Eisenack, 1969

Cordosphaeridium cantharellus (Brosius, 1963) Gocht, 1969
Cordosphaeridium fibrospinosum Davey & Williams, 1966
Cordosphaeridium spp.
Coronifera cf. striolata (Deflandre, 1937) Stover & Evitt, 1978
Coronifera oceanica subsp. hebospina Yun Hyesu, 1981
Coronifera spp.
Corradinisphaeridium horridum (Deflandre, 1937) Masure, 1986;

emend. Masure, 1986

Cribroperdinium? edwardsii (Cookson & Eisenack, 1958) Davey,

1969

Cribroperidinium cooksoniae Norvick, 1976
Cribroperidinium orthoceras (Eisenack, 1958) Davey, 1969;

emend. Sarjeant, 1985

Cribroperidinium spp.
Cyclonephelium filoreticulatum (Slimani, 1994) Prince et al., 1999
Cyclonephelium spp.
Cymososphaeridium validum Davey, 1982
Damassadinium californicum (Drugg, 1967) Fensome et al., 1993
Dapsilidinium laminaspinosum (Davey & Williams, 1966) Lentin

& Williams, 1981

Deflandrea antarctica Wilson, 1967
Deflandrea cygniformis Pöthe de Baldis, 1966
Deflandrea delineata Cookson & Eisenack, 1965
Deflandrea galeata (Lejeune-Carpentier, 1942) Lentin & Williams,

1973; emend. Lejeune-Carpentier & Sarjeant, 1981

Deflandrea phosphoritica Eisenack, 1938
Deflandrea spp.
Dinocyst sp. 1
Dinocyst sp. 2
Dinocyst sp. 3
Dinogymnium acuminatum Evitt et al., 1967
Dinogymnium denticulatum (Alberti, 1961) Evitt et al., 1967

Appendix A. Species list

List of dinocyst taxa encountered in this study, arranged in alphabetical order of genus name. The references to species’ authors are given
by Fensome et al. (2008) and the taxonomy of dinocyst taxa is generally based on Fensome et al. (1993).

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Dinogymnium spp.
Diphyes colligerum (Deflandre & Cookson, 1955) Cookson, 1965;

emend. Cookson, 1965

Diphyes ficusoides Islam, 1983
Disphaerogena carposphaeropsis Wetzel, 1933; emend. Sarjeant,

1985

Disphaerogena irregularis (Wilson, 1988) Lentin & Williams, 1993
Dissiliodinium spp.
Downiesphaeridium? aciculare (Davey, 1969) Islam, 1993
Eisenackia circumtabulata Drugg, 1967
Eisenackia margarita (Harland, 1979) Quattrocchio & Sarjeant, 2003
Eisenackia reticulata (Damassa, 1979) Quattrocchio & Sarjeant,

2003

Elytrocysta druggii Stover & Evitt, 1978
Endoscrinium asymmetricum Riding, 1987
Eocladopyxis peniculata Morgenroth, 1966; emend. McLean, 1976
Exochosphaeridium bifidum (Clarke & Verdier, 1967) Clarke et al.,

1968; emend. Davey, 1969

Exochosphaeridium phragmites Davey et al., 1966
Fibrocysta bipolaris (Cookson & Eisenack, 1965) Stover & Evitt,

1978

Florentinia aculeata Kirsch, 1991
Florentinia deanei (Davey & Williams, 1966) Davey & Verdier, 1973
Florentinia ferox (Deflandre, 1937) Duxbury, 1980
Florentinia hypomagna Yun Hyesu, 1981
Florentinia mayi Kirsch, 1991
Florentinia spp.
Florentinia laciniata Davey & Verdier, 1973
Glaphyrocysta cf. expansa (Corradini, 1973) Roncaglia & Corradini,

1997, p. 187; emend. Roncaglia & Corradini, 1997

Glaphyrocysta exuberans (Deflandre & Cookson, 1955 ex Eaton,

1976) Stover & Evitt, 1978; emend. Sarjeant, 1986

Glaphyrocysta ordinata (Williams & Downie, 1966) Stover &

Evitt, 1978

Glaphyrocysta perforata Hultberg & Malmgren, 1985
Glaphyrocysta semiticta (Bujak in Bujak et al., 1980) Lentin &

Williams, 1981

Glaphyrocysta spp.
Glaphyrocysta wilsonii Kirsch, 1991
Gonyaulacysta dualis (Brideaux & Fisher, 1976) Stover & Evitt, 1978
Gonyaulacysta spp.
Gonyaulacysta? kleithria Duxbury, 1983
Hafniasphaera delicata Fensome et al., 2009
Hapsocysta dictyota Davey, 1979
Hapsocysta peridictya (Eisenack & Cookson, 1960) Davey, 1979;

emend. Davey, 1979

Hapsocysta? benteae N

hr-Hansen, 1993

Heterosphaeridium cordiforme Yun Hyesu, 1981
Heterosphaeridium spinaconjunctum Yun Hyesu, 1981
Homotryblium spp.
Homotryblium tenuispinosum Davey & Williams, 1966
Hystrichodinium pulchrum Deflandre, 1935
Hystrichodinium pulchrum subsp. areatum Marheinecke, 1992
Hystrichokolpoma cinctum Klumpp, 1953
Hystrichokolpoma bulbosum (Ehrenberg, 1838) Morgenroth, 1968;

emend. Morgenroth, 1968

Hystrichokolpoma cf. rigaudiae Deflandre & Cookson, 1955
Hystrichokolpoma reductum Zevenboom & Santarelli in Zevenboom,

1995

Hystrichokolpoma spp.
Hystrichokolpoma truncatum Biffi & Manum, 1988
Hystrichokolpoma unispinum Williams & Downie, 1966
Hystrichosphaeridium bowerbankii Davey & Williams, 1966
Hystrichosphaeridium conispiniferum Yun Hyesu, 1981
Hystrichosphaeridium recurvatum (White, 1842) Lejeune-Carpentier,

1940

Hystrichosphaeridium salpingophorum Deflandre, 1935 ex Deflandre,

1937; emend. Davey & Williams, 1966

Hystrichosphaeridium spp.
Hystrichosphaeridium tenuitubatum Marheinecke, 1992
Hystrichosphaeridium tubiferum (Ehrenberg, 1838) Deflandre, 1937;

emend. Davey & Williams, 1966

Hystrichosphaeridium tubiferum subsp. brevispinum (Davey &

Williams, 1966) Lentin & Williams, 1973; emend. Marheinecke,
1992

Hystrichosphaerina schindewolfii Alberti, 1961
Hystrichosphaeropsis obscura Habib, 1972
Hystrichosphaeropsis ovum Deflandre, 1935
Hystrichostrogylon membraniphorum Agelopoulos, 1964; emend.

Eaton, 1976

Impagidinium maculatum (Cookson & Eisenack, 1961) Stover &

Evitt, 1978

Impagidinium spp.
Impagidinium? ovum (Sah et al., 1970) Stover & Evitt, 1978
Isabelidinium cooksoniae (Alberti, 1959) Lentin & Williams, 1977
Isabelidinium spp.
Kallosphaeridium brevibarbatum de Coninck, 1969; emend. Jan du

Chęne et al., 1985

Kenleyia lophophora Cookson & Eisenack, 1965
Kenleyia spp.
Kleithriasphaeridium loffrense Davey & Verdier, 1976
Leberidocysta chlamydata (Cookson & Eisenack, 1962) Stover &

Evitt, 1978; emend. Fechner, 1985 and Marheinecke, 1992

Leberidocysta spinosa Pestchevitskaya, 2009
Leberidocysta? laticaudata (Vozzhennikova, 1967) Stover & Evitt,

1978

Lejeunecysta communis Biffi & Grignani, 1983
Lejeunecysta hyalina (Gerlach, 1961) Artzner & Dörhöfer, 1978;

emend. Sarjeant, 1984

Lejeunecysta spp.
Magallanesium densispinatum (Stanley, 1965) Quattrocchio &

Sarjeant, 2003

Magallanesium macmurdoense Wilson, 1967
Manumiella druggii (Stover, 1974) Bujak & Davies, 1983
Manumiella seelandica (Lange, 1969) Bujak & Davies 1983; emend.

Firth, 1987

Manumiella seymourensis Askin, 1999
Manumiella spp.
Manumiella? cf. cretacea (Cookson, 1956) Bujak & Davies, 1983
Manumiella? hemmoorensis Marheinecke, 1992
Membranilarnacia polycladiata Cookson & Eisenack in Eisenack,

1963

Membranilarnacia spp.
Membranilarnacia? tenella Morgenroth, 1968
Membranophoridium aspinatum Gerlach, 1961
Muderongia australis Helby, 1987; emend. Monteil, 1991
Muderongia extensiva Duxbury, 1977
Muderongia spp.
Nematosphaeropsis downiei Brown, 1986
Odontochitina operculata (Wetzel, 1933) Deflandre & Cookson,

1955

Odontochitina porifera Cookson, 1956
Odontochitina spp.
Oligosphaeridium albertense (Pocock, 1962) Davey & Williams,

1969

Oligosphaeridium complex (White, 1842) Davey & Williams, 1966
Oligosphaeridium pulcherrimum (Deflandre & Cookson, 1955)

Davey & Williams, 1966

Oligosphaeridium? asterigerum (Gocht, 1959) Davey & Williams,

1969

Operculodinium centrocarpum (Deflandre & Cookson, 1955) Wall,

1967

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Operculodinium severinii (Cookson & Cranwell, 1967) Islam, 1983
Palaeocystodinium australinum (Cookson, 1965) Lentin & Williams,

1976; emend. Malloy, 1972

Palaeocystodinium golzowense Alberti, 1961
Palaeocystodinium lidiae (Górka, 1963) Davey, 1969; emend. Davey,

1969

Palaeocystodinium spp.
Palaeohystrichophora infusorioides Deflandre, 1935
Palaeoperidinium cretaceum (Pocock, 1962) Lentin & Williams,

1976; emend. Harding, 1990 and Evitt et al., 1998

Palaeoperidinium pyrophorum (Ehrenberg, 1838 ex Wetzel, 1933)

Sarjeant, 1967; emend. Sarjeant, 1967

Palaeotetradinium silicorum Deflandre, 1936; emend. Deflandre &

Sarjeant, 1970

Palynodinium biculleus Kirsch, 1991
Palynodinium grallator Gocht, 1970
Paralecaniella indentata (Deflandre & Cookson, 1955) Cookson &

Eisenack, 1970; emend. Elsik, 1977

Pareodinia ceratophora Deflandre, 1947; emend. Gocht, 1970
Pentadinium laticinctum Gerlach, 1961; emend. Benedek et al.,

1982

Pentadinium sabulum Mao Shaozhi & Norris, 1988
Pervosphaeridium granaciculare Fensome et al., 2009
Pervosphaeridium pseudhystrichodinium (Deflandre, 1937) Yun

Hyesu, 1981; emend. Davey, 1969

Pervosphaeridium spp.
Pervosphaeridium truncatum (Davey, 1969) Below, 1982; emend.

Masure, 1988

Phelodinium magnificum (Stanley, 1965) Stover & Evitt, 1978
Phelodinium tricuspe (Wetzel, 1933) Stover & Evitt, 1978; emend.

Lejeune-Carpentier & Sarjeant, 1981

Prolixosphaeridium conulum Davey, 1969
Prolixosphaeridium parvispinum (Deflandre, 1937) Davey et al.,

1969

Protoellipsodinium spinosum Davey & Verdier, 1971
Pseudoceratium anaphrissum (Sarjeant, 1966) Bint, 1986; emend.

Harding, 1990

Pseudoceratium pelliferum Gocht, 1957; emend. Dörhöfer &

Davies, 1980

Pterodinium agadirense Below, 1981
Pterodinium aliferum Eisenack, 1958; emend. Sarjeant, 1985
Pterodinium cingulatum subsp. cingulatum (Wetzel, 1933) Below,

1981

Pterodinium cingulatum subsp. polygonale (Clarke & Verdier,

1967) Paul et al., 1994

Pterodinium cingulatum subsp. reticulatum (Davey & Williams,

1966) Lentin & Williams, 1981

Pterodinium? cornutum Cookson & Eisenack, 1962
Pterodinium? pterotum (Cookson & Eisenack, 1958) Pavlishina,

1990; emend. Pavlishina, 1990

Pyxidinopsis spp.
Pyxidinopsis waipawaensis Wilson, 1988
Raetiadinium evittigratia Kirsch, 1991
Raetiaedinium truncigerum (Deflandre, 1937) Kirsch, 1991
Rhiptocorys veligera (Deflandre, 1937) Lejeune-Carpentier &

Sarjeant, 1983; emend. Lejeune-Carpentier & Sarjeant, 1983

Rhombodella paucispina (Alberti, 1961) Duxbury, 1980
Riculacysta amplexa Kirsch, 1991
Riculacysta spp.
Riculacysta? pala Kirsch, 1991
Rigaudella aemula (Deflandre, 1939) Below, 1982; emend. Below,

1982

Rigaudella apenninica (Corradini, 1973) Below, 1982
Rottnestia borussica (Eisenack, 1954) Cookson & Eisenack, 1961
Rottnestia wetzelii (Deflandre, 1937) Slimani, 1994
Scriniodinium crystallinum (Deflandre, 1939) Klement, 1960;

emend. Riding & Fensome, 2003

Scriniodinium spp.
Senoniasphaera inornata (Drugg, 1970) Stover & Evitt, 1978
Senoniasphaera protrusa Clarke & Verdier, 1967; emend. Prince et

al., 1999

Spinidinium echinoideum (Cookson & Eisenack, 1960) Lentin &

Williams, 1976; emend. Sverdlove & Habib, 1974

Spinidinium echinoideum subsp. rhombicum (Cookson & Eisenack,

1974) Lentin & Williams, 1976

Spinidinium spp.
Spiniferella cornuta (Gerlach, 1961) Stover & Hardenbol, 1994;

emend. Stover & Hardenbol, 1994

Spiniferites cf. bulloideus (Deflandre & Cookson, 1955) Sarjeant,

1970

Spiniferites dentatus (Gocht, 1959) Lentin & Williams, 1973;

emend. Duxbury, 1977

Spiniferites membranaceus (Rossignol, 1964) Sarjeant, 1970
Spiniferites multibrevis (Davey & Williams, 1966) Below, 1982
Spiniferites pseudofurcatus (Klumpp, 1953) Sarjeant, 1970; emend.

Sarjeant, 1981

Spiniferites ramosus (Ehrenberg, 1838) Mantell, 1854
Spiniferites ramosus subsp. brevifurcatus (Cookson & Eisenack,

1974) Lentin & Williams, 1977

Spiniferites ramosus subsp. cavispinosus Hansen, 1977
Spiniferites ramosus subsp. gracile (Davey & Williams, 1966) Lentin

& Williams, 1973

Spiniferites ramosus subsp. granosus (Davey & Williams, 1966)

Lentin & Williams, 1973

Spiniferites scabrosus (Clarke & Verdier, 1967) Lentin & Williams,

1975

Spiniferites spp.
Spiniferites? spongiosus Duxbury, 2001
Spongodinium delitiense (Ehrenberg, 1838) Deflandre, 1936; emend.

Lucas-Clark, 1987

Stanfordella fastigiata (Duxbury, 1977) Helenes & Lucas-Clark,

1997; emend. Helenes & Lucas-Clark, 1997

Subtilisphaera perlucida (Alberti, 1959) Jain & Millepied, 1973
Subtilisphaera terrula (Davey, 1974) Lentin & Williams, 1976;

emend. Harding, 1986

Surculosphaeridium belowii Yun Hyesu, 1981
Surculosphaeridium longifurcatum (Firtion, 1952) Davey et al.,

1966

Surculosphaeridium trunculum Davey, 1979
Surculosphaeridium? basifurcatum Yun Hyesu, 1981
Systematophora cretacea Davey, 1979
Systematophora? septata Wilson, 1988
Tanyosphaeridium boletus Davey, 1974
Tanyosphaeridium xanthiopyxides (Wetzel, 1933 ex Deflandre,

1937) Stover & Evitt, 1978; emend. Sarjeant, 1985

Tenua hystrix Eisenack, 1958; emend. Sarjeant, 1985
Thalassiphora delicata Williams & Downie, 1966; emend. Eaton,

1976

Thalassiphora inflata Heilmann-Clausen in Thomsen & Heilmann-

Clausen, 1985

Thalassiphora pelagica (Eisenack, 1954) Eisenack & Gocht, 1960;

emend. Benedek & Gocht, 1981

Thalassiphora reticulata Morgenroth, 1966
Trabeculidium pusulosum (Morgenroth, 1966) Duxbury, 1980
Trabeculidium quinquetrum Duxbury, 1980
Trichodinium ciliatum (Gocht, 1959) Eisenack & Klement, 1964
Trigonopyxidia ginella (Cookson & Eisenack, 1960) Downie &

Sarjeant, 1965

Trithyrodinium cf. evittii Drugg, 1967
Trithyrodinium evittii Drugg, 1967
Trithyrodinium robustum Benson, 1976
Trithyrodinium suspectum (Manum & Cookson, 1964) Davey, 1969

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Turbiosphaera filosa (Wilson, 1967) Archangelsky, 1969
Turnhosphaera hypoflata (Yun Hyesu, 1981) Slimani, 1994;

emend. Slimani, 1994

Wetzeliella symmetrica Weiler, 1956
Xenascus cf. asperatus Stover & Helby, 1987
Xenascus gochtii (Corradini, 1973) Stover & Evitt, 1978

Xenascus sarjeantii (Corradini, 1973) Stover & Evitt, 1978
Xenascus spp.
Xiphophoridium alatum (Cookson & Eisenack, 1962) Sarjeant,

1966; emend. Sarjeant, 1966

Ynezidinium brevisulcatum (Michoux, 1985) Lucas-Clark & Helenes,

2000

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1420

120

1470

35 17

1 1

1520

1 12

1530

1580

1630

1680

1730

1780

1830

1880

1930

1980

2030

3 1

2080

4 1

2130

1 1

2180

2230

2280

2330

2380

2430

2470

2480

2530

1 1

2560

(

)

2565

Appendix C (continued)

1220

1270

1320

1370

1420

1470

2 38

1520

7 1

1530

9 1

1580

1630

1 1

1680

3 1

1730

1780

1830

1880

1930

1980

2030

2080

1 1

2130

2180

2230

2280

2330

2380

2430

2470

2480

1 1

2530

1 15

1 1

2560

(

)

2565

Appendix C (continued)

1220

1270

1320

1370

1420

1470

1 11

2 1

1520

1530

38 1

1580

1 47

1630

1680

1 26

1730

1 36

1780

1830

1880

1930

3 72

1980

2030

2080

2130

2180

2230

2280

2330

1 30

2380

2430

2470

2480

2530

2560

(

)

2565

Appendix C (continued)

1220

5 1 141

1270

3 3 160

1320

12 2 117

1370

124

1420

2 1 306

1470

308

1520

1 32

296

1530

241

1580

4 2 198

1630

150

1680

8 11

4 10

163

1730

6 12

218

1780

1830

1880

1930

246

1980

2030

2080

2130

263

2180

2230

2280

8 14

2330

7 12

114

2380

2430

2470

2480

17 16 27

125

2530

35 13 16

204

2560

(

)

2565

Appendix C (continued)

xii